Geology topics
Frank T. van Manen
Publications and source records attributed to Frank T. van Manen.
Primarily resident grizzly bears respond to late-season elk harvest
Autumn ungulate hunting in the Greater Yellowstone Ecosystem carries the risk of hunter–grizzly bear ( Ursus arctos ) conflict and creates a substantial challenge for managers. For Grand Teton National Park, Wyoming, USA, a key information need is whether increased availability of elk ( Cervus canadensis ) carcasses during a late autumn (Nov–Dec) harvest within the national park attracts grizzly bears and increases the potential for conflict with hunters. Using a robust design analysis with 6 primary sampling periods during 2014–2015, we tested the hypothesis that the elk harvest resulted in temporary movements of grizzly bears into the hunt areas, thus increasing bear numbers. We detected 31 unique individuals (6 F, 25 M) through genetic sampling and retained 26 encounter histories for analysis. Markovian movement models had more support than a null model of no temporary movement. Contrary to our research hypothesis, temporary movements into the study area occurred between the July–August (no hunt; N̄ 2014–2015 = 5) and September–October (no hunt; N̄ 2014–2015 = 24) primary periods each year, rather than during the transition from September–October (no hunt) to November–December (hunt; N̄ 2014–2015 = 15). A post hoc analysis indicated that September–October population estimates were biased high by detections of transient bears. Grizzly bear presence during the elk hunt was limited to approximately 15 resident bears that specialized in accessing elk carcasses. The late timing of the elk hunt likely moderated the effect of carcasses as a food attractant because it coincides with the onset of hibernation. From a population response perspective, the current timing of the elk harvest likely represents a scenario of low relative risk of hunter–bear conflicts. The risk of hunter–grizzly bear encounters remains, but may be more a function of factors that operate at the level of individual bears and hunters, such as hunter movements and bear responses to olfactory cues.
Grizzly bear depredation on grazing allotments in the Yellowstone ecosystem
Grizzly bear (Ursus arctos) conflicts with humans, including livestock depredation on public land grazing allotments, have increased during the last several decades within the Greater Yellowstone Ecosystem (GYE) in the western United States as the grizzly bear population has grown in number and occupied range. Minimizing conflicts and improving conservation efficacy requires information on the relationships between livestock depredations, allotment management, grizzly bear habitat conditions, and their interactions. We used generalized linear mixed models to evaluate spatio-temporal relationships between grizzly bear depredation of livestock and the characteristics of 316 United States Department of Agriculture Forest Service and National Park Service grazing allotments in the GYE during 1992–2014. We evaluated relationships at 2 spatial extents, representing daily and annual grizzly bear activity areas. During the study period, more grazing allotments became occupied by grizzly bears and most livestock depredations were associated with these areas of population expansion. Number of livestock (beta = 1.15 +/- 0.19 [SE]) and grizzly bear density index (beta = 1.13 +/- 0.10) had the greatest effects on the number of livestock depredation events relative to other allotment attributes. Estimated number of depredation events increased by approximately 20% when cow-calf pairs increased by 100 pairs and grizzly bear density index increased by 1 bear/196 km2 (the average annual home-range size of a female grizzly bear in the GYE). Additionally, grazing allotment size was positively related to the number of depredation events (beta = 0.56 +/- 0.16), whereas the presence of bull cattle or horses was associated with an approximately 50% reduction in depredations (beta = -0.71 +/- 0.37). Livestock depredation events were greater for allotments with lower road density (beta = -0.89 +/- 0.28), less rugged terrain (beta = -0.57 +/- 0.25), higher vegetative primary productivity (beta = 0.33 +/- 0.16), and more whitebark pine coverage (beta = 0.30 +/- 0.15). Relationships between depredations and grizzly bear habitat conditions varied across spatial extents. As the grizzly bear population continues to expand, natural resource managers and livestock producers could focus efforts on allotments with a higher density of grizzly bears, fewer roads, and quality grizzly bear habitat, including higher vegetative productivity, when developing cooperative management plans and preventative measures to reduce the likelihood of depredation. The perspectives gained from our analysis provide context for long-term, landscape-level planning to accommodate livestock production on public lands while meeting conservation goals for grizzly bears.
