Scavengers generally rely on patchily distributed, unpredictable carrion. A long-standing hypothesis suggests scavenging ravens reliably locate such food by directly following large carnivores to their kills. However, by satellite tracking 69 ravens, 20 wolves, and 11 cougars in Yellowstone National Park, we found that following of predators over large distances rarely occurred. Instead, ravens routinely revisited sites where wolf kills were common-returning from distances of up to 155 kilometers to find carrion. Much like navigating to permanent anthropogenic subsidies, ravens appear to remember potential sources of carrion shaped by previous encounters with wolves or their kills. These findings suggest that spatial memory and navigation play a considerably greater role than previously assumed among scavengers, and possibly other wide-ranging species, in search of ephemeral resources.

How similar species overlap in space and resource use within communities is a long-standing question in ecology. For mobile species, how individuals move shapes their resource use and interactions with other animals. Yet links between individual movements and community resource use remain underdeveloped as resource overlap is primarily assessed across species' diets, and individual movements are generally assessed within single species. Here, we bridge movement and community ecology by evaluating how individual movements influence resource use across a community of sympatric, migratory ungulates. Using remotely sensed environmental data and GPS locations of 5 ungulates in Yellowstone National Park, US (2016-2021), we show that variation of individual movements drove differences in community resource use, supporting the classic niche variation hypothesis. Species with lower between-individual variation in their movements (bighorn sheep and pronghorn) overlapped less in their resource use with other species than species with higher between-individual variation (elk, mule deer, and bison). The extent of resource-use overlap across species was determined by the daily and seasonal movements and habitat use of individuals. Tracking of plant phenology and snow conditions by individuals drove the use of, or exposure to, differing qualities of resources in different locations across species. Despite variation across species, Yellowstone ungulates were overall highly similar in their resource use, suggesting that interspecies competition was not a prominent driver of their resource use. Our findings confirm a strong link between individual movements and community resource use, underscoring the importance of individual behaviors in shaping communities.

Given the importance of protected areas for biodiversity, the growth of visitation to many areas has raised concerns about the effects of humans on wildlife. In 2020, the COVID-19 pandemic led to temporary closure of national parks in the United States, offering a pseudonatural experiment to tease apart the effects of permanent infrastructure and transient human presence on animals. We compiled GPS tracking data from 229 individuals of 10 mammal species in 14 parks and used third-order hierarchical resource selection functions to evaluate the influence of the human footprint on animal space use in 2019 and 2020. Averaged across all parks and species, animals avoided the human footprint, whether the park was open or closed. However, although animals in remote areas showed consistent avoidance, on average those in more developed areas switched from avoidance to selection when protected areas were closed. Findings varied across species: some responded consistently negatively to the footprint (wolves, mountain goats), some positively (mule deer, red fox) and others had a strong exposure-mediated response (elk, mountain lion). Furthermore, some species responded more strongly to the park closure (black bear, moose). This study advances our understanding of complex interactions between recreation and wildlife in protected areas.

Behavioral plasticity, the alteration of behavior in response to stimuli, is becoming increasingly important in the context of human-induced rapid environmental change. Theoretical and empirical studies suggest that the expression and magnitude of behavioral plasticity are likely facilitated or constrained primarily by two factors: environmental variation and endogenous traits such as body size. The contextual role of these factors on behavioral plasticity, however, is poorly understood; there are relatively few studies that have compared the magnitude and potential drivers of behavioral plasticity at different levels (i.e., population and individual) across species, especially in free-ranging animals with diverse behavioral traits such as large mammals. Here, we quantify and test potential hypotheses for the mechanisms underpinning behavioral plasticity at the individual and population level in response to variation in summer temperatures for 1068 animal-years in 17 populations across nine species of large mammals. All populations displayed behavioral plasticity in response to increased temperatures, modifying their relative selection for heat-relieving habitat attributes (e.g., elevation) and heat-generating behavior (i.e., movement speed). We found strong support for the hypothesis that the variability of the physical environment is an important driver of behavioral plasticity-both mean population behavioral plasticity and variation among individuals within each population in plasticity were lower with increased heterogeneity of habitat attributes such as tree cover. Yet, the variability in environmental conditions (i.e., the magnitude of the temperature increase) had no effect on behavioral plasticity within and among populations. We did not detect an effect of endogenous traits on the expression of behavioral plasticity; however, we note that data availability limited our tests of this hypothesis to a select few endogenous traits (body size, feeding guild, and sex of the tracked individuals) that predominantly vary at the species level, for which we had one to three replicate populations per species. Our results provide an integrative and generalizable understanding of the expression of behavioral plasticity among populations of large mammals in temperate environments and emphasize the important but nuanced role of environmental variation in determining the scope of behavioral plasticity in these populations.

