Recreation and hunting differentially affect deer behaviour and sapling performance.

Humans are increasingly acknowledged as apex predators that shape landscapes of fear to which herbivores adapt their behaviour. Here, we investigate how humans modify deer space-use and their effects on vegetation at two spatial scales; zones with different types of human use (large-scale risk factor) and, nested within that, trails (fine-scale risk factor). In zones with three contrasting types of human activities: (1) no recreation, no hunting, (2) with recreation, no hunting, and (3) with recreation and hunting, we linked deer space-use (dropping counts) to browsing intensity, relative growth, and survival of planted saplings. Plots were located at two distances to trails (20 vs. 100m) to test how trails affect deer space-use and sapling performance. Additionally, plots were distributed over forest and heathland as risk effects are habitat-dependent. Deer space-use was highest in the zone without recreation or hunting, resulting in higher browsing levels and lower sapling growth and survival, but only in heathland. In contrast, deer space-use and sapling performance did not differ between zones with recreation only and zones with recreation and hunting. Deer dropping counts were lower near trails used for recreation, but this was not associated with browsing impact or sapling performance. Our results show that recreational use modifies deer space-use which is associated with browsing impact on woody vegetation, while seasonal hunting activities in zones with recreation did not have additive year-round effects. Yet, effects were only observed at the larger scale of recreation zones and not near trails. Furthermore, deer space-use was only associated with sapling performance in open heathland, where high visibility presumably increases avoidance behaviour because it increases detectability and decreases escape possibilities. This suggests that recreation creates behaviourally mediated cascading effects that influence vegetation development, yet these effects are context-dependent. We advocate incorporating human-induced fear effects in conservation, management and research.