Corridor-based approach with spatial cross-validation reveals scale-dependent effects of geographic distance, human footprint and canopy cover on grizzly bear genetic connectivity.

Understanding how human infrastructure and other landscape attributes affect genetic differentiation in animals is an important step for identifying and maintaining dispersal corridors for these species. We built upon recent advances in the field of landscape genetics by using an individual-based and multiscale approach to predict landscape-level genetic connectivity for grizzly bears (Ursus arctos) across ~100,000 km2 in Canada's southern Rocky Mountains. We used a genetic dataset with 1156 unique individuals genotyped at nine microsatellite loci to identify landscape characteristics that influence grizzly bear gene flow at multiple spatial scales and map predicted genetic connectivity through a matrix of rugged terrain, large protected areas, highways and a growing human footprint. Our corridor-based modelling approach used a machine learning algorithm that objectively parameterized landscape resistance, incorporated spatial cross validation and variable selection and explicitly accounted for isolation by distance. This approach avoided overfitting, discarded variables that did not improve model performance across withheld test datasets and spatial predictive capacity compared to random cross-validation. We found that across all spatial scales, geographic distance explained more variation in genetic differentiation in grizzly bears than landscape variables. Human footprint inhibited connectivity across all spatial scales, while open canopies inhibited connectivity at the broadest spatial scale. Our results highlight the negative effect of human footprint on genetic connectivity, provide strong evidence for using spatial cross-validation in landscape genetics analyses and show that multiscale analyses provide additional information on how landscape variables affect genetic differentiation.

Toxic elements in arctic and sub-arctic brown bears: Blood concentrations of As, Cd, Hg and Pb in relation to diet, age, and human footprint.

Contamination with arsenic (As), cadmium (Cd), mercury (Hg) and lead (Pb) is a global concern impairing resilience of organisms and ecosystems. Proximity to emission sources increases exposure risk but remoteness does not alleviate it. These toxic elements are transported in atmospheric and oceanic pathways and accumulate in organisms. Mercury accumulates in higher trophic levels. Brown bears (Ursus arctos), which often live in remote areas, are long-lived omnivores, feeding on salmon (Oncorhynchus spp.) and berries (Vaccinium spp.), resources also consumed by humans. We measured blood concentrations of As, Cd, Hg and Pb in bears (n = 72) four years and older in Scandinavia and three national parks in Alaska, USA (Lake Clark, Katmai and Gates of the Arctic) using high-resolution, inductively-coupled plasma sector field mass spectrometry. Age and sex of the bears, as well as the typical population level diet was associated with blood element concentrations using generalized linear regression models. Alaskan bears consuming salmon had higher Hg blood concentrations compared to Scandinavian bears feeding on berries, ants (Formica spp.) and moose (Alces). Cadmium and Pb blood concentrations were higher in Scandinavian bears than in Alaskan bears. Bears using marine food sources, in addition to salmon in Katmai, had higher As blood concentrations than bears in Scandinavia. Blood concentrations of Cd and Pb, as well as for As in female bears increased with age. Arsenic in males and Hg concentrations decreased with age. We detected elevated levels of toxic elements in bears from landscapes that are among the most pristine on the planet. Sources are unknown but anthropogenic emissions are most likely involved. All study areas face upcoming change: Increasing tourism and mining in Alaska and more intensive forestry in Scandinavia, combined with global climate change in both regions. Baseline contaminant concentrations as presented here are important knowledge in our changing world.

Survey for Selected Parasites in Alaska Brown Bears (Ursus arctos).

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

Supplementing Environmental Assessments with Cumulative Effects Scenario Modeling for Grizzly Bear Connectivity in the Bow Valley, Alberta, Canada.

