Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates.

Recolonizing carnivores can have a large impact on the status of wild ungulates, which have often modified their behavior in the absence of predation. Therefore, understanding the dynamics of reestablished predator-prey systems is crucial to predict their potential ecosystem effects. We decomposed the spatial structure of predation by recolonizing wolves (Canis lupus) on two sympatric ungulates, moose (Alces alces) and roe deer (Capreolus capreolus), in Scandinavia during a 10-year study. We monitored 18 wolves with GPS collars, distributed over 12 territories, and collected records from predation events. By using conditional logistic regression, we assessed the contributions of three main factors, the utilization patterns of each wolf territory, the spatial distribution of both prey species, and fine-scale landscape structure, in determining the spatial structure of moose and roe deer predation risk. The reestablished predator-prey system showed a remarkable spatial variation in kill occurrence at the intra-territorial level, with kill probabilities varying by several orders of magnitude inside the same territory. Variation in predation risk was evident also when a spatially homogeneous probability for a wolf to encounter a prey was simulated. Even inside the same territory, with the same landscape structure, and when exposed to predation by the same wolves, the two prey species experienced an opposite spatial distribution of predation risk. In particular, increased predation risk for moose was associated with open areas, especially clearcuts and young forest stands, whereas risk was lowered for roe deer in the same habitat types. Thus, fine-scale landscape structure can generate contrasting predation risk patterns in sympatric ungulates, so that they can experience large differences in the spatial distribution of risk and refuge areas when exposed to predation by a recolonizing predator. Territories with an earlier recolonization were not associated with a lower hunting success for wolves. Such constant efficiency in wolf predation during the recolonization process is in line with previous findings about the naive nature of Scandinavian moose to wolf predation. This, together with the human-dominated nature of the Scandinavian ecosystem, seems to limit the possibility for wolves to have large ecosystem effects and to establish a behaviorally mediated trophic cascade in Scandinavia.