Estimating distemper virus dynamics among wolves and grizzly bears using serology and Bayesian state‐space models
Many parasites infect multiple hosts, but estimating the transmission across host species remains a key challenge in disease ecology. We investigated the within and across host species dynamics of canine distemper virus (CDV) in grizzly bears ( Ursus arctos ) and wolves ( Canis lupus ) of the Greater Yellowstone Ecosystem (GYE). We hypothesized that grizzly bears may be more likely to be exposed to CDV during outbreaks in the wolf population because grizzly bears often displace wolves while scavenging carcasses. We used serological data collected from 1984 to 2014 in conjunction with Bayesian state‐space models to infer the temporal dynamics of CDV. These models accounted for the unknown timing of pathogen exposure, and we assessed how different testing thresholds and the potential for testing errors affected our conclusions. We identified three main CDV outbreaks (1999, 2005, and 2008) in wolves, which were more obvious when we used higher diagnostic thresholds to qualify as seropositive. There was some evidence for increased exposure rates in grizzly bears in 2005, but the magnitude of the wolf effect on bear exposures was poorly estimated and depended upon our prior distributions. Grizzly bears were exposed to CDV prior to wolf reintroduction and during time periods outside of known wolf outbreaks, thus wolves are only one of several potential routes for grizzly bear exposures. Our modeling approach accounts for several of the shortcomings of serological data and is applicable to many wildlife disease systems, but is most informative when testing intervals are short. CDV circulates in a wide range of carnivore species, but it remains unclear whether the disease persists locally within the GYE carnivore community or is periodically reintroduced from distant regions with larger host populations.
Carnivore hotspots in Peninsular Malaysia and their landscape attributes
Mammalian carnivores play a vital role in ecosystem functioning. However, they are prone to extinction because of low population densities and growth rates, and high levels of persecution or exploitation. In tropical biodiversity hotspots such as Peninsular Malaysia, rapid conversion of natural habitats threatens the persistence of this vulnerable group of animals. Here, we carried out the first comprehensive literature review on 31 carnivore species reported to occur in Peninsular Malaysia and updated their probable distribution. We georeferenced 375 observations of 28 species of carnivore from 89 unique geographic locations using records spanning 1948 to 2014. Using the Getis-Ord Gi*statistic and weighted survey records by IUCN Red List status, we identified hotspots of species that were of conservation concern and built regression models to identify environmental and anthropogenic landscape factors associated with Getis-Ord Gi* z scores. Our analyses identified two carnivore hotspots that were spatially concordant with two of the peninsula’s largest and most contiguous forest complexes, associated with Taman Negara National Park and Royal Belum State Park. A cold spot overlapped with the southwestern region of the Peninsula, reflecting the disappearance of carnivores with higher conservation rankings from increasingly fragmented natural habitats. Getis-Ord Gi* z scores were negatively associated with elevation, and positively associated with the proportion of natural land cover and distance from the capital city. Malaysia contains some of the world’s most diverse carnivore assemblages, but recent rates of forest loss are some of the highest in the world. Reducing poaching and maintaining large, contiguous tracts of lowland forests will be crucial, not only for the persistence of threatened carnivores, but for many mammalian species in general.
Size‐assortative choice and mate availability influences hybridization between red wolves (Canis rufus) and coyotes (Canis latrans)
Anthropogenic hybridization of historically isolated taxa has become a primary conservation challenge for many imperiled species. Indeed, hybridization between red wolves ( Canis rufus ) and coyotes ( Canis latrans ) poses a significant challenge to red wolf recovery. We considered seven hypotheses to assess factors influencing hybridization between red wolves and coyotes via pair‐bonding between the two species. Because long‐term monogamy and defense of all‐purpose territories are core characteristics of both species, mate choice has long‐term consequences. Therefore, red wolves may choose similar‐sized mates to acquire partners that behave similarly to themselves in the use of space and diet. We observed multiple factors influencing breeding pair formation by red wolves and found that most wolves paired with similar‐sized conspecifics and wolves that formed congeneric pairs with nonwolves (coyotes and hybrids) were mostly female wolves, the smaller of the two sexes. Additionally, we observed that lower red wolf abundance relative to nonwolves and the absence of helpers increased the probability that wolves consorted with nonwolves. However, successful pairings between red wolves and nonwolves were associated with wolves that maintained small home ranges. Behaviors associated with territoriality are energetically demanding and behaviors (e.g., aggressive interactions, foraging, and space use) involved in maintaining territories are influenced by body size. Consequently, we propose the hypothesis that size disparities between consorting red wolves and coyotes influence positive assortative mating and may represent a reproductive barrier between the two species. We offer that it may be possible to maintain wild populations of red wolves in the presence of coyotes if management strategies increase red wolf abundance on the landscape by mitigating key threats, such as human‐caused mortality and hybridization with coyotes. Increasing red wolf abundance would likely restore selection pressures that increase mean body and home‐range sizes of red wolves and decrease hybridization rates via reduced occurrence of congeneric pairs.