Persistence of sensitive wildlife in populated regions requires conservation strategies that address gradual expansion of development footprint and human activity. The project-based environmental assessment regime for municipal development is poorly suited to provide necessary strategic perspective, given its focus on local and short-term impacts. We used the ALCES cumulative effects model to strategically assess impacts to grizzly bears (Ursus arctos) in the Bow Valley of Alberta, Canada. Landscape simulation mapped expansion of past and potential future development footprint in the region over multiple decades. Consequences to movement connectivity for grizzly bears were estimated by applying a least cost path analysis to the landscape simulation. An index of recreational activity was derived from fitness tracking data and integrated with the landscape simulation to model change in recreational activity through time. Maps of grizzly bear connectivity and recreational activity were combined to calculate human-bear conflict risk. The analysis suggests that connectivity has been altered through displacement to upslope areas by settlement expansion, such that surrounding natural areas have become important for grizzly bear connectivity. These areas are also popular for outdoor recreation, resulting in elevated human-bear conflict risk which can be expected to increase if development and human activity continue to expand in high connectivity areas. Conservation of wildlife in populated regions will be supported by broadening the scope of environmental assessment to address cumulative effects of development footprint and human activity over large spatial and temporal scales.

How Can We Do Citizen Science Better? A Case Study Evaluating Grizzly Bear Citizen Science Using Principles of Good Practice in Alberta, Canada.

Citizen science offers an excellent opportunity to engage the public in scientific data collection, educational opportunities, and applied management. However, the practicalities of developing and implementing citizen science programming are often more complex than considered. Some challenges to effective citizen science include scientists' skepticism about the ability of public participants to rigorously collect quality data; a lack of clarity on or confidence in the utility of data; scientists' hesitancy in engaging the public in projects; limited financial commitments; and challenges associated with the temporal and geographic scales of projects. To address these challenges, and provide a foundation upon which practitioners, scientists, and the public can credibly engage in citizen science, the Government of Alberta developed a set of citizen science principles. These principles offer a framework for planning, designing, implementing, and evaluating citizen science projects that extend beyond Alberta. Here, we present a case study using these principles to evaluate GrizzTracker, a citizen science program developed to help inform provincial species-at-risk recovery efforts. While we found that GrizzTracker applied each of the six principles in some way, including successful public engagement, strengthened relationships, and raising public awareness about northwest Alberta's grizzly bears, we also identified a number of challenges. These included ongoing skepticism from the traditional scientific community about the utility of citizen science and governance challenges related to program leadership, staff capacity, and funding. By using the principles as a guideline, we provide policy recommendations for future citizen science efforts, including considerations for program design, implementation, and evaluation.

Conservation and management of the culture of bears.

Culture is widely accepted as an important social factor present across a wide range of species. Bears have a culture as defined as behavioral traditions inherited through social learning usually from mothers to offspring. Successful bear cultures can enhance fitness and resource exploitation benefits. In contrast, some bear cultures related to response to humans and human-related foods can be maladaptive and result in reduced fitness and direct mortality. In environments with minimal human influence most bear culture has evolved over generations to be beneficial and well adapted to enhance fitness. However, most bears across the world do not live in areas with minimal human influence and in these areas, bear culture is often changed by bear interactions with humans, usually to the detriment of bear survival. We highlight the importance of identifying unique bear cultural traits that allow efficient use of local resources and the value of careful management to preserve these adaptive cultural behaviors. It is also important to select against maladaptive cultural behaviors that are usually related to humans in order to reduce human-bear conflicts and high bear mortality. We use examples from Yellowstone National Park to demonstrate how long-term management to reduce maladaptive bear cultures related to humans has resulted in healthy bear populations and a low level of human-bear conflict in spite of a high number of Yellowstone National Park visitors in close association with bears.

About pace: How variations in method and definition affect quantification of pacing in bears.

