Unreliable knowledge about economic impacts of large carnivores on bovine calves

Sommers et al. (2010) reported that recolonizing predators increased bovine calf mortality rates in the Upper Green River Cattle Allotment in western Wyoming. However, Sommers et al. (2010) failed to consider multiple competing hypotheses explaining calf loss rates, increasing the likelihood that their results are actually spurious. I reanalyzed their data using a multiple competing hypotheses framework that considered effects of livestock density, summer precipitation, bias in reporting rates, and whether mortality by different predator species was compensatory. I found support for a confounded web of factors influencing calf losses. Calf losses increased with livestock density (which increased during the study), but also during drier summers and with increasing rancher reporting rates. Although both wolves (Canis lupus) and grizzly bears (Ursus arctos) did increase calf losses, the presence of just grizzly bears alone did not significantly increase calf losses. Unconditional estimates of the effects of wolves and grizzly bears on calf losses were only 2.0% (95% CI 0.53–3.81), compared to 3.6% reported by Sommers et al. (2010). Most importantly, however, I report bias in favor of livestock producers in the authors' assumptions that cast further doubt on the rigor of their results. In conclusion, I recommend managers not consider the spurious predator compensation factors reported by Sommers et al. (2010) to be reliable. © 2011 The Wildlife Society.     

Effects of predator treatments,individual traits,and environment on moose survival in Alaska

We studied moose (Alces alces) survival, physical condition, and abundance in a 3-predator system in western Interior Alaska, USA, during 2001–2007. Our objective was to quantify the effects of predator treatments on moose population dynamics by investigating changes in survival while evaluating the contribution of potentially confounding covariates. In May 2003 and 2004, we reduced black bear (Ursus americanus) and brown bear (U. arctos) numbers by translocating bears ≥240 km from the study area. Aircraft-assisted take reduced wolf (Canis lupus) numbers markedly in the study area during 2004–2007. We estimated black bears were reduced by approximately 96% by June 2004 and recovered to within 27% of untreated numbers by May 2007. Brown bears were reduced approximately 50% by June 2004. Late-winter wolf numbers were reduced by 75% by 2005 and likely remained at these levels through 2007. In addition to predator treatments, moose hunting closures during 2004–2007 reduced harvests of male moose by 60% in the study area. Predator treatments resulted in increased calf survival rates during summer (primarily from reduced black bear predation) and autumn (primarily from reduced wolf predation). Predator treatments had little influence on survival of moose calves during winter; instead, calf survival was influenced by snow depth and possibly temperature. Increased survival of moose calves during summer and autumn combined with relatively constant winter survival in most years led to a corresponding increase in annual survival of calves following predator treatments. Nonpredation mortalities of calves increased following predator treatments; however, this increase provided little compensation to the decrease in predation mortalities resulting from treatments. Thus, predator-induced calf mortality was primarily additive. Summer survival of moose calves was positively related to calf mass (β > 0.07, SE = 0.073) during treated years and lower (β = −0.82, SE = 0.247) for twins than singletons during all years. Following predator treatments, survival of yearling moose increased 8.7% for females and 21.4% for males during summer and 2.2% for females and 15.6% for males during autumn. Annual survival of adult (≥2 yr old) female moose also increased in treated years and was negatively (β = −0.21, SE = 0.078) related to age. Moose density increased 45%, from 0.38 moose/km² in 2001 to 0.55 moose/km² in 2007, which resulted from annual increases in overall survival of moose, not increases in reproductive rates. Indices of nutritional status remained constant throughout our study despite increased moose density. This information can be used by wildlife managers and policymakers to better understand the outcomes of predator treatments in Alaska and similar environments. © 2011 The Wildlife Society.     

Bear-Baiting May Exacerbate Wolf-Hunting Dog Conflict

Background

The influence of policy on the incidence of human-wildlife conflict can be complex and not entirely anticipated. Policies for managing bear hunter success and depredation on hunting dogs by wolves represent an important case because with increasing wolves, depredations are expected to increase. This case is challenging because compensation for wolf depredation on hunting dogs as compared to livestock is less common and more likely to be opposed. Therefore, actions that minimize the likelihood of such conflicts are a conservation need.

