Contrasting patterns of risk from human and non-human predators shape temporal activity of prey.

Spatiotemporal variation in predation risk arises from interactions between landscape heterogeneity, predator densities and predator hunting mode, generating landscapes of fear for prey species that can have important effects on prey behaviour and ecosystem dynamics. As widespread apex predators, humans present a significant source of risk for hunted animal populations. Spatiotemporal patterns of risk from hunters can overlap or contrast with patterns of risk from other predators. Human infrastructure can also reshape spatial patterns of risk by facilitating or impeding hunter or predator movement, or deterring predators that are themselves wary of humans. We examined how anthropogenic and natural landscape features interact with hunting modes of rifle hunters and mountain lions Puma concolor to generate spatiotemporal patterns of risk for their primary prey. We explored the implications of human-modified landscapes of fear for Columbian black-tailed deer Odocoileus hemionus columbianus in Mendocino County, California. We used historical harvest records, hunter GPS trackers and camera trap records of mountain lions to model patterns of risk for deer. We then used camera traps to examine deer spatial and temporal activity patterns in response to this variation in risk. Hunters and mountain lions exhibited distinct, contrasting patterns of spatiotemporal activity. Risk from rifle hunters, who rely on long lines of sight, was highest in open grasslands and near roads and was confined to the daytime. Risk from mountain lions, an ambush predator, was highest in dense shrubland habitat, farther from developed areas, and during the night and crepuscular periods. Areas of human settlement provided a refuge from both hunters and mountain lions. We found no evidence that deer avoided risk in space at the scale of our observations, but deer adjusted their temporal activity patterns to reduce the risk of encounters with humans and mountain lions in areas of higher risk. Our study demonstrates that interactions between human infrastructure, habitat cover and predator hunting mode can result in distinct spatial patterns of predation risk from hunters and other predators that may lead to trade-offs for prey species. However, distinct diel activity patterns of predators may create vacant hunting domains that reduce costly trade-offs for prey. Our study highlights the importance of temporal partitioning as a mechanism of predation risk avoidance.