Reintroduced wolves and hunting limit the abundance of a subordinate apex predator in a multi-use landscape

Top-down effects exhibited by apex predators are modulated by human impacts on community composition and species abundances. For these reasons, research supporting strong top-down effects of apex predators occurs almost entirely within protected areas rather than the much more common multi-use landscapes dominating modern ecosystems. Based on 16 years of monitoring, we developed an integrated population model to disentangle the concurrent contributions of a reintroduced apex predator, the gray wolf, human hunting, and prey abundances on vital rates and abundance of a subordinate apex predator, the puma. Increasing wolf numbers had strong negative effects on puma fecundity, and subadult and adult survival. Puma survival was also influenced by density-dependence. Overall, our results showed that puma dynamics in our multi-use landscape were more strongly influenced by top-down forces exhibited by a reintroduced apex predator, than by human hunting or bottom-up forces (prey abundance) subs...