Potential paths for male-mediated gene flow to and from an isolated grizzly bear population
For several decades, grizzly bear populations in the Greater Yellowstone Ecosystem (GYE) and the Northern Continental Divide Ecosystem (NCDE) have increased in numbers and range extent. The GYE population remains isolated and although effective population size has increased since the early 1980s, genetic connectivity between these populations remains a long-term management goal. With only ~110 km distance separating current estimates of occupied range for these populations, the potential for gene flow is likely greater now than it has been for many decades. We sought to delineate potential paths that would provide the opportunity for male-mediated gene flow between the two populations. We first developed step-selection functions to generate conductance layers using ecological, physical, and anthropogenic landscape features associated with non-stationary GPS locations of 124 male grizzly bears (199 bear-years). We then used a randomized shortest path (RSP) algorithm to estimate the average number of net passages for all grid cells in the study region, when moving from an origin to a destination node. Given habitat characteristics that were the basis for the conductance layer, movements follow certain grid cell sequences more than others and the resulting RSP values thus provide a measure of movement potential. Repeating this process for 100 pairs of random origin and destination nodes, we identified paths for three levels of random deviation (θ) from the least-cost path. We observed broad-scale concordance between model predictions for paths originating in the NCDE and those originating in the GYE for all three levels of movement exploration. Model predictions indicated that male grizzly bear movement between the ecosystems could involve a variety of routes, and verified observations of grizzly bears outside occupied range supported this finding. Where landscape features concentrated paths into corridors (e.g., because of anthropogenic influence), they typically followed neighboring mountain ranges, of which several could serve as pivotal stepping stones. The RSP layers provide detailed, spatially explicit information for land managers and organizations working with land owners to identify and prioritize conservation measures that maintain or enhance the integrity of potential areas conducive to male grizzly bear dispersal.
Using diets of Canis breeding pairs to assess resource partitioning between sympatric red wolves and coyotes
Foraging behaviors of red wolves ( Canis rufus ) and coyotes ( Canis latrans ) are complex and their ability to form congeneric breeding pairs and hybridize further complicates our understanding of factors influencing their diets. Through scat analysis, we assessed prey selection of red wolf, coyote, and congeneric breeding pairs formed by red wolves and coyotes, and found that all 3 had similar diets. However, red wolf and congeneric pairs consumed more white-tailed deer ( Odocoileus virginianus ) than coyote pairs. Coyotes forming breeding pairs with red wolves had 12% more white-tailed deer in their diet than conspecifics paired with coyotes. Contrary to many studies on coyotes in the southeastern United States, we found coyotes in eastern North Carolina to be primarily carnivorous with increased consumption of deer during winter. Although prey selection was generally similar among the 3 groups, differences in diet among different breeding pairs were strongly associated with body mass. Larger breeding pairs consumed more white-tailed deer, and fewer rabbits ( Sylvilagus spp.) and other small mammals. Partitioning of food resources by sympatric red wolves and coyotes is likely via differences in the proportions of similar prey consumed, rather than differences in types of prey exploited. Consequently, our results suggest coexistence of red wolves and coyotes in the southeastern United States may not be possible because there are limited opportunities for niche partitioning to reduce competitive interactions.
