Cougars, wolves, and humans drive a dynamic landscape of fear for elk.

ABSTRACT

To manage predation risk, prey navigate a dynamic landscape of fear, or spatiotemporal variation in risk perception, reflecting predator distributions, traits, and activity cycles. Prey may seek to reduce risk across this landscape using habitat at times and in places when predators are less active. In multipredator landscapes, avoiding one predator could increase vulnerability to another, making the landscape of fear difficult to predict and navigate. Additionally, humans may shape interactions between predators and prey, and induce new sources of risk. Humans can function as a shield, providing a refuge for prey from human-averse carnivores, and as a predator, causing mortality through hunting and vehicle collisions and eliciting a fear response that can exceed that of carnivores. We used telemetry data collected between 2017 and 2021 from 63 Global Positioning System-collared elk (Cervus canadensis), 42 cougars (Puma concolor), and 16 wolves (Canis lupus) to examine how elk habitat selection changed in relation to carnivores and humans in northeastern Washington, USA. Using step selection functions, we evaluated elk habitat use in relation to cougars, wolves, and humans, diel period (daytime vs. nighttime), season (summer calving season vs. fall hunting season), and habitat structure (open vs. closed habitat). The diel cycle was critical to understanding elk movement, allowing elk to reduce encounters with predators where and when they would be the largest threat. Elk strongly avoided cougars at night but had a near-neutral response to cougars during the day, whereas elk avoided wolves at all times of day. Elk generally used more open habitats where cougars and wolves were most active, rather than altering the use of habitat structure depending on the predator species. Elk avoided humans during the day and ~80% of adult female mortality was human caused, suggesting that humans functioned as a "super predator" in this system. Simultaneously, elk leveraged the human shield against wolves but not cougars at night, and no elk were confirmed to have been killed by wolves. Our results add to the mounting evidence that humans profoundly affect predator-prey interactions, highlighting the importance of studying these dynamics in anthropogenic areas.