RESEARCH ARTICLE Behaviour-Related Scalar Habitat Use by

Studies of habitat use by animals must consider behavioural resource requirements at dif-ferent scales, which could influence the functional value of different sites. Using Cape buf-falo (Syncerus caffer caffer) in the Okavango Delta, Botswana, we tested the hypotheses that behaviour affected use between and within habitats, hereafter referred to as macro-and microhabitats, respectively. We fitted GPS-enabled collars to fifteen buffalo and used the distances and turning angles between consecutive fixes to cluster the resulting data into resting, grazing, walking and relocating behaviours. Distance to water and six vegetation characteristic variables were recorded in sites used for each behaviour, except for relocat-ing, which occurred too infrequently. We used multilevel binomial and multinomial logistic regressions to identify variables that characterised seasonally-preferred macrohabitats and microhabitats used for different behaviours. Our results showed that macrohabitat use was linked to behaviour, although this was least apparent during the rainy season, when resources were most abundant. Behaviour-related microhabitat use was less significant, but variation in forage characteristics could predict some behaviour within all macrohabitats. The variables predicting behaviour were not consistent, but resting and grazing sites were more readily identifiable than walking sites. These results highlight the significance of rest-ing, as well as foraging, site availability in buffalo spatial processes. Our results emphasise the importance of considering several behaviours and scales in studies of habitat use to understand the links between environmental resources and animal behavioural and spatial ecology