Methodology/Principal Findings

We used data from two US states with similar wolf populations but markedly different wolf/hunting dog depredation patterns to examine the influence of bear hunting regulations, bear hunter to wolf ratios, hunter method, and hunter effort on wolf depredation trends. Results indicated that the ratio of bear hunting permits sold per wolf, and hunter method are important factors affecting wolf depredation trends in the Upper Great Lakes region, but strong differences exist between Michigan and Wisconsin related in part to the timing and duration of bear-baiting (i.e., free feeding). The probability that a wolf depredated a bear-hunting dog increases with the duration of bear-baiting, resulting in a relative risk of depredation 2.12–7.22× greater in Wisconsin than Michigan. The net effect of compensation for hunting dog depredation in Wisconsin may also contribute to the difference between states.

Conclusions/Significance

These results identified a potential tradeoff between bear hunting success and wolf/bear-hunting dog conflict. These results indicate that management options to minimize conflict exist, such as adjusting baiting regulations. If reducing depredations is an important goal, this analysis indicates that actions aside from (or in addition to) reducing wolf abundance might achieve that goal. This study also stresses the need to better understand the relationship among baiting practices, the effect of compensation on hunter behavior, and depredation occurrence.     

Wolf depredation on livestock in central Greece

We studied wolfCanis lupus Linnaeus, 1758 — livestock conflict in central Greece by investigating patterns of 267 verified wolf attacks on livestock for 21 months. Wolves attacked adult goats 43% and cattle 218% more than expected, whereas sheep 41% less than expected from their availability. Wolves killed less than four sheep or goats in 79%, and one cow or calf in 74% of depredation events, respectively. We recorded higher attack rates during wolf post-weaning season. Wolf attacks on strayed, or kept inside non predator-proof enclosures, sheep and goats, were on average two to four times respectively more destructive than those when livestock was guarded by a shepherd. Sheepdog use reduced losses per attack. Optimal sheepdog number ranged from 3 to 9 animals depending on flock size. Losses per attack were positively related to the number of wolves involved. Total losses per farm were positively correlated with the size of livestock unit but percentage losses per capita increased with decreasing flock size. Management implications to mitigate livestock depredation are discussed.     

Contrasting Activity Patterns of Sympatric and Allopatric Black and Grizzly Bears

ABSTRACT The distribution of grizzly (Ursus arctos) and American black bears (U. americanus) overlaps in western North America. Few studies have detailed activity patterns where the species are sympatric and no studies contrasted patterns where populations are both sympatric and allopatric. We contrasted activity patterns for sympatric black and grizzly bears and for black bears allopatric to grizzly bears, how human influences altered patterns, and rates of grizzly-black bear predation. Activity patterns differed between black bear populations, with those sympatric to grizzly bears more day-active. Activity patterns of black bears allopatric with grizzly bears were similar to those of female grizzly bears; both were crepuscular and day-active. Male grizzly bears were crepuscular and night-active. Both species were more night-active and less day-active when ≤1 km from roads or developments. In our sympatric study area, 2 of 4 black bear mortalities were due to grizzly bear predation. Our results suggested patterns of activity that allowed for intra- and inter-species avoidance. National park management often results in convergence of locally high human densities in quality bear habitat. Our data provide additional understanding into how bears alter their activity patterns in response to other bears and humans and should help park managers minimize undesirable bear-human encounters when considering needs for temporal and spatial management of humans and human developments in bear habitats.     

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.     

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1.?Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2.?Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3.?Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4.?In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5.?Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.     

