Death comes for us all: relating movement-integrated habitat selection and social behavior to human-associated and disease-related mortality among gray wolves.

Avoiding death affects biological processes, including behavior. Habitat selection, movement, and sociality are highly flexible behaviors that influence the mortality risks and subsequent fitness of individuals. In the Anthropocene, animals are experiencing increased risks from direct human causes and increased spread of infectious diseases. Using integrated step selection analysis, we tested how the habitat selection, movement, and social behaviors of gray wolves vary in the two months prior to death due to humans (being shot or trapped) or canine distemper virus (CDV). We further tested how those behaviors vary as a prelude to death. We studied populations of wolves that occurred under two different management schemes: a national park managed for conservation and a provincially managed multi-use area. Behaviors that changed prior to death were strongly related to how an animal eventually died. Wolves killed by humans moved slower than wolves that survived and selected to be nearer roads closer in time to their death. Wolves that died due to CDV moved progressively slower as they neared death and reduced their avoidance of wet habitats. All animals, regardless of dying or living, maintained selection to be near packmates across time, which seemingly contributed to disease dynamics in the packs infected with CDV. There were no noticeable differences in behavior between the two management areas. Overall, habitat selection, movement, and sociality interact to put individuals and groups at greater risks, influencing their cause-specific mortality.

Wolf spatial behavior promotes encounters and kills of abundant prey.

Predators use different spatial tactics to track the prey on the landscape. Three hypotheses describe spatial tactics: prey abundance for prey that are aggregated in space; prey habitat for uniformly distributed prey; and prey catchability for prey that are difficult to catch and kill. The gray wolf (Canis lupus) is a generalist predator that likely employs more than one spatial hunting tactic to match their diverse prey with distinct distributions and behavior that are available. We conducted a study on 17 GPS collared wolves in 6 packs in Riding Mountain National Park, Manitoba, Canada where wolves prey on moose (Alces alces) and elk (Cervus canadensis). We evaluated wolf selection for prey density, habitat selection and catchability on the landscape through within-territory habitat selection analysis. We reveal support for both the prey habitat and prey catchability hypotheses. For moose, their primary prey, wolves employed a mixed habitat and catchability tactic. Wolves used spaces described by the intersection of moose habitat and moose catchability. Wolves selected for the catchability of elk, their secondary prey, but not elk habitat. Counter to our predictions, wolves avoided areas of moose and elk density, likely highlighting the ongoing space race between predator and prey. We illustrate that of the three hypotheses the primary driver was prey catchability, where the interplay of both prey habitat with catchability culminate in predator spatial behaviour in a multiprey system.

The Ecology of Individual Differences Empirically Applied to Space-Use and Movement Tactics.

Movement provides a link between individual behavioral ecology and the spatial and temporal variation in an individual's landscape. Individual variation in movement traits is an important axis of animal personality, particularly in the context of foraging ecology. We tested whether individual caribou (Rangifer tarandus) displayed plasticity in movement and space-use behavior across a gradient of resource aggregation. We quantified first-passage time and range-use ratio as proxies for movement-related foraging behavior and examined how these traits varied at the individual level across a foraging resource gradient. Our results suggest that individuals adjusted first-passage time but not range-use ratio to maximize access to high-quality foraging resources. First-passage time was repeatable, and intercepts for first-passage time and range-use ratio were negatively correlated. Individuals matched first-passage time but not range-use ratio to the expectations of our patch-use model that maximized access to foraging resources, a result that suggests that individuals acclimated their movement patterns to accommodate both intra- and interannual variation in foraging resources on the landscape. Collectively, we highlight repeatable movement and space-use tactics and provide insight into how individual plasticity in movement interacts with landscape processes to affect the distribution of behavioral phenotypes and potentially fitness and population dynamics.

Friends because of foes: synchronous movement within predator-prey domains.

For prey, movement synchrony represents a potent antipredator strategy. Prey, however, must balance the costs and benefits of using conspecifics to mediate risk. Thus, the emergent patterns of risk-driven sociality depend on variation in space and in the predators and prey themselves. We applied the concept of predator-prey habitat domain, the space in which animals acquire food resources, to test the conditions under which individuals synchronize their movements relative to predator and prey habitat domains. We tested the response of movement synchrony of prey to predator-prey domains in two populations of ungulates that vary in their gregariousness and predator community: (i) elk, which are preyed on by wolves; and (ii) caribou, which are preyed on by coyotes and black bears. Prey in both communities responded to cursorial predators by increasing synchrony during seasons of greater predation pressure. Elk moved more synchronously in the wolf habitat domain during winter and caribou moved more synchronously in the coyote habitat domains during spring. In the winter, caribou increased movement synchrony when coyote and caribou domains overlapped. By integrating habitat domains with movement ecology, we provide a compelling argument for social behaviours and collective movement as an antipredator response. This article is part of the theme issue 'The spatial-social interface: A theoretical and empirical integration'.