Repetitive pacing behavior is exhibited by many species in zoos and is particularly prevalent in carnivores with large home ranges, such as bears. Pacing can be a behavioral indicator of poor welfare, however, understanding this behavior can be challenging. As many bears that pace are singly housed, efforts to systematically examine and ameliorate pacing may be strengthened by multi-institutional studies. However, there is currently no standardized method to quantify pacing, which makes cross-institutional analyses of causal factors and intervening measures challenging. The purpose of this study was to compare multiple sampling methods and definitions for quantifying pacing in bears to understand how they affect outcome measures. We analyzed video recordings of two grizzly and two black bears pacing, using three sampling methods (continuous, instantaneous 30-s interval, instantaneous 1-min interval), and three definitions of pacing (AB-two repetitions of the path, ABA-three repetitions, ABAB-four repetitions). A generalized linear mixed model revealed that continuous and instantaneous 30-s interval methods captured more pacing than instantaneous 1-min methods, and definitions captured a decreasing amount of pacing from AB to ABA to ABAB. AB also captured the highest number of pacing bouts. The importance of comparability across institutions is growing, and a standard methodology and definition for recording pacing would be useful. We suggest that the combination of instantaneous sampling and the ABA definition presents a good balance between capturing the right data and being flexible enough for a variety of institutions to implement.

INVESTIGATION OF SARCOCYSTIS SPP. INFECTION IN FREE-RANGING AMERICAN BLACK BEARS (URSUS AMERICANUS) AND GRIZZLY BEARS (URSUS ARCTOS HORRIBILIS) IN BRITISH COLUMBIA, CANADA.

Sarcocystis spp. are protozoan parasites that cause a spectrum of lesions in various hosts. Hepatic sarcocystosis and encephalitis have been described in captive American black bears (Ursus americanus) and polar bears (Ursus maritimus), and in a free-ranging grizzly bear (Ursus arctos horribilis), but have not previously been reported in free-ranging American black bears. This study aimed to characterize the presence and lesions associated with Sarcocystis spp. in free-ranging bears in British Columbia, Canada from samples submitted to the provincial diagnostic laboratory. From 2007 to 2019, 102 free-ranging American black bear and grizzly bear tissues were examined postmortem for sarcocystosis using histopathology and follow-up molecular diagnostics. Sarcocystosis was confirmed in 41 (40%) free-ranging bears including 39 American black bears and two grizzly bears. Microscopic lesions included multifocal necrotizing hepatitis, nonsuppurative encephalitis, and/or intramuscular sarcocysts with or without associated inflammation. Sarcocystosis was considered the cause of death in eight (20%) of these bears, exclusively in cubs of the year (<1 yr old). Sarcocystis canis was identified in 22/32 (69%) cases where molecular characterization was performed and was the etiologic agent associated with bears that died of sarcocystosis. Confirmed cases were distributed widely across British Columbia. While there was an alternate proximate cause of death in the other confirmed bears, sarcocystosis may have contributed. Age was a significant risk factor, with yearlings presenting more often with fulminant lesions; however, there was a sampling bias toward juvenile bear submissions due to size and ease of transport. Further research is needed to understand the disease epidemiology and significance to population health.

Cortisol levels in blood and hair of unanesthetized grizzly bears (Ursus arctos) following intravenous cosyntropin injection.

Hair cortisol concentration (HCC) is being used increasingly to evaluate long-term stress in many mammalian species. Most of the cortisol is assumed to passively diffuse from circulating blood into hair follicles and gradually accumulate in growing hair. However, our research with free-ranging grizzly bears (Ursus arctos) suggests HCC increases significantly within several hours following capture, a time too brief to be explained by this mechanism alone. In this study with captive grizzly bears, we sought to determine if a brief spike in blood cortisol concentration, thus mimicking a single stressful event, would cause an increase in HCC over a 7-day period. To do this, we administered a single intravenous dose (5 µg/kg) of cosyntropin to three captive unanaesthetised adult female grizzly bears on two occasions, during April when hair growth was arrested and during August when hair was growing. In both trials, the cosyntropin caused a two-fold or greater increase in serum cortisol levels within 1 hr but did not appear to influence HCC at 1, 48, and 168 hr following cosyntropin administration. We conclude the cosyntropin-induced cortisol spike was likely insignificant when compared to the adrenocortical response that occurs in free-ranging bears when captured. We suggest further study with a larger sample of captive bears to evaluate the combined effects of anaesthesia and multiple doses of cosyntropin administered over several hours would better simulate the adrenocortical response of free-ranging grizzly bears during capture.