Influence of Predator Harvest,Biological Factors,and Landscape on Elk Calf Survival in Idaho

ABSTRACT We evaluated survival of elk (Cervus elaphus) calves on 2 contrasting study areas in north-central Idaho, USA, from 1997 to 2004. Recruitment was modest (>30 calves:100 F [calves of either sex: F elk 1 yr old]) and stable on the South Fork study area and low (<20 calves:100 F) and declining on the Lochsa study area. The primary proximate cause of calf mortality on both study areas was predation by black bears (Ursus americanus) and mountain lions (Puma concolor). We experimentally manipulated populations of black bears and mountain lions on a portion of each study area. Black bear harvest (harvest density/600km²) initially doubled on the Lochsa treatment after manipulating season bag limits. Mountain lion harvest also increased by 60% but varied widely during the manipulation period. Harvest seasons were closed for black bears and mountain lions on the treatment portion of the South Fork study area. Using the Andersen—Gill formulation (A-G) of the Cox proportional hazards model, we examined effects of landscape structure, predator harvest levels, and biological factors on summer calf survival. We used Akaike's Information Criterion (AIC_c) and multimodel inference to assess some potentially useful predictive factors relative to calf survival. We generated risk ratios for both the best models and for model-averaged coefficients. Our models predicted that calf survival was influenced by biological factors, landscape surrounding calf locations, and predator harvest levels. The model that best explained mortality risk to calves on the Lochsa included black bear harvest (harvest density/600 km²), estimated birth mass of calves, and percentage of shrub cover surrounding calf locations. Incorporating a shrub X time interaction allowed us to correct for nonproportionality and detect that effect of shrub cover was only influential during the first 14 days of a calf's life. Model-averaging indicated that estimated birth mass of calves and black bear harvest were twice as important as the next variables, but age of calves at capture was also influential in calf survival. The model that best explained mortality risk to calves on the South Fork included black bear harvest, age of calves at capture, and gender of calves. Model-averaging indicated that age at capture and black bear harvest were twice as important as the next variable, forest with 33–66% canopy cover (Canopy 33–66). Risk to calves decreased when calves occupied areas with more of this forest cover type. Model-averaging also indicated that increased mountain lion harvest lowered calf mortality risk 4% for every 1-unit increase in lion harvest (harvest density/600 km²) but was lower (<25%) in importance compared to age at capture and black bear harvest. Our results suggest that levels of predator harvest, and presumably predator density, resource limitations expressed through calf birth mass, and habitat structure had substantial effects on calf survival. Our results can be generalized to other areas where managers are dealing with low calf elk recruitment. However, because factors vary spatially, a single management strategy applied in different areas will probably not have the same effect on calf survival.     

Modeling female brown bear kill rates on moose calves using global positioning satellite data

The recent development in Global Positioning System (GPS) techniques has started a new era in predation studies. Estimates of kill rates based on animal movements and GPS relocation clusters have proven to be valid in several obligatory carnivores. The main focus has been to obtain accurate mean predation estimates for the management of wildlife populations. We present a model to estimate individual kill rates of moose calves by adult female brown bears in Sweden, based on spatiotemporal clustering of 30,889 bear GPS relocations and 71 moose calves verified killed during 714 field investigations in 2004–2006. In this virtually single-predator single large prey system, the omnivorous brown bear is an efficient predator on moose calves up to 4 weeks of age. The top model set only included models with cluster radii of 30 m or 50 m, indicating very high kill-site fidelity. The best model included a cluster radius of 30 m and number of periods of bear activity at the kill site as a single covariate. The mean estimated individual kill rate of 7.6 ± 0.71 (n = 18, ± SE) moose calves per calving season is comparable to the estimate of 6.8 from a previous study of radio-tracked moose in our study area, though at a lower moose/bear ratio. The mean annual kill rates varied from 6.1 to 9.4 calves per bear. The estimated individual kill rates ranged from 2 to 15 calves per season, indicating a large individual variation in hunting skills and possibly effort. Predation and livestock depredation represent a core conflict between humans and carnivores in rural Scandinavia. Accurate predation estimates represent an important step in quantifying costs of carnivores and reducing human–carnivore conflicts. Our technique may be applied in the exploration of predation mechanisms and predator–prey interactions, and contribute to the old and global debate of problem individuals in livestock depredation. © 2012 The Wildlife Society.     

Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park

Grizzly bears (Ursus arctos) and American black bears (U. americanus) are sympatric in much of Yellowstone National Park. Three primary bear foods, cutthroat trout (Oncorhynchus clarki), whitebark pine (Pinus albicaulis) nuts, and elk (Cervus elaphus), have declined in recent years. Because park managers and the public are concerned about the impact created by reductions in these foods, we quantified bear diets to determine how bears living near Yellowstone Lake are adjusting. We estimated diets using: 1) stable isotope and mercury analyses of hair samples collected from captured bears and from hair collection sites established along cutthroat trout spawning streams and 2) visits to recent locations occupied by bears wearing Global Positioning System collars to identify signs of feeding behavior and to collect scats for macroscopic identification of residues. Approximately 45 ± 22% ( ± SD) of the assimilated nitrogen consumed by male grizzly bears, 38 ± 20% by female grizzly bears, and 23 ± 7% by male and female black bears came from animal matter. These assimilated dietary proportions for female grizzly bears were the same as 10 years earlier in the Lake area and 30 years earlier in the Greater Yellowstone Ecosystem. However, the proportion of meat in the assimilated diet of male grizzly bears decreased over both time frames. The estimated biomass of cutthroat trout consumed by grizzly bears and black bears declined 70% and 95%, respectively, in the decade between 1997–2000 and 2007–2009. Grizzly bears killed an elk calf every 4.3 ± 2.7 days and black bears every 8.0 ± 4.0 days during June. Elk accounted for 84% of all ungulates consumed by both bear species. Whitebark pine nuts continue to be a primary food source for both grizzly bears and black bears when abundant, but are replaced by false-truffles (Rhizopogon spp.) in the diets of female grizzly bears and black bears when nut crops are minimal. Thus, both grizzly bears and black bears continue to adjust to changing resources, with larger grizzly bears continuing to occupy a more carnivorous niche than the smaller, more herbivorous black bear. © 2012 The Wildlife Society.     

Summer elk calf survival in a partially migratory population

Decomposing variation in juvenile recruitment is a key component of understanding population dynamics for partially migratory ungulates. We investigated reproductive parameters of adult female elk (Cervus canadensis) with calves at heel, and survivorship, cause-specific mortality, and intrinsic and extrinsic factors affecting risk of mortality for calves in a partially migratory elk population from 2013–2016 in Alberta, Canada. Elk calves born to resident mothers had 45% lower survivorship on average compared to migrant calves (0.24 vs. 0.69) and nearly twice the mortality rate (0.37 vs. 0.19) from bears (Ursus spp.), the dominant source of mortality. Contrary to our predictions, we found that increasing levels of maternal ingesta-free body fat were associated with increasing risk of calf mortality, indicating predation may have overwhelmed nutritional effects. We found no evidence that timing of calf birth or birth weight differed between migratory tactics or influenced mortality risk. We found that as percentage of cut forest increased, risk of calf mortality marginally decreased, which benefited migrant elk that were exposed to more clear-cuts compared to residents. Exposure to bear predation risk was unimportant during the hiding phase (≤10 days after birth) for either migratory tactic, presumably because neonatal hiding behavior reduced vulnerability. In contrast, bear predation risk was important for mortality risk after 10 days in age, especially for resident elk calves, which were exposed to higher bear predation risk compared to migrants. We conclude that relative differences in bear predation between migratory tactics are contributing to the dynamics of partial migration in this population through additive effects on calf mortality. Thus, wildlife managers should anticipate that recovering grizzly bear (U. arctos) populations may substantially lower elk recruitment through effects on summer calf survival, especially in areas with diverse carnivore assemblages.     

