Variable strategies to solve risk-reward tradeoffs in carnivore communities.

Mesopredator release theory suggests that dominant predators suppress subordinate carnivores and ultimately shape community dynamics, but the assumption that subordinate species are only negatively affected ignores the possibility of facilitation through scavenging. We examined the interplay within a carnivore community consisting of cougars, coyotes, black bears, and bobcats using contemporaneous Global Positioning System telemetry data from 51 individuals; diet analysis from 972 DNA-metabarcoded scats; and data from 128 physical investigations of cougar kill sites, 28 of which were monitored with remote cameras. Resource provisioning from competitively dominant cougars to coyotes through scavenging was so prolific as to be an overwhelming determinant of coyote behavior, space use, and resource acquisition. This was evident via the strong attraction of coyotes to cougar kill sites, frequent scavenging of cougar-killed prey, and coyote diets that nearly matched cougars in the magnitude of ungulate consumption. Yet coyotes were often killed by cougars and used space to minimize encounters, complicating the fitness benefits gained from scavenging. We estimated that 23% (95% CI: 8 to 55%) of the coyote population in our study area was killed by cougars annually, suggesting that coyote interactions with cougars are a complex behavioral game of risk and reward. In contrast, we found no indication that bobcat space use or diet was influenced by cougars. Black bears avoided cougars, but there was no evidence of attraction to cougar kill sites and much lower levels of ungulate consumption and carcass visitation than for coyotes. Interspecific interactions among carnivores are multifaceted, encompassing both suppression and facilitation.

Habitat predicts local prevalence of migratory behaviour in an alpine ungulate.

The resource hierarchy hypothesis predicts that the most important factors limiting a species' distribution act at the coarsest spatial scales. However, resource selection behaviour affords mobile organisms the opportunity to adopt a range of tactics for navigating spatial trade-offs between competing biotic and abiotic constraints. Throughout the animal kingdom, partial migration (where some individuals migrate, and others remain resident year round) offers a pervasive example of such behavioural polymorphism. Identifying the differences between these behaviours is therefore central to understanding the conditions (habitat) needed to sustain migrant and resident populations. Here we test an extension of the resource hierarchy hypothesis. We hypothesized that rather than responding to a single limiting factor, migration and residency represent contrasting scale-specific approaches to managing trade-offs between forage, predation risk and severe winter conditions. Furthermore, we predicted that the distribution of habitat selected by migrants and residents is predictive of the local prevalence of migratory behaviour. To test these hypotheses, we quantified migratory status- (resident/migrant) and season-specific (winter/summer) differences in resource selection by eight populations of federally endangered Sierra Nevada bighorn sheep Ovis canadensis sierrae across three spatial scales: population range, individual range and within individual range. We then integrated these spatial predictions to produce separate spatial predictions of migrant and resident winter habitat. As predicted, model selection provided strong evidence for the importance of status-specific differences in resource selection. Residents showed stronger coarse-scale selection for terrain associated with predator avoidance and stronger fine-scale selection for greenness, while in migrants this pattern was reversed. Availability of migrant habitat predicted the local prevalence of migration (top model pseudo R2 of .87). Our ability to respond to global declines of migratory species depends on improving our understanding of the conditions required to maintain migratory behaviour. Through explicitly contrasting migrant and resident behaviour, our results illustrate seasonal differences in migrant and resident habitat and how these two behaviours represent responses to different limiting conditions. Our analyses provides a novel empirical basis for assessing the local prevalence of migratory behaviour across large landscapes.

Fire history influences large-herbivore behavior at circadian, seasonal, and successional scales.

Recurrent environmental changes often prompt animals to alter their behavior leading to predictable patterns across a range of temporal scales. The nested nature of circadian and seasonal behavior complicates tests for effects of rarer disturbance events like fire. Fire can dramatically alter plant community structure, with important knock-on effects at higher trophic levels, but the strength and timing of fire's effects on herbivores remain unclear. We combined prescribed fire treatments with fine-scale location data to quantify herbivore responses to fire across three temporal scales. Between 2001 and 2003, 26 stands of fir (Abies spp.) and Douglas-fir (Pseudotsuga menziesii) were thinned and burned; 27 similar stands were left untreated as experimental controls. Analyzing female elk (Cervus canadensis) locations across 21 yr (1996-2016), we found crepuscular, seasonal, and successional shifts in behavioral responses to fire. Elk displayed "commuting" behavior, avoiding burns during the day, but selecting them at night. Elk selection for burns was strongest in early summer and the relative probability of elk using burns peaked quickly (5 yr post burn) before gradually returning to pre-treatment levels (15 yr post burn). Our results demonstrate that fire history has complex, persistent effects on herbivore behavior, and suggest that herbivores benefit from heterogeneous landscapes containing a range of successional stages.

Standardizing protocols for determining the cause of mortality in wildlife studies.

Mortality site investigations of telemetered wildlife are important for cause-specific survival analyses and understanding underlying causes of observed population dynamics. Yet, eroding ecoliteracy and a lack of quality control in data collection can lead researchers to make incorrect conclusions, which may negatively impact management decisions for wildlife populations. We reviewed a random sample of 50 peer-reviewed studies published between 2000 and 2019 on survival and cause-specific mortality of ungulates monitored with telemetry devices. This concise review revealed extensive variation in reporting of field procedures, with many studies omitting critical information for the cause of mortality inference. Field protocols used to investigate mortality sites and ascertain the cause of mortality are often minimally described and frequently fail to address how investigators dealt with uncertainty. We outline a step-by-step procedure for mortality site investigations of telemetered ungulates, including evidence that should be documented in the field. Specifically, we highlight data that can be useful to differentiate predation from scavenging and more conclusively identify the predator species that killed the ungulate. We also outline how uncertainty in identifying the cause of mortality could be acknowledged and reported. We demonstrate the importance of rigorous protocols and prompt site investigations using data from our 5-year study on survival and cause-specific mortality of telemetered mule deer (Odocoileus hemionus) in northern California. Over the course of our study, we visited mortality sites of neonates (n = 91) and adults (n = 23) to ascertain the cause of mortality. Rapid site visitations significantly improved the successful identification of the cause of mortality and confidence levels for neonates. We discuss the need for rigorous and standardized protocols that include measures of confidence for mortality site investigations. We invite reviewers and journal editors to encourage authors to provide supportive information associated with the identification of causes of mortality, including uncertainty.