Vital rates of two small populations of brown bears in Canada and range-wide relationship between population size and trend.

Identifying mechanisms of population change is fundamental for conserving small and declining populations and determining effective management strategies. Few studies, however, have measured the demographic components of population change for small populations of mammals (<50 individuals). We estimated vital rates and trends in two adjacent but genetically distinct, threatened brown bear (Ursus arctos) populations in British Columbia, Canada, following the cessation of hunting. One population had approximately 45 resident bears but had some genetic and geographic connectivity to neighboring populations, while the other population had <25 individuals and was isolated. We estimated population-specific vital rates by monitoring survival and reproduction of telemetered female bears and their dependent offspring from 2005 to 2018. In the larger, connected population, independent female survival was 1.00 (95% CI: 0.96-1.00) and the survival of cubs in their first year was 0.85 (95% CI: 0.62-0.95). In the smaller, isolated population, independent female survival was 0.81 (95% CI: 0.64-0.93) and first-year cub survival was 0.33 (95% CI: 0.11-0.67). Reproductive rates did not differ between populations. The large differences in age-specific survival estimates resulted in a projected population increase in the larger population (λ = 1.09; 95% CI: 1.04-1.13) and population decrease in the smaller population (λ = 0.84; 95% CI: 0.72-0.95). Low female survival in the smaller population was the result of both continued human-caused mortality and an unusually high rate of natural mortality. Low cub survival may have been due to inbreeding and the loss of genetic diversity common in small populations, or to limited resources. In a systematic literature review, we compared our population trend estimates with those reported for other small populations (<300 individuals) of brown bears. Results suggest that once brown bear populations become small and isolated, populations rarely increase and, even with intensive management, recovery remains challenging.

High Prevalence and Intensity of Trichinella Infection in Yukon American Black (Ursus americanus) and Grizzly (Ursus arctos) Bears.

Trichinella spp. nematodes are commonly found in bear species (Ursidae) and can pose severe health risks to humans when infective first-stage larvae are ingested in meat. Samples of tongue or masseter muscle from 22 male and 11 female American black bears (Ursus americanus; mean age 6.5 yr, range 1-16 yr) and 22 male, eight female, and one unknown sex grizzly bears (Ursus arctos; mean age 8.8 yr, range 2-28 yr), from Yukon, Canada, were tested to determine prevalence and intensity of Trichinella spp. infection. For black bears, prevalence was 20% and mean intensity was 401 larvae per gram of tissue (LPG), whereas for grizzly bears, prevalence was 71%, and mean infection intensity was 35 LPG. Isolates from all positive samples were identified as genotype Trichinella-T6 by multiplex PCR. For black bears, prevalence is the highest reported in Canada and infection intensity the highest recorded in North America. One black bear had a larval burden of 1,173 LPG, the second highest recorded in any host species. The prevalence in grizzly bears was the highest reported in Canada for this host. In total, 90% (27 of 30) of infected bears had infection burdens above the human food safety threshold of ≥1 LPG, reinforcing the importance of communicating the health risks to people consuming bear meat.

Automated facial recognition for wildlife that lack unique markings: A deep learning approach for brown bears.

