Abstract:Elephant are believed to be one of the main ecological drivers in the conversion of savanna woodlands to grassland. We assessed the impacts of elephant on large trees (≥5 m in height) in the southern section of the Kruger National Park. Tree dimensions and utilization by elephant were recorded for 3082 individual trees across 22 transects (average length of 3 km and 10 m wide). Sixty per cent of the trees exhibited elephant utilization and 4% were dead as a direct result of elephant foraging behaviour. Each height class of tree was utilized in proportion to abundance. However, the size of the tree and the species influenced the intensity of utilization and foraging approach. Sclerocarya birrea was actively selected for and experienced the highest proportional utilization (75% of all trees). Interestingly, the proportion of large trees that were utilized and pushed over increased with distance from permanent water, a result which has implications for the provision of water in the KNP. We conclude that mortality is likely to be driven by a combination of factors including fire, drought and disease, rather than the actions of elephant alone. Further investigation is also required regarding the role of senescence and episodic mortality.

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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.

The vegetation dynamics of the savanna ecosystem are driven by complex interactions between biotic and abiotic factors, and thus are expected to exhibit emergent properties of biocomplexity. We explore the relative importance of static and dynamic drivers in explaining the patterns of mortality of large trees in the Kruger National Park, South Africa. Data on large trees were collected from 22 transects in April 2006, and these transects were re‐sampled in November 2008. Of the 2546 individually‐identified trees that were re‐sampled, 290 (11.4%) died in the interim. We tested several competing hypotheses with varying levels of complexity, and found that mortality of large trees was affected mainly by both static (geophysical and landscape characteristics) and dynamic (elephant damage and fire) factors that were either additive or interactive in their effects. Elephant damage was the main predictor of tree mortality, but fire also played an important role depending on the landscape type. Other static variables such as position‐on‐slope, height below canopy, and altitude had weak effects in explaining tree mortality. These results indicate that keystone features such as large trees, show differential vulnerability to mortality that is landscape‐specific. For conservation managers, this implies that the dynamic drivers (elephant and fire) of tree mortality have to be managed at the specific landscape‐level. We suggest that this emergent biocomplexity in the spatial and temporal patterns of large tree mortality is not unique to the African savannas, but is likely widespread across heterogeneous landscapes.

Summary 1.Globally, both climatic patterns and nitrogen deposition rates show directional changes over time. It is uncertain how woody seedlings, which coexist with herbaceous plants in savannas, respond to concurrent changes in water and nutrient availability. 2. We investigated competition effects between herbaceous vegetation and tree seedlings ( Colophospermum mopane ) under changed water and nutrient (fertilized) conditions in a garden experiment situated in a semi-arid savanna. 3. Herbaceous competition significantly suppressed woody seedling growth. The effect of herbaceous competition on woody seedling growth remained constant with both increasing water and nutrient availability. However, during a wet-season drought, herbaceous competition apparently caused premature leaf senescence in non-irrigated treatments. Fertilization exacerbated negative competition effects during the drought, while irrigation prevented leaf loss of tree seedlings in spite of herbaceous competition and fertilization. 4. Based on a conceptual model, we propose that the vigorous response of herbaceous plants to increased nutrient availability leads to faster depletion of soil water, which increasingly causes water stress in woody seedlings if the interval between watering events is prolonged, e.g. during wet-season droughts. 5. Synthesis . Our data support the notion that changes in drought frequency are of greater importance to woody recruitment success than changes in annual rainfall amount. Based on the water and nutrient interactions observed in our experiment, we suggest that the effect of increased nitrogen deposition on woody seedling recruitment is contingent on water availability.

Aim Artificial water points are often used in protected savanna ecosystems to maintain populations of large herbivores. However, these interventions lead to increased ranging and foraging pressure and can negatively impact important habitats and species. This study investigated the influence of artificial water provision on the foraging and movement paths of an African elephant population and modelled the impact of changing water availability on sensitive habitat types.Location Tembe Elephant Park (TEP), KwaZulu-Natal, South Africa.Methods We mapped and classified 414 km of elephant movement and foraging paths in a 300-km 2 fenced protected area. The data were analysed to determine the relationship between path size, distribution and distance to water. We also used a logistic modelling approach to explore the predicted effects of removing artificial water points on path distribution.Results Elephant paths were unevenly distributed throughout the habitats of TEP and the most established and heavily used paths were found closest to water. We also discovered a number of elephant 'rest areas' along the paths, which were distinct clearings that tended to be close to water and in sand forest habitat. Our model predicted that the removal of artificial water points would reduce the area crossed by elephant paths by 79%, leading to an 89% reduction in the presence of elephant paths in sand forest.Main conclusions Our study provides further evidence that manipulating surface water availability can be a useful tool for managing large herbivore impacts on vegetation and acts as the basis for further research on the trade-offs between conservation objectives.

Elephants (Loxodonta africana) exhibit pronounced sexual dimorphism, and in this study we test the prediction that the differences in body size and sociality are significant enough to drive divergent foraging strategies and ultimately sexual segregation. Body size influences the foraging behaviour of herbivores through the differential scaling coefficients of metabolism and gut size, with larger bodied individuals being able to tolerate greater quantities of low-quality, fibrous vegetation, whilst having lower mass-specific energy requirements. We test two distinct theories: the scramble competition hypothesis (SCH) and the forage selection hypothesis (FSH). Comprehensive behavioural data were collected from the Pongola Game Reserve and the Phinda Private Game Reserve in South Africa over a 2.5-year period. The data were analysed using sex as the independent variable. Adult females targeted a wider range of species, adopted a more selective foraging approach and exhibited greater bite rates as predicted by the body size hypothesis and the increased demands of reproductive investment (lactation and pregnancy). Males had longer feeding bouts, displayed significantly more destructive behaviour (31% of observations, 11% for females) and ingested greater quantities of forage during each feeding bout. The independent ranging behaviour of adult males enables them to have longer foraging bouts as they experience fewer social constraints than females. The SCH was rejected as a cause of sexual segregation due to the relative abundance of low quality forage, and the fact that feeding heights were similar for both males and females. However, we conclude that the differences in the foraging strategies of the sexes are sufficient to cause spatial segregation as postulated by