Social, behavioural and temporal components of wolf (Canis lupus) responses to anthropogenic landscape features in the central Apennines, Italy

Understanding large carnivores’ behavioural adaptations to habitat modifications is critical for their persistence in human-modified environments. Based on 10 Global Positioning System-collared wolves in a protected area of central Italy, we investigated wolf responses to anthropogenic features such as roads and settlements by using Step and Resource Selection Functions. We revealed that responses by wolves to anthropogenic features varied according to behavioural state (travelling vs. resting) and social affiliation (pack members vs. floaters), while accounting for seasonal and circadian effects. During summer, pack members strongly avoided roads and settlements throughout all day, both when travelling and resting, and complemented this response by selecting forested areas and shrublands. Conversely, during fall and winter, pack members relaxed the avoidance towards anthropogenic features by travelling closer to main roads and settlements, but they still selected resting sites farther from anthropogenic features and located them in denser cover and along steeper slopes. Compared to pack members, floaters, when travelling, showed a weaker avoidance of main roads and settlements and did not show any selection pattern towards environmental variables. When resting, contrary to pack members, floaters selected sites closer to main roads and settlements, even though these were still located along denser cover and steeper slopes. Our findings suggest that wolves living in human-modified landscapes adapt behaviourally to the spatial and temporal distribution of perceived interference by humans and that their response is complementary to the expected seasonal and circadian variation in human activity. These adaptations appear to be aided by the selection of habitat features that enhance security and allow segregation from humans. Maintenance of such habitat characteristics appears of critical importance to ensure the functionality of behavioural adaptations by wolves living in human-modified landscapes.