Space use and habitat selection by resident and transient red wolves ( Canis rufus )
Recovery of large carnivores remains a challenge because complex spatial dynamics that facilitate population persistence are poorly understood. In particular, recovery of the critically endangered red wolf ( Canis rufus ) has been challenging because of its vulnerability to extinction via human-caused mortality and hybridization with coyotes ( Canis latrans ). Therefore, understanding red wolf space use and habitat selection is important to assist recovery because key aspects of wolf ecology such as interspecific competition, foraging, and habitat selection are well-known to influence population dynamics and persistence. During 2009–2011, we used global positioning system (GPS) radio-telemetry to quantify space use and 3 rd -order habitat selection for resident and transient red wolves on the Albemarle Peninsula of eastern North Carolina. The Albemarle Peninsula was a predominantly agricultural landscape in which red wolves maintained spatially stable home ranges that varied between 25 km 2 and 190 km 2 . Conversely, transient red wolves did not maintain home ranges and traversed areas between 122 km 2 and 681 km 2 . Space use by transient red wolves was not spatially stable and exhibited shifting patterns until residency was achieved by individual wolves. Habitat selection was similar between resident and transient red wolves in which agricultural habitats were selected over forested habitats. However, transients showed stronger selection for edges and roads than resident red wolves. Behaviors of transient wolves are rarely reported in studies of space use and habitat selection because of technological limitations to observed extensive space use and because they do not contribute reproductively to populations. Transients in our study comprised displaced red wolves and younger dispersers that competed for limited space and mating opportunities. Therefore, our results suggest that transiency is likely an important life-history strategy for red wolves that facilitates metapopulation dynamics through short- and long-distance movements and eventual replacement of breeding residents lost to mortality.
Diet and macronutrient optimization in wild ursids: A comparison of grizzly bears with sympatric and allopatric black bears
When fed ad libitum, ursids can maximize mass gain by selecting mixed diets wherein protein provides 17 ± 4% of digestible energy, relative to carbohydrates or lipids. In the wild, this ability is likely constrained by seasonal food availability, limits of intake rate as body size increases, and competition. By visiting locations of 37 individuals during 274 bear-days, we documented foods consumed by grizzly ( Ursus arctos ) and black bears ( Ursus americanus ) in Grand Teton National Park during 2004–2006. Based on published nutritional data, we estimated foods and macronutrients as percentages of daily energy intake. Using principal components and cluster analyses, we identified 14 daily diet types. Only 4 diets, accounting for 21% of days, provided protein levels within the optimal range. Nine diets (75% of days) led to over-consumption of protein, and 1 diet (3% of days) led to under-consumption. Highest protein levels were associated with animal matter (i.e., insects, vertebrates), which accounted for 46–47% of daily energy for both species. As predicted: 1) daily diets dominated by high-energy vertebrates were positively associated with grizzly bears and mean percent protein intake was positively associated with body mass; 2) diets dominated by low-protein fruits were positively associated with smaller-bodied black bears; and 3) mean protein was highest during spring, when high-energy plant foods were scarce, however it was also higher than optimal during summer and fall. Contrary to our prediction: 4) allopatric black bears did not exhibit food selection for high-energy foods similar to grizzly bears. Although optimal gain of body mass was typically constrained, bears usually opted for the energetically superior trade-off of consuming high-energy, high-protein foods. Given protein digestion efficiency similar to obligate carnivores, this choice likely supported mass gain, consistent with studies showing monthly increases in percent body fat among bears in this region.
Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data
Global positioning system (GPS) wildlife collars have revolutionized wildlife research. Studies of predation by free-ranging carnivores have particularly benefited from the application of location clustering algorithms to determine when and where predation events occur. These studies have changed our understanding of large carnivore behavior, but the gains have concentrated on obligate carnivores. Facultative carnivores, such as grizzly/brown bears ( Ursus arctos ), exhibit a variety of behaviors that can lead to the formation of GPS clusters. We combined clustering techniques with field site investigations of grizzly bear GPS locations ( n = 732 site investigations; 2004–2011) to produce 174 GPS clusters where documented behavior was partitioned into five classes (large-biomass carcass, small-biomass carcass, old carcass, non-carcass activity, and resting). We used multinomial logistic regression to predict the probability of clusters belonging to each class. Two cross-validation methods—leaving out individual clusters, or leaving out individual bears—showed that correct prediction of bear visitation to large-biomass carcasses was 78–88%, whereas the false-positive rate was 18–24%. As a case study, we applied our predictive model to a GPS data set of 266 bear-years in the Greater Yellowstone Ecosystem (2002–2011) and examined trends in carcass visitation during fall hyperphagia (September–October). We identified 1997 spatial GPS clusters, of which 347 were predicted to be large-biomass carcasses. We used the clustered data to develop a carcass visitation index, which varied annually, but more than doubled during the study period. Our study demonstrates the effectiveness and utility of identifying GPS clusters associated with carcass visitation by a facultative carnivore.
