Summary1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is important for ecological research, biodiversity conservation and ecosystem management. However, this understanding is complicated by the effects of spatial scale, because the scale of analysis affects the quantification of species-environment relationships. 2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on the analysis and prediction of habitat selection by African elephants in Kruger National Park, South Africa. 3. We focused on the spatial scaling of the elephants' response to their main resources, forage and water, and found that the quantification of habitat selection strongly depended on the scales at which environmental context was considered. Moreover, the inclusion of environmental context at characteristic scales (i.e. those at which habitat selectivity was maximized) increased the predictive capacity of habitat suitability models. 4. The elephants responded to their environment in a scale-dependent and perhaps hierarchical manner, with forage characteristics driving habitat selection at coarse spatial scales, and surface water at fine spatial scales. 5. Furthermore, the elephants exhibited sexual habitat segregation, mainly in relation to vegetation characteristics. Male elephants preferred areas with high tree cover and low herbaceous biomass, whereas this pattern was reversed for female elephants. 6. We show that the spatial distribution of elephants can be better understood and predicted when scale-dependent species-environment relationships are explicitly considered. This demonstrates the importance of considering the influence of spatial scale on the analysis of spatial patterning in ecological phenomena.

Free-ranging African elephants Loxodonta africana use their front feet frequently during the process of foraging and this could be the reason for the high prevalence of physical injuries to these parts of the body. Although the occurrence of severe lameness caused by foot lesions in adult elephants has already been investigated and the clinical and pathological findings have been reported, the effect of foot injuries on glucocorticoid levels as a potential physiological stress response has not been examined. Given the practical difficulties involved in monitoring unpredictable events in free-ranging animals, like the occurrence of foot injuries in elephants, it is not surprising that information regarding the endocrine correlates of physical injury is still limited for elephants. In our study we investigated the effects of foot injuries on concentrations of faecal glucocorticoid metabolites (GCM), body condition score (BCS) and reproductive behaviour in two GPS/radio-collared elephant bulls in the Kruger National Park, South Africa. We monitored the bulls aged 40+ (Bull 1) and 30+ (Bull 2) 2-3 times per week for 13 months starting in June 2007 and frequently collected faecal samples for non-invasive hormone monitoring. Faecal samples were lyophilised, extracted and assayed with an enzyme immunoassay which detects GCM with a 3a-hydroxy-11-oxo-structure. Both bulls acquired foot injuries (right-front), which caused temporary lameness, but the effect of injury on GCM concentration differed between bulls (P , 0.001). In Bull 1 the injury lasted 6 250 days and was associated with an up to four-fold increase in GCM concentrations (P , 0.001) and his BCS reduced from 'good' to 'very thin' by the end of the injury period. In Bull 2 the injury lasted 65 days and was associated with a smaller increase in GCM concentrations (P ¼ 0.03) together with a reduced loss in condition when compared to Bull 1. Following recovery, the condition of both bulls improved progressively and faecal GCM returned to baseline concentrations. Collectively, the data clearly underlined the value of non-invasive hormone measurements as a tool to provide information on the level of stress experienced by elephants. Thus, monitoring GCM levels could help improve the assessment of an elephant's state of health.

Human footprint and protected areas shape elephant range across Africa Highlights d 229 African elephants were tracked across forest, Sahel, savanna, and bushveld sites d Protected areas and the human footprint both constrain elephant range size d 62% of the African continent is potential elephant habit, of which elephants use 17% d 85% of potential elephant habitat lies outside current protected areas

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