Abstract
Large herbivores influence ecosystem functioning via their effects on vegetation at different spatial scales. It is often overlooked that the spatial distribution of large herbivores results from their responses to interacting top-down and bottom-up ecological gradients that create landscape-scale variation in the structure of the entire community. We studied the complexity of these cascading interactions using high-resolution camera trapping and remote sensing data in the best-preserved European lowland forest, Białowieża Forest, Poland. We showed that the variation in spatial distribution of an entire community of large herbivores is explained by species-specific responses to both environmental bottom-up and biotic top-down factors in combination with human-induced (cascading) effects. We decomposed the spatial variation in herbivore community structure and identified functionally distinct landscape-scale herbivory regimes ('herbiscapes'), which are predicted to occur in a variety of ecosystems and could be an important mechanism creating spatial variation in herbivory maintaining vegetation heterogeneity.