Emerging technologies support a new era of applied wildlife research, generating data on scales from individuals to populations. Computer vision methods can process large datasets generated through image-based techniques by automating the detection and identification of species and individuals. With the exception of primates, however, there are no objective visual methods of individual identification for species that lack unique and consistent body markings. We apply deep learning approaches of facial recognition using object detection, landmark detection, a similarity comparison network, and an support vector machine-based classifier to identify individuals in a representative species, the brown bear Ursus arctos. Our open-source application, BearID, detects a bear's face in an image, rotates and extracts the face, creates an "embedding" for the face, and uses the embedding to classify the individual. We trained and tested the application using labeled images of 132 known individuals collected from British Columbia, Canada, and Alaska, USA. Based on 4,674 images, with an 80/20% split for training and testing, respectively, we achieved a facial detection (ability to find a face) average precision of 0.98 and an individual classification (ability to identify the individual) accuracy of 83.9%. BearID and its annotated source code provide a replicable methodology for applying deep learning methods of facial recognition applicable to many other species that lack distinguishing markings. Further analyses of performance should focus on the influence of certain parameters on recognition accuracy, such as age and body size. Combining BearID with camera trapping could facilitate fine-scale behavioral research such as individual spatiotemporal activity patterns, and a cost-effective method of population monitoring through mark-recapture studies, with implications for species and landscape conservation and management. Applications to practical conservation include identifying problem individuals in human-wildlife conflicts, and evaluating the intrapopulation variation in efficacy of conservation strategies, such as wildlife crossings.

The ecology of human-carnivore coexistence.

With a shrinking supply of wilderness and growing recognition that top predators can have a profound influence on ecosystems, the persistence of large carnivores in human-dominated landscapes has emerged as one of the greatest conservation challenges of our time. Carnivores fascinate society, yet these animals pose threats to people living near them, resulting in high rates of carnivore death near human settlements. We used 41 y of demographic data for more than 2,500 brown bears-one of the world's most widely distributed and conflict-prone carnivores-to understand the behavioral and demographic mechanisms promoting carnivore coexistence in human-dominated landscapes. Bear mortality was high and unsustainable near people, but a human-induced shift to nocturnality facilitated lower risks of bear mortality and rates of conflict with people. Despite these behavioral shifts, projected population growth rates for bears in human-dominated areas revealed a source-sink dynamic. Despite some female bears successfully reproducing in the sink areas, bear persistence was reliant on a supply of immigrants from areas with minimal human influence (i.e., wilderness). Such mechanisms of coexistence reveal a striking paradox: Connectivity to wilderness areas supplies bears that likely will die from people, but these bears are essential to avert local extirpation. These insights suggest carnivores contribute to human-carnivore coexistence through behavioral and demographic mechanisms, and that connected wilderness is critical to sustain coexistence landscapes.

Railway-Associated Attractants as Potential Contaminants for Wildlife.

Grizzly bears (Ursus arctos) appear to be attracted to natural and anthropogenic forage along railways, which may increase collision vulnerability, but also potentially causes exposure to contaminants associated with railway infrastructure. We assessed contaminant exposure for a vulnerable population of grizzly bears in the Canadian Rocky Mountains by determining if (1) dandelions (Taraxacum officinale) growing adjacent to a railway and grain spilled from hopper cars contain heavy metals, polycyclic aromatic hydrocarbons (PAHs), and mycotoxins and (2) metal concentrations from hair samples of individual bears correlates with use of the railway or other anthropogenic features. We used principle components analysis to represent 10 heavy metals and 16 PAHs and then compared their concentrations in railway-associated sources of grain and dandelions to reference samples that we purchased (grain) or sampled from nearby sites (dandelions). We also measured metal concentrations in the hair of bears that were captured and fitted with GPS collars. We found significantly higher concentrations in railway-associated samples of dandelion and grain for both metals (particularly lead, iron, and chromium), and the sum of 16 PAHs. Several metals and PAHs in railway-associated samples exceeded regulatory standards for soil or animal feed. Mycotoxins were detectable in grain samples, but occurred well below permissible standards. Metal concentrations in bear hair were not predicted by railway use, but higher metal concentrations occurred in male bears and two individuals that used ski hills during fall. As mitigation to reduce wildlife exposure to contaminants, particularly in protected areas, we encourage removal of railway grain deposits, regular maintenance of railway infrastructure, such as lubricating stations, and investigation of contaminants associated with other human infrastructures, such as ski hills.