New challenges for grizzly bear management in Yellowstone National Park
A key factor contributing to the success of grizzly bear Ursus arctos conservation in the Greater Yellowstone Ecosystem has been the existence of a large protected area, Yellowstone National Park. We provide an overview of recovery efforts, how demographic parameters changed as the population increased, and how the bear management program in Yellowstone National Park has evolved to address new management challenges over time. Finally, using the management experiences in Yellowstone National Park, we present comparisons and perspectives regarding brown bear management in Shiretoko National Park.
Multiple estimates of effective population size for monitoring a long-lived vertebrate: An application to Yellowstone grizzly bears
Effective population size ( N e ) is a key parameter for monitoring the genetic health of threatened populations because it reflects a population's evolutionary potential and risk of extinction due to genetic stochasticity. However, its application to wildlife monitoring has been limited because it is difficult to measure in natural populations. The isolated and well-studied population of grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem provides a rare opportunity to examine the usefulness of different N e estimators for monitoring. We genotyped 729 Yellowstone grizzly bears using 20 microsatellites and applied three single-sample estimators to examine contemporary trends in generation interval (GI), effective number of breeders ( N b ) and N e during 1982–2007. We also used multisample methods to estimate variance ( N eV ) and inbreeding N e ( N eI ). Single-sample estimates revealed positive trajectories, with over a fourfold increase in N e (≈100 to 450) and near doubling of the GI (≈8 to 14) from the 1980s to 2000s. N eV (240–319) and N eI (256) were comparable with the harmonic mean single-sample N e (213) over the time period. Reanalysing historical data, we found N eV increased from ≈80 in the 1910s–1960s to ≈280 in the contemporary population. The estimated ratio of effective to total census size ( N e /N c ) was stable and high (0.42–0.66) compared to previous brown bear studies. These results support independent demographic evidence for Yellowstone grizzly bear population growth since the 1980s. They further demonstrate how genetic monitoring of N e can complement demographic-based monitoring of N c and vital rates, providing a valuable tool for wildlife managers.
Density dependence, whitebark pine, and vital rates of grizzly bears
Understanding factors influencing changes in population trajectory is important for effective wildlife management, particularly for populations of conservation concern. Annual population growth of the grizzly bear ( Ursus arctos ) population in the Greater Yellowstone Ecosystem, USA has slowed from 4.2–7.6% during 1983–2001 to 0.3–2.2% during 2002–2011. Substantial changes in availability of a key food source and bear population density have occurred. Whitebark pine ( Pinus albicaulis ), the seeds of which are a valuable but variable fall food for grizzly bears, has experienced substantial mortality primarily due to a mountain pine beetle ( Dendroctonus ponderosae ) outbreak that started in the early 2000s. Positive growth rates of grizzly bears have resulted in populations reaching high densities in some areas and have contributed to continued range expansion. We tested research hypotheses to examine if changes in vital rates detected during the past decade were more associated with whitebark pine decline or, alternatively, increasing grizzly bear density. We focused our assessment on known-fate data to estimate survival of cubs-of-the-year (cubs), yearlings, and independent bears (≥2 yrs), and reproductive transition of females from having no offspring to having cubs. We used spatially and temporally explicit indices for grizzly bear density and whitebark pine mortality as individual covariates. Models indicated moderate support for an increase in survival of independent male bears over 1983–2012, whereas independent female survival did not change. Cub survival, yearling survival, and reproductive transition from no offspring to cubs all changed during the 30-year study period, with lower rates evident during the last 10–15 years. Cub survival and reproductive transition were negatively associated with an index of grizzly bear density, indicating greater declines where bear densities were higher. Our analyses did not support a similar relationship for the index of whitebark pine mortality. The results of our study support the interpretation that slowing of population growth during the last decade was associated more with increasing grizzly bear density than the decline in whitebark pine. Grizzly bear density and its potential effect on vital rates and population trajectory warrant consideration for management of the grizzly bear population in the Greater Yellowstone Ecosystem.