Using spatially‐explicit capture–recapture models to explain variation in seasonal density patterns of sympatric ursids

Understanding how environmental factors interact to determine the abundance and distribution of animals is a primary goal of ecology, and fundamental to the conservation of wildlife populations. Studies of these relationships, however, often assume static environmental conditions, and rarely consider effects of competition with ecologically similar species. In many parts of their shared ranges, grizzly bears Ursus arctos and American black bears U. americanus have nearly complete dietary overlap and share similar life history traits. We therefore tested the hypothesis that density patterns of both bear species would reflect seasonal variation in available resources, with areas of higher primary productivity supporting higher densities of both species. We also hypothesized that interspecific competition would influence seasonal density patterns. Specifically, we predicted that grizzly bear density would be locally reduced due to the ability of black bears to more efficiently exploit patchy food resources such as seasonally abundant fruits. To test our hypotheses, we used detections of 309 grizzly and 597 black bears from two independent genetic sampling methods in spatially‐explicit capture–recapture (SECR) models. Our results suggest grizzly bear density was lower in areas of high black bear density during spring and summer, although intraspecific densities were also important, particularly during the breeding season. Black bears had lower densities in areas of high grizzly bear density in spring; however, density of black bears in early and late summer was best explained by primary productivity. Our results are consistent with the hypothesis that smaller‐bodied, more abundant black bears may influence the density patterns of behaviorally‐dominant grizzly bears through exploitative competition. We also suggest that seasonal variation in resource availability be considered in efforts to relate environmental conditions to animal density.     

Trends in intensive management of Alaska's grizzly bears, 1980–2010

Hunting regulations for grizzly bears (Ursus arctos) in much of Alaska since 1980 increasingly were designed to reduce bear abundance in the expectation such regulations would lead to increased harvests by hunters of moose (Alces alces) and caribou (Rangifer tarandus). Regulations were liberalized during 1980–2010 primarily in the area we termed the Liberal Grizzly Bear Hunting Area (hereafter Liberal Hunt Area) which encompassed 76.2% of Alaska. By 2010, these changes resulted in longer hunting seasons (100% of Liberal Hunt Area had seasons > 100 days, 99.7% > 200 days, and 67.8% > 300 days), more liberal bag limits (99.1% of the Liberal Hunt Area with a bag limit ≥ 1/yr and 10.1% with a bag limit ≥ 2/yr), and widespread waiver of resident tag fees (waived in 95.7% of the Liberal Hunt Area). During 1995–2010, there were 124 changes that made grizzly bear hunting regulations more liberal and two making them more conservative. The 4-year mean for grizzly bear kills by hunters increased 213% between 1976–1980 (387 grizzly bears) and 2005–2008 (823 grizzly bears). Since 2000, long-term research studies on grizzly populations in the Liberal Hunt Area have been terminated without replacement. Management of large predators by the State of Alaska is constrained by a 1994 state statute mandating “intensive management” in areas classified as important for human consumptive use of ungulates. Current grizzly bear management in the Liberal Hunt Area is inconsistent with the recommendations of the National Research Council's 1997 report on predator management in Alaska. Current attitudes, policies and absence of science-based management of grizzly bears in Alaska are increasingly similar to those that resulted in the near extirpation of grizzly bears south of Canada in the 19th and 20th centuries. If current trends continue, they increase risks to portions of the largest and most intact population of grizzly bears in North America. © 2011 The Wildlife Society.     

A Conservation-Based Approach to Compensation for Livestock Depredation: The Florida Panther Case Study