Salmonid species diversity predicts salmon consumption by terrestrial wildlife.

Resource waves-spatial variation in resource phenology that extends feeding opportunities for mobile consumers-can affect the behaviour and productivity of recipient populations. Interspecific diversity among Pacific salmon species (Oncorhynchus spp.) creates staggered spawning events across space and time, thereby prolonging availability to terrestrial wildlife. We sought to understand how such variation might influence consumption by terrestrial predators compared with resource abundance and intra- and interspecific competition. Using stable isotope analysis, we investigated how the proportion of salmon in the annual diet of male black bears (Ursus americanus; n = 405) varies with species diversity and density of spawning salmon biomass, while also accounting for competition with sympatric black and grizzly bears (U. arctos horribilis), in coastal British Columbia, Canada. We found that the proportion of salmon in the annual diet of black bears was ≈40% higher in the absence of grizzly bears, but detected little effect of relative black bear density and salmon biomass density. Rather, salmon diversity had the largest positive effect on consumption. On average, increasing diversity from one salmon species to ~four (with equal biomass contributions) approximately triples the proportion of salmon in diet. Given the importance of salmon to bear life histories, this work provides early empirical support for how resource waves may increase the productivity of consumers at population and landscape scales. Accordingly, terrestrial wildlife management might consider maintaining not only salmon abundance but also diversity.

The Genome of the North American Brown Bear or Grizzly: Ursus arctos ssp. horribilis.

The grizzly bear (Ursus arctos ssp. horribilis) represents the largest population of brown bears in North America. Its genome was sequenced using a microfluidic partitioning library construction technique, and these data were supplemented with sequencing from a nanopore-based long read platform. The final assembly was 2.33 Gb with a scaffold N50 of 36.7 Mb, and the genome is of comparable size to that of its close relative the polar bear (2.30 Gb). An analysis using 4104 highly conserved mammalian genes indicated that 96.1% were found to be complete within the assembly. An automated annotation of the genome identified 19,848 protein coding genes. Our study shows that the combination of the two sequencing modalities that we used is sufficient for the construction of highly contiguous reference quality mammalian genomes. The assembled genome sequence and the supporting raw sequence reads are available from the NCBI (National Center for Biotechnology Information) under the bioproject identifier PRJNA493656, and the assembly described in this paper is version QXTK01000000.

Estimating unrecorded human-caused mortalities of grizzly bears in the Flathead Valley, British Columbia, Canada.

Managing the number of grizzly bear (Ursus arctos) mortalities to a sustainable level is fundamental to bear conservation. All known grizzly bear deaths are recorded by management agencies but the number of human-caused grizzly bear deaths that are not recorded is generally unknown, causing considerable uncertainty in the total number of mortalities. Here, we compare the number of bears killed legally by hunters to the number killed by people for all other reasons, for bears wearing functioning radiocollars and for uncollared bears recorded in the British Columbia (BC) government mortality database for the Flathead Valley in southeast BC. Between 1980 and 2016, permitted hunters killed 10 collared bears and 12 (9 known, 3 suspected) were killed by people for other reasons. This ratio differed (p < 0.0001) from the uncollared bears in the government database where 71 were killed by hunters while only 10 were killed for other reasons. We estimate that 88% (95% CI; 67-96%) of the human-caused mortalities that were not by permitted hunters were unreported. The study area may have low reporting rates because it is >40 km on a gravel road from a Conservation Officer office, so reporting is difficult and there are no human residences so there is little concern of a neighbor contacting an officer. Our results are likely indicative of other places that are road-accessed but far from settlements. We discuss the implications of sampling individuals for collaring and the possible implications of wearing a collar on the animal's fate.

Functional macronutritional generalism in a large omnivore, the brown bear.

We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log-ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log-ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human-sourced foods have different dietary macronutrient proportions relative to populations with natural diets.

Are pumas subordinate carnivores, and does it matter?

