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

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.