Human avoidance, selection for darkness and prey activity explain wolf diel activity in a highly cultivated landscape

Wildlife that share habitats with humans with limited options for spatial avoidance must either tolerate frequent human encounters or concentrate their activity into those periods with the least risk of encountering people. Based on 5259 camera trap images of adult wolves from eight territories, we analyzed the extent to which diel activity patterns in a highly cultivated landscape with extensive public access (Denmark) could be explained from diel variation in darkness, human activity, and prey (deer) activity. A resource selection function that contrasted every camera observation (use) with 24 alternative hourly observations from the same day (availability), revealed that diel activity correlated with all three factors simultaneously with human activity having the strongest effect (negative), followed by darkness (positive) and deer activity (positive). A model incorporating these three effects had lower parsimony and classified use and availability observations just as well as a ‘circadian' model that smoothed the use‐availability ratio as function of time of the day. Most of the selection for darkness was explained by variation in human activity, supporting the notion that nocturnality (proportion of observations registered at night vs. day at the equinox) is a proxy for temporal human avoidance. Contrary to our expectations, wolves were no more nocturnal in territories with unrestricted public access than in territories where public access was restricted to roads, possibly because wolves in all territories had few possibilities to walk more than few hundred meters without crossing roads. Overall, Danish wolf packs were 6.5 (95% CI: 4.6–9.6) times more active at night than at daylight, which make them amongst the most nocturnally active wolves reported so far. These results confirm the prediction that wolves in habitats with limited options for spatial human avoidance, invest more in temporal avoidance.