Direct competition for resources is especially fierce among predators, leading to disproportionately strong effects on fitness and functional roles. These competitive effects are exacerbated in complex predator guilds with dominance hierarchies that have clear winners and losers. The direct costs of losing these competitions are well understood, but the drivers of such interactions, and their indirect effects on prey, are not. We evaluate the drivers of interference competition for cougars, and how such competition affects cougar-prey dynamics, by leveraging 23 years of cougar predation data from Yellowstone National Park, USA. We show that the effect of prey size is context-dependent, negatively affecting how often cougars kill ungulate prey but positively affecting how often wolves/bears find and steal cougar kills. Further, cougars increasingly kill smaller prey as larger, primary prey density decreases. Handling time is shorter for smaller prey, leading to less kleptoparasitism by wolves and bears when primary prey density is lower. Our study counters the theory suggesting that interference competition should increase at kills when prey density declines, interspecific competitor density increases, or kill rates increase. We demonstrate that predator, competitor, and prey traits drive the strength of and even dampen interference competition, possibly increasing coexistence in complex communities.

The population dynamics of animals involve a complex interplay between intrinsic and extrinsic factors. Intrinsic regulation, characterized as population self-regulation, encompasses mechanisms that moderate growth rates before reaching maximum food-dependent densities. Conversely, extrinsically regulated populations are constrained by environmental variables such as food availability, predation, and disease. What regulates wolf populations has been debated for decades, with most publications concluding that wolves are extrinsically regulated, despite evidence of intrinsic mechanisms. This paper summarizes one perspective from a debate at the 'Wolves Across Borders' conference in Stockholm, Sweden in May 2023. Since this paper is a summary of a public debate, the paper is neither a review nor research paper, rather a research history and a recommendation on how the long-term disagreement might be resolved. Wolf population regulation is likely an interplay between both intrinsic and extrinsic factors which is best studied longitudinally by monitoring one population through time.

The mammalian mouth is colonized by complex microbial communities, adapted to specific niches, and in homeostasis with the host. Individual microbes interact metabolically and rely primarily on nutrients provided by the host, with which they have potentially co-evolved along the mammalian lineages. The oral environment is similar across mammals, but the diversity, specificity, and evolution of community structure in related or interacting mammals are little understood. Here, we compared the oral microbiomes of dogs with those of wild wolves and humans. In dogs, we found an increased microbial diversity relative to wolves, possibly related to the transition to omnivorous nutrition following domestication. This includes a larger diversity of Patescibacteria than previously reported in any other oral microbiota. The oral microbes are most distinct at bacterial species or strain levels, with few if any shared between humans and canids, while the close evolutionary relationship between wolves and dogs is reflected by numerous shared taxa. More taxa are shared at higher taxonomic levels including with humans, supporting their more ancestral common mammalian colonization followed by diversification. Phylogenies of selected oral bacterial lineages do not support stable human-dog microbial transfers but suggest diversification along mammalian lineages (apes and canids). Therefore, despite millennia of cohabitation and close interaction, the host and its native community controls and limits the assimilation of new microbes, even if closely related. Higher resolution metagenomic and microbial physiological studies, covering a larger mammalian diversity, should help understand how oral communities assemble, adapt, and interact with their hosts.IMPORTANCENumerous types of microbes colonize the mouth after birth and play important roles in maintaining oral health. When the microbiota-host homeostasis is perturbed, proliferation of some bacteria leads to diseases such as caries and periodontitis. Unlike the gut microbiome, the diversity of oral microbes across the mammalian evolutionary space is not understood. Our study compared the oral microbiomes of wild wolves, dogs, and apes (humans, chimpanzees, and bonobos), with the aim of identifying if microbes have been potentially exchanged between humans and dogs as a result of domestication and cohabitation. We found little if any evidence for such exchanges. The significance of our research is in finding that the oral microbiota and/or the host limit the acquisition of exogenous microbes, which is important in the context of natural exclusion of potential novel pathogens. We provide a framework for expanded higher-resolution studies across domestic and wild animals to understand resistance/resilience.