Calf (Bos taurus) depredation by the federally endangered Florida panther (Puma concolor coryi) on ranches in southwest Florida is an important issue because ranches represent mixed landscapes that provide habitat critical to panther recovery. The objectives of this study were to (1) quantify calf depredation by panthers on two ranches in southwest Florida, and (2) develop a habitat suitability model to evaluate the quality of panther hunting habitat on ranchlands, assess whether the model could predict predation risk to calves, and discuss its potential to be incorporated into an incentive-based compensation program. We ear-tagged 409 calves with VHF transmitters on two ranches during 2011–2013 to document calf mortality. We developed a model to evaluate the quality of panther hunting habitat on private lands in southwest Florida using environmental variables obtained from the Florida Natural Areas Inventory (FNAI) Cooperative Landcover Database and nocturnal GPS locations of panthers provided by the Florida Fish and Wildlife Conservation Commission (FWC). We then tested whether the model could predict the location of calf depredation sites. Tagged calf loss to panthers varied between the two ranches (0.5%/yr to 5.3%/yr) and may have been influenced by the amount of panther hunting habitat on each ranch as the ranch that experienced higher depredation rates contained a significantly higher probability of panther presence. Depredation sites of tagged calves had a significantly greater probability of panther presence than depredation sites of untagged calves that were found by ranchers in open pastures. This suggests that there may be more calves killed in high risk environments than are being found and reported by ranchers and that panthers can hunt effectively in open environments. It also suggests that the model may provide a means for evaluating the quality of panther hunting habitat and the corresponding risk of depredation to livestock across the landscape. We suggest that our approach could be applied to prioritize and categorize private lands for participation in a Payment for Ecosystem Services program that compensates landowners for livestock loss and incentivizes conserving high quality habitat for large carnivores where livestock depredation is a concern.     

Carnivore-livestock conflicts: determinants of wolf (Canis lupus) depredation on sheep farms in Finland

Wolves (Canis lupus) have recently expanded their distribution range into western and southern Finland, which has not hosted breeding wolves for over 100 years. This has raised concerns and public debate over wolf-livestock conflicts. Between 1998 and 2004 there were 45 wolf attacks on sheep on 34 farms. To assess the risk wolves may pose to sheep husbandry, we used data on depredation, sheep management, landscape structure and moose and wolf populations from continental Finland outside the area of reindeer husbandry to build models of the factors that may predispose sheep farms to wolf depredation. Our results provided evidence that sheep farms with the highest risk of wolf depredation were those located in regions where wolves were abundant. These farms were usually located close to the Russian border where the landscape is a mosaic of forest, wetlands and clear cut areas. These regions are sparsely populated by humans and farms are located far from each other. Finally, we generated probability maps based on generalised additive modelling to predict the risk of wolf predation on livestock in farms of southern Finland.     

Grizzly bear predation links the loss of native trout to the demography of migratory elk in Yellowstone

The loss of aquatic subsidies such as spawning salmonids is known to threaten a number of terrestrial predators, but the effects on alternative prey species are poorly understood. At the heart of the Greater Yellowstone ecosystem, an invasion of lake trout has driven a dramatic decline of native cutthroat trout that migrate up the shallow tributaries of Yellowstone Lake to spawn each spring. We explore whether this decline has amplified the effect of a generalist consumer, the grizzly bear, on populations of migratory elk that summer inside Yellowstone National Park (YNP). Recent studies of bear diets and elk populations indicate that the decline in cutthroat trout has contributed to increased predation by grizzly bears on the calves of migratory elk. Additionally, a demographic model that incorporates the increase in predation suggests that the magnitude of this diet shift has been sufficient to reduce elk calf recruitment (4–16%) and population growth (2–11%). The disruption of this aquatic–terrestrial linkage could permanently alter native species interactions in YNP. Although many recent ecological changes in YNP have been attributed to the recovery of large carnivores—particularly wolves—our work highlights a growing role of human impacts on the foraging behaviour of grizzly bears.     