BACKGROUND: Interspecific competition affects species fitness, community assemblages and structure, and the geographic distributions of species. Established dominance hierarchies among species mitigate the need for fighting and contribute to the realized niche for subordinate species. This is especially important for apex predators, many of which simultaneous contend with the costs of competition with more dominant species and the costs associated with human hunting and lethal management. METHODS: Pumas are a widespread solitary felid heavily regulated through hunting to reduce conflicts with livestock and people. Across their range, pumas overlap with six apex predators (gray wolf, grizzly bear, American black bear, jaguar, coyote, maned wolf), two of which (gray wolf, grizzly bear) are currently expanding in North America following recovery efforts. We conducted a literature search to assess whether pumas were subordinate or dominant with sympatric apex predators, as well as with three felid mesocarnivores with similar ecology (ocelot, bobcat, Canada lynx). We also conducted an analysis of the spatial distributions of pumas and their dominant sympatric competitors to estimate in what part of their range, pumas are dominant versus subordinate. RESULTS: We used 64 sources to assess dominance among pumas and other apex predators, and 13 sources to assess their relationships with felid mesocarnivores. Evidence suggested that wolves, grizzly bears, black bears, and jaguars are dominant over pumas, but that pumas are dominant over coyotes and maned wolves. Evidence suggested that pumas are also dominant over all three felid mesocarnivores with which they share range. More broadly, pumas are subordinate to at least one other apex carnivore in 10,799,252 (47.5%) of their 22,735,268 km2 range across North and South America. DISCUSSION: Subordinate pumas change their habitat use, suffer displacement at food sources, likely experience increased energetic demands from harassment, exhibit increased starvation, and are sometimes directly killed in competitive interactions with dominant competitors. Nevertheless, we lack research clearly linking the costs of competition to puma fitness. Further, we lack research that assesses the influence of human effects simultaneous with the negative effects of competition with other sympatric carnivores. Until the time that we understand whether competitive effects are additive with human management, or even potentially synergistic, we encourage caution among managers responsible for determining harvest limits for pumas and other subordinate, apex carnivores in areas where they are sympatric with dominant species. This may be especially important information for managers working in regions where wolves and brown bears are recolonizing and recovering, and historic competition scenarios among multiple apex predators are being realized.

Space-time clusters for early detection of grizzly bear predation.

Accurate detection and classification of predation events is important to determine predation and consumption rates by predators. However, obtaining this information for large predators is constrained by the speed at which carcasses disappear and the cost of field data collection. To accurately detect predation events, researchers have used GPS collar technology combined with targeted site visits. However, kill sites are often investigated well after the predation event due to limited data retrieval options on GPS collars (VHF or UHF downloading) and to ensure crew safety when working with large predators. This can lead to missing information from small-prey (including young ungulates) kill sites due to scavenging and general site deterioration (e.g., vegetation growth). We used a space-time permutation scan statistic (STPSS) clustering method (SaTScan) to detect predation events of grizzly bears (Ursus arctos) fitted with satellite transmitting GPS collars. We used generalized linear mixed models to verify predation events and the size of carcasses using spatiotemporal characteristics as predictors. STPSS uses a probability model to compare expected cluster size (space and time) with the observed size. We applied this method retrospectively to data from 2006 to 2007 to compare our method to random GPS site selection. In 2013-2014, we applied our detection method to visit sites one week after their occupation. Both datasets were collected in the same study area. Our approach detected 23 of 27 predation sites verified by visiting 464 random grizzly bear locations in 2006-2007, 187 of which were within space-time clusters and 277 outside. Predation site detection increased by 2.75 times (54 predation events of 335 visited clusters) using 2013-2014 data. Our GLMMs showed that cluster size and duration predicted predation events and carcass size with high sensitivity (0.72 and 0.94, respectively). Coupling GPS satellite technology with clusters using a program based on space-time probability models allows for prompt visits to predation sites. This enables accurate identification of the carcass size and increases fieldwork efficiency in predation studies.