Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest, Poland

Large carnivores can either directly influence ungulate populations or indirectly affect their behaviour. Knowledge from European systems, in contrast to North American systems, on how this might lead to cascading effects on lower trophic levels is virtually absent. We studied whether wolves Canis lupus via density‐mediated and behaviorally‐mediated effects on their ungulate prey species influence patterns of browsing and tree regeneration inside the Białowieża National Park, Poland. Browsing intensity of tree saplings (height class < 150 cm), irrespective of tree species or forest type, was lower inside a wolf core area (50.5%) where predator presence is highest, than in the remainder of the wolf pack’s home range (58.3%). Additionally, browsing intensity was reduced when the amount of coarse woody debris (CWD), which can act as a ʻungulate escape impedimentʼ, increased (within 5‐m radius) inside the wolf core area. No relationship existed outside the core area. As a result, the proportion of trees growing out of herbivore control increased more strongly with increasing amount of CWD inside compared to outside the wolf core area. This suggests that next to direct effects of wolves on ungulate density caused by a higher predation pressure inside the core area, risk effects are important and are enhanced by habitat characteristics. These results indicate that behaviorally‐mediated effects of predators on prey can become more important than density‐mediated effects in affecting lower trophic levels. This is the first study we are aware of, that shows CWD can create fine‐scale risk effects on ungulates with the potential for cascading effects of large predators on patterns of tree regeneration for a European forest system. This knowledge broadens the discussion on how the impact of large predators on ecosystem functioning depends on the physical landscape, by illustrating these effects for a system which largely contrasts in this respect to the North American systems.