Hunting and Outdoor Recreation Affect Large Herbivore Activity Patterns More Than Natural Predators in a Human-Dominated Landscape.

Across Europe, landscapes where large carnivores, large herbivores and human communities coexist are expanding, reflecting the widespread recovery of large mammal populations in recent decades. The influence of top-down effects of wolves on large herbivores has been extensively studied in areas with relatively little anthropogenic disturbance, but less is known about their effect in human-dominated landscapes. We systematically collected camera-trap data over five consecutive autumn hunting seasons in an area of the eastern Alps which is intensely frequented by tourists and trekkers, and partially open to ungulate hunting. We used a quasi-experimental design, with half of the sampling sites located within nonhunting areas and half outside. Applying generalised additive mixed models (GAMMs) with cyclic cubic splines we investigated the effect of wolf, as well as lethal (hunting) and nonlethal (recreational) human activities on red deer spatiotemporal activity pattern. Similarly, we analysed the effect of recreational activities and red deer site-use on the spatiotemporal activity pattern of wolves. Hunting was associated with overall lower red deer activity, as well as reduced dawn-dusk peaks and diurnality. Crucially, hunting interacted with outdoor recreation exacerbating its impact, with major changes to red deer activity curve. Wolf site-use did not have a significant effect on the shape of red deer temporal curve. Wolves were markedly more active in areas highly used by red deer, and remained strongly nocturnal even where human activity was scarce. Our results show that humans, through both lethal and nonlethal activities, elicit stronger responses in red deer than their natural predator. Behavioural constraints imposed by humans on red deer, coupled with the cursorial predatory strategy of wolves, likely limit the possibility of wolf avoidance by red deer. In human-dominated European landscapes, human disturbance can therefore override natural predator-prey dynamics, reshaping behavioural landscapes and potentially increasing predator and prey spatiotemporal co-occurrence.