Space use and habitat selection by resident and transient coyotes ( Canis latrans )
Little information exists on coyote ( Canis latrans ) space use and habitat selection in the southeastern United States and most studies conducted in the Southeast have been carried out within small study areas (e.g., ≤1,000 km 2 ). Therefore, studying the placement, size, and habitat composition of coyote home ranges over broad geographic areas could provide relevant insights regarding how coyote populations adjust to regionally varying ecological conditions. Despite an increasing number of studies of coyote ecology, few studies have assessed the role of transiency as a life-history strategy among coyotes. During 2009–2011, we used GPS radio-telemetry to study coyote space use and habitat selection on the Albemarle Peninsula of northeastern North Carolina. We quantified space use and 2 nd - and 3 rd -order habitat selection for resident and transient coyotes to describe space use patterns in a predominantly agricultural landscape. The upper limit of coyote home-range size was approximately 47 km 2 and coyotes exhibiting shifting patterns of space use of areas >65 km 2 were transients. Transients exhibited localized space use patterns for short durations prior to establishing home ranges, which we defined as “biding” areas. Resident and transient coyotes demonstrated similar habitat selection, notably selection of agricultural over forested habitats. However, transients exhibited stronger selection for roads than resident coyotes. Although transient coyotes are less likely to contribute reproductively to their population, transiency may be an important life history trait that facilitates metapopulation dynamics through dispersal and the eventual replacement of breeding residents lost to mortality.
Book Reivew: A chance for lasting survival: Ecology and behavior of wild giant pandas
“If we watch species going extinct in front of us, how useful is that we publish 100 or even 1,000 papers by studying them?” (p. 330). This quote from senior author Pan Wenshi captures an important essence of this book. A translation of a 2001 monograph originally published in Chinese, this volume details the findings of a 15-year research program in the Qinling Mountains by Wenshi and his students. Starting in 1984, this Chinese research team from Peking University was only the second to study free-ranging pandas. This is the remarkable journey of a devoted group of field researchers who helped changed the course of giant panda conservation, events that few conservationists outside of China have been aware of until now. Review info: A chance for lasting survival: Ecology and behavior of wild giant pandas. By Pan Wenshi, Lü Zhi, Zhu Xiaojian, Wang Dajun, Wang Hao, Long Yu, Fu Dali, and Zhou Xin; edited by, William J. McShea, Richard B. Harris, David L. Garshelis, and Wang Dajun, 2014. ISBN: 978-1-935623-17-5, 349pp.
Forty years of grizzly bear recovery in the Greater Yellowstone Ecosystem
No abstract available.
Influence of whitebark pine decline on fall habitat use and movements of grizzly bears in the Greater Yellowstone Ecosystem
When abundant, seeds of the high-elevation whitebark pine (WBP; Pinus albicaulis ) are an important fall food for grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem. Rates of bear mortality and bear/human conflicts have been inversely associated with WBP productivity. Recently, mountain pine beetles ( Dendroctonus ponderosae ) have killed many cone-producing WBP trees. We used fall (15 August–30 September) Global Positioning System locations from 89 bear years to investigate temporal changes in habitat use and movements during 2000–2011. We calculated Manly–Chesson (MC) indices for selectivity of WBP habitat and secure habitat (≥500 m from roads and human developments), determined dates of WBP use, and documented net daily movement distances and activity radii. To evaluate temporal trends, we used regression, model selection, and candidate model sets consisting of annual WBP production, sex, and year. One-third of sampled grizzly bears had fall ranges with little or no mapped WBP habitat. Most other bears (72%) had a MC index above 0.5, indicating selection for WBP habitats. From 2000 to 2011, mean MC index decreased and median date of WBP use shifted about 1 week later. We detected no trends in movement indices over time. Outside of national parks, there was no correlation between the MC indices for WBP habitat and secure habitat, and most bears (78%) selected for secure habitat. Nonetheless, mean MC index for secure habitat decreased over the study period during years of good WBP productivity. The wide diet breadth and foraging plasticity of grizzly bears likely allowed them to adjust to declining WBP. Bears reduced use of WBP stands without increasing movement rates, suggesting they obtained alternative fall foods within their local surroundings. However, the reduction in mortality risk historically associated with use of secure, high-elevation WBP habitat may be diminishing for bears residing in multiple-use areas.