RESEARCH ARTICLE Coping with Spatial Heterogeneity and Temporal Variability in Resources and Risks: Adaptive Movement Behaviour by a Large Grazing Herbivore

Movement is a key mean for mobile species to cope with heterogeneous environments. While in herbivorous mammals large-scale migration has been widely investigated, fine-scale movement responses to local variations in resources and predation risk remain much less studied, especially in savannah environments. We developed a novel approach based on complementary movement metrics (residence time, frequency of visits and regularity of visits) to relate movement patterns of a savannah grazer, the blue wildebeest Conno-chaetes taurinus, to fine-scale variations in food availability, predation risk and water avail-ability in the Kruger National Park, South Africa. Wildebeests spent more time in grazing lawns where the grass is of higher quality but shorter than in seep zones, where the grass is of lower quality but more abundant. Although the daily distances moved were longer during the wet season compared to the dry season, the daily net displacement was lower, and the residence time higher, indicating a more frequent occurrence of area-concentred searching. In contrast, during the late dry season the foraging sessions were more fragmented and wil-debeests moved more frequently between foraging areas. Surprisingly, predation risk ap