Separating spatial search and efficiency rates as components of predation risk.

Predation risk is an important driver of ecosystems, and local spatial variation in risk can have population-level consequences by affecting multiple components of the predation process. I use resource selection and proportional hazard time-to-event modelling to assess the spatial drivers of two key components of risk--the search rate (i.e. aggregative response) and predation efficiency rate (i.e. functional response)--imposed by wolves (Canis lupus) in a multi-prey system. In my study area, both components of risk increased according to topographic variation, but anthropogenic features affected only the search rate. Predicted models of the cumulative hazard, or risk of a kill, underlying wolf search paths validated well with broad-scale variation in kill rates, suggesting that spatial hazard models provide a means of scaling up from local heterogeneity in predation risk to population-level dynamics in predator-prey systems. Additionally, I estimated an integrated model of relative spatial predation risk as the product of the search and efficiency rates, combining the distinct contributions of spatial heterogeneity to each component of risk.