Large carnivores are globally threatened due to habitat fragmentation and loss, prey depletion and human exploitation. Human exploitation includes both legal and illegal hunting and trapping. Protected areas can create refugia from hunting and trapping; however, hunting can still threaten wide-ranging large carnivores when they leave these areas. Large carnivore reintroductions to protected areas are often motivated to restore ecological processes, including wolf reintroduction to Yellowstone National Park (YNP). Determining if harvest is compensatory or additive is essential for informed conservation strategies, as it influences the overall impact on wolf populations and their ecosystems. If harvest was compensatory, then increasing harvest pressure outside YNP should not decrease overall survival for transboundary wolves. Alternatively, if increasing harvest was additive, then increasing harvest pressure outside YNP should decrease overall survival for transboundary wolves. We tested the effects of variable harvest pressure following delisting on the survival of YNP grey wolves (Canis lupus) from 1995 to 2022. We defined three harvest levels: no harvest, harvest with limited quotas and unlimited harvest. We used Cox-proportional hazards models and cumulative incidence functions to estimate survival rates, factors affecting survival and cause-specific mortality between these three harvest periods to test predictions of the additive mortality hypothesis. Most harvested wolves that primarily lived in YNP were killed adjacent to the park border. Cox-proportional hazards models revealed that mortality was highest during years of unlimited harvest during winter outside YNP. Cause-specific mortality analyses showed that natural mortality from other wolves and harvest were the two leading causes of death, but that harvest mortality had additive effects on wolf mortality. Wolf survival decreased with increased harvest mortality, while natural mortality remained relatively unchanged. Synthesis and applications. High rates of additive harvest mortality of wolves could negatively impact wolf survival in YNP. Harvest mortality of transboundary wolves is additive possibly due to source-sink dynamics of uneven spatial susceptibility of wolves to harvest mortality across protected area borders, as well as effects of harvest on complex social dynamics of wolves in YNP. Transboundary management of large carnivores is challenging, yet cooperation between agencies is vital for wolf management in and around YNP. Our results support the use of small quota zones surrounding protected areas, that minimize transboundary mortality impacts on large carnivores living primarily inside protected areas.

A central debate in ecology has been the long-running discussion on the role of apex predators in affecting the abundance and dynamics of their prey. In terrestrial systems, research has primarily relied on correlational approaches, due to the challenge of implementing robust experiments with replication and appropriate controls. A consequence of this is that we largely suffer from a lack of mechanistic understanding of the population dynamics of interacting species, which can be surprisingly complex. Mechanistic models offer an opportunity to examine the causes and consequences of some of this complexity. We present a bioenergetic mechanistic model of a tritrophic system where the primary vegetation resource follows a seasonal growth function, and the herbivore and carnivore species are modeled using two integral projection models (IPMs) with body mass as the phenotypic trait. Within each IPM, the demographic functions are structured according to bioenergetic principles, describing how animals acquire and transform resources into body mass, energy reserves, and breeding potential. We parameterize this model to reproduce the population dynamics of grass, elk, and wolves in northern Yellowstone National Park (USA) and investigate the impact of wolf reintroduction on the system. Our model generated predictions that closely matched the observed population sizes of elk and wolf in Yellowstone prior to and following wolf reintroduction. The introduction of wolves into our basal grass-elk bioenergetic model resulted in a population of 99 wolves and a reduction in elk numbers by 61% (from 14,948 to 5823) at equilibrium. In turn, vegetation biomass increased by approximately 25% in the growing season and more than threefold in the nongrowing season. The addition of wolves to the model caused the elk population to switch from being food-limited to being predator-limited and had a stabilizing effect on elk numbers across different years. Wolf predation also led to a shift in the phenotypic composition of the elk population via a small increase in elk average body mass. Our model represents a novel approach to the study of predator-prey interactions, and demonstrates that explicitly considering and linking bioenergetics, population demography and body mass phenotypes can provide novel insights into the mechanisms behind complex ecosystem processes.