Hazards Affecting Grizzly Bear Survival in the Greater Yellowstone Ecosystem

Abstract: During the past 2 decades, the grizzly bear (Ursus arctos) population in the Greater Yellowstone Ecosystem (GYE) has increased in numbers and expanded its range. Early efforts to model grizzly bear mortality were principally focused within the United States Fish and Wildlife Service Grizzly Bear Recovery Zone, which currently represents only about 61% of known bear distribution in the GYE. A more recent analysis that explored one spatial covariate that encompassed the entire GYE suggested that grizzly bear survival was highest in Yellowstone National Park, followed by areas in the grizzly bear Recovery Zone outside the park, and lowest outside the Recovery Zone. Although management differences within these areas partially explained differences in grizzly bear survival, these simple spatial covariates did not capture site-specific reasons why bears die at higher rates outside the Recovery Zone. Here, we model annual survival of grizzly bears in the GYE to 1) identify landscape features (i.e., foods, land management policies, or human disturbances factors) that best describe spatial heterogeneity among bear mortalities, 2) spatially depict the differences in grizzly bear survival across the GYE, and 3) demonstrate how our spatially explicit model of survival can be linked with demographic parameters to identify source and sink habitats. We used recent data from radiomarked bears to estimate survival (1983–2003) using the known-fate data type in Program MARK. Our top models suggested that survival of independent (age ≥ 2 yr) grizzly bears was best explained by the level of human development of the landscape within the home ranges of bears. Survival improved as secure habitat and elevation increased but declined as road density, number of homes, and site developments increased. Bears living in areas open to fall ungulate hunting suffered higher rates of mortality than bears living in areas closed to hunting. Our top model strongly supported previous research that identified roads and developed sites as hazards to grizzly bear survival. We also demonstrated that rural homes and ungulate hunting negatively affected survival, both new findings. We illustrate how our survival model, when linked with estimates of reproduction and survival of dependent young, can be used to identify demographically the source and sink habitats in the GYE. Finally, we discuss how this demographic model constitutes one component of a habitat-based framework for grizzly bear conservation. Such a framework can spatially depict the areas of risk in otherwise good habitat, providing a focus for resource management in the GYE.     

Habitat use by black bears in relation to conspecifics and competitors

Sympatric black bears (Ursus americanus) and brown bears (Ursus arctos) are common in many boreal systems; however, few predator assemblages are known to coexist on a single seasonally abundant large prey item. In lowland southwestern interior Alaska, black bears and brown bears are considered the primary cause of moose (Alces alces) calf mortality during the first 6 weeks of life. The objective of this study was to document habitat use of global-positioning system (GPS)-collared black bears during peak and non-peak seasons of black bear-induced and brown bear-induced moose calf mortality within southwestern interior Alaska, in spring 2002. We compared habitats of GPS-collared black bears to those of presumably uncollared black bears and brown bears at their moose calf mortality sites. Results from this study suggest that GPS-collared black bears use similar habitat as conspecifics more than expected during the peak period of black bear predation on moose calves, whereas they use habitat in proportion to home range availability during the peak in brown bear predation on moose calves. Sex-specific Ivlev's electivity indices describe greater than expected use of mixed-deciduous forest and needleleaf forest by male GPS-collared black bears during the peak of moose calf predation, whereas females have a tendency to use these habitats less than expected. Juvenile GPS-collared black bears largely use the same habitat as other sympatric predators during the peak of moose calf predation, whereas during the non-peak period juveniles use opposite habitats as adult GPS-collared black bears. The outcome of this study offers possible explanations (e.g., sex, age) for spatial overlap or segregation in one member of a complex predator guild in relation to a seasonal pulse of preferred prey.     

