Summary 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.

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 (AICc) 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 recruitmen...

Genetic data are increasingly used to describe the structure of wildlife populations and to infer landscape influences on functional connectivity. To accomplish this, genetic structure can be described with a multitude of methods that vary in their assumptions, advantages and limitations. While some methods discriminate distinct subpopulations separated by sharp genetic boundaries (i.e. barrier detection or clustering methods), other methods estimate gradient genetic structures using individual-based genetic distances. We present an analytical framework based on individual ancestry values that combines these different approaches and can be used to a) test for local barriers to gene flow and b) evaluate effects of landscape gradients through individual-based genetic distances that account for hierarchical genetic structure. We illustrate the approach with a data set of 371 cougars Puma concolor from a 217 000 km 2 study area in Idaho and western Montana (USA) that were genotyped at 12 microsatellite loci. Results suggest that cougars in the region show a complex, hierarchical genetic structure that is influenced by a local barrier to gene flow (an urban population cluster connected by high traffic volumes), different landscape features (the Snake River Plain, forested habitat), and geographic distance. Our novel approach helped to elucidate the relative influence of these factors on different hierarchical levels of population structure, which was not possible when using either clustering methods or standard genetic distances. Results obtained with our analytical framework highlight the need for multi-scale management of cougars in the region and show that landscape heterogeneity can create complex genetic structures, even in generalist species with high dispersal capabilities.

The identification of carnivores responsible for preying on wild or domestic ungulates often is of interest to wildlife managers. Typically, field personnel collect a variety of data at mortality sites including scat or hair samples that may have been deposited by the predator. We compared mitochondrial DNA (mtDNA) analysis of hair and scat samples (n = 122) collected at elk (Cervus elaphus) mortality sites between 1997 and 2004 in north‐central Idaho, USA, with field identification of carnivore presence. We amplified mtDNA from samples via a 2‐step process involving an initial screening for American black bears (Ursus americanus), brown bears (Ursus arctos), and gray wolves (Canis lupus) using a length variation in the 5′ hypervariable section of the control region. Samples that failed the first screening subsequently were analyzed using conserved mtDNA primers that amplify a wide array of vertebrates. Species identification success rate was high (88.5%) and established the presence of 3 predators at elk mortality sites including black bears (55.7%), cougars (Puma concolor; 27.9%), and coyotes (Canis latrans; 6.6%). Attempts at hair and scat identification by field personnel were correct for 58% of hair samples and 79% of fecal samples. Results from these analyses demonstrate the merits of combining field mortality assessments with mtDNA species identification to aid wildlife managers in more accurately pinpointing predators involved in either predation or depredation events.