Consequences of ratio‐dependent predation by wolves for elk population dynamics

Abstract A growing number of studies suggest ratio‐dependence may be common in many predator–prey systems, yet in large mammal systems, evidence is limited to wolves and their prey in Isle Royale and Yellowstone. More importantly, the consequences of ratio‐dependent predation have not been empirically examined to understand the implications for prey. Wolves recolonized Banff National Park in the early 1980s, and recovery was correlated with significant elk declines. I used time‐series data of wolf kill rates of elk, wolf and elk densities in winter from 1985–2007 to test for support for prey‐, ratio‐, or predator dependent functional and numeric responses of wolf killing rate to elk density. I then combined functional and numeric responses to estimate the total predation response to identify potential equilibrium states. Evidence suggests wolf predation on elk was best described by a type II ratio‐dependent functional response and a type II numeric response that lead to inversely density‐dependent predation rate on elk. Despite support for ratio‐dependence, like other wolf‐prey systems, there was considerable uncertainty amongst functional response models, especially at low prey densities. Consistent with predictions from ratio‐dependent models, however, wolves contributed to elk population declines of over 80 % in our Banff system. Despite the statistical signature for ratio‐dependence, the biological mechanism remains unknown and may be related to multi‐prey dynamics in our system. Regardless, ratio‐dependent models strike a parsimonious balance between theory and empiricism, and this study suggests that large mammal ecologists need to consider ratio‐dependent models in predator–prey dynamics.