Effects of wolf pack size and winter conditions on elk mortality

Elk (Cervus canadensis) are high-profile game animals for many states in the western United States, yet over the past several decades some populations have experienced a persistent and broad-scale decline in recruitment. Over this same period, gray wolves (Canis lupus) have become an integral component of many western landscapes and agencies are increasingly challenged to maximize hunting opportunities of ungulates via predator management while simultaneously ensuring wolf conservation. To better understand the implications of predator management on elk populations, we monitored survival of 1,244 adult female elk and 806 6-month-old calves from 29 populations distributed throughout Idaho, USA, from 2004 to 2016. We developed predictive models of mortality that related mortality risk to wolf pack size, winter conditions, and individual-level characteristics. Annual mortality rates (excluding harvest) for adult females and calves were 0.09 and 0.40, respectively. Calf mortality was predicted best with a model that included additive effects of chest girth at time of capture, mean size of surrounding wolf packs, and snow depth. Adult female mortality was predicted best with a model that included female age, mean size of surrounding wolf packs, and snow depth. Based on a sensitivity analysis, chest girth had the largest effect on risk of mortality for calves followed by pack size and snow depth. Other than the effect of senescence in the oldest (>15 yr) individuals, pack size and snow depth had the largest effect on risk of mortality for adult females. We estimated cause-specific mortality and predation was the dominant cause of known-fate mortalities for adult females (35% mountain lion [Puma concolor] and 32% wolf) and calves (45% mountain lion and 28% wolf), whereas malnutrition accounted for 9% and 10% of adult female and calf mortalities, respectively. Wolves preferentially selected smaller calves and older adult females, whereas mountain lions showed little preference for calf size or age class of adult females. Our study indicates managers can increase elk survival by reducing wolf pack sizes on surrounding winter ranges, especially in areas where, or during years when, snow is deep. Additionally, managers interested in improving over-winter calf survival can implement actions to increase the size of calves entering winter by increasing the nutritional quality of summer and early fall forage resources. Although our study was prompted by management questions related to wolves, mountain lions killed more elk than wolves and differences in selection of individual elk indicate mountain lions may have comparably more of an effect on elk population dynamics. Although we were unable to relate changes in mountain lion populations to elk survival in our study, future research should seek a better understanding of multi-predator systems, including how management of one predator affect others and ultimately how these interactions affect elk survival. © 2019 The Wildlife Society     

No long-term effect of black bear removal on elk calf recruitment in the southern Appalachians

In 2001 and 2002, 52 elk (Cervus canadensis; 21 males, 31 females), originally obtained from Elk Island National Park, Alberta, Canada, were transported and released into Cataloochee Valley in the northeastern portion of Great Smoky Mountains National Park (GRSM, Park), North Carolina, USA. The annual population growth rate (λ) was negative (0.996, 95% CI = 0.945–1.047) and predation by black bears (Ursus americanus) on elk calves was identified as an important determinant of population growth. From 2006 to 2008, 49 bears from the primary elk calving area (i.e., Cataloochee Valley) were trapped and translocated about 70 km to the southwestern portion of the Park just prior to elk calving. Per capita recruitment (i.e., the number of calves produced per adult female that survive to 1 year of age) increased from 0.306 prior to bear translocation (2001–2005) to 0.544 during years when bears were translocated (2006–2008) and λ increased to 1.118 (95% CI = 1.096–1.140). Our objective was to determine whether per capita calf recruitment rates after bear removal (2009–2019) at Cataloochee were similar to the higher rates estimated during bear removal (i.e., long-term response) or if they returned to rates before bear removal (i.e., short-term response), and how those rates compared with recruitment from portions of our study area where bears were not relocated. We documented 419 potential elk calving events and monitored 129 yearling and adult elk from 2001 to 2019. Known-fate models based on radio-telemetry and observational data supported calf recruitment returning to pre-2006 levels at Cataloochee (short-term response); recruitment of Cataloochee elk before and after bear relocation was lower (0.184) than during bear relocation (0.492). Recruitment rates of elk outside the removal area during the bear relocation period (0.478) were similar to before and after rates (0.420). In the Cataloochee Valley, cause-specific annual calf mortality rates due to predation by bears were 0.319 before, 0.120 during, and 0.306 after bear relocation. In contrast, the cause-specific annual mortality rate of calves in areas where bears were not relocated was 0.033 after the bear relocation period, with no bear predation on calves before or during bear relocation. The mean annual population growth rate for all monitored elk was 1.062 (95% CI = 0.979–1.140) after bear relocation based on the recruitment and survival data. Even though the effects of bear removal were temporary, the relocations were effective in achieving a short-term increase in elk recruitment, which was important for the reintroduction program given that the elk population was small and vulnerable to extirpation.