Behavioral responses of prey to predation risk have ecological impacts that can be as great as the direct mortality. Theory often suggests that prey pair responses to risks based on the hunting mode of the predator (hunting mode hypothesis), but prey may ignore hunting mode to prioritize responding to the most lethal predators (lethality hypothesis). To test these hypotheses, we evaluated the behavioral responses of white-tailed deer (Odocoileus virginianus) to risks from two natural mesopredators and human sources of mortality. Specifically, we determined, for each source of risk, whether deer responded with behavioral state changes or spatial avoidance and whether this behavior changed with time (diurnally and annually). We collared and tracked 40 female and 29 male deer. To determine the response of deer to risk, we collected data on the distribution of coyotes (Canis latrans), bobcats (Lynx rufus), human modification, hunters, and roads. We used hidden Markov models (HMM) to determine whether each covariate impacted the probability of transitioning between behavioral states and selection functions (SSF) to determine whether deer spatially avoided each covariate.Generally, deer changed behavioral state in response to both mesopredators but avoided human modification. In response to mesopredators, deer consistently shifted to slower movement behavioral states. Spatial responses to human modification varied depending on the time of day. During daylight hours, deer selected for human modification, but during the crepuscular and nighttime period, deer avoided human modification. Space use was most strongly related to more lethal humans, providing support for the lethality hypothesis. Despite prioritizing humans, mesopredators impacted behavioral state, suggesting that mesopredators still have important impacts on prey behavior. Finally, temporal patterns of avoidance align with other studies that indicate avoidance of predators is time-dependent, but further highlight the complex push-pull relationship of human modified areas on wildlife.

Spatial behavior, including home-ranging behaviors, habitat selection, and movement, can be extremely informative in estimating how animals respond to landscape heterogeneity. Responses in these spatial behaviors to factors such as human modification and resources on the landscape can highlight a species' spatial strategy to maximize fitness and minimize risk. These strategies can vary on spatial, temporal, and individual scales, and the combination of behaviors on these scales can lead to very different strategies among species. Harnessing the variation present at these scales, we developed a framework for predicting how species may respond to changes in their environments on a gradient ranging from generic, where a species exhibits broad-stroke spatial responses to their environment, to nuanced, in which a species uses a combination of temporal and spatial strategies paired with functional responses in selection behaviors. Using 46 GPS-tracked bobcats and coyotes inhabiting a landscape encompassing a range of human modification, we evaluated where each species falls along the generic-to-nuanced gradient. Bobcats and coyotes studied occupied opposite ends of this gradient, using different strategies in response to human modification in their home ranges, with bobcats broadly expanding their home range with increases in human modification and clearly selecting for or avoiding features on the landscape with temporal consistency. Meanwhile, coyotes did not expand their home ranges with human modification, but instead displayed temporal and spatial adjustments in their functional responses to human modification. These differences in response to habitat, resources, and risk between the two species highlighted the variation in spatial behaviors animals can use to exist in anthropogenic environments influenced by interspecific variation in behavioral plasticity. Categorizing animal spatial behavior based on the generic-to-nuanced gradient can help in predicting how a species will respond to future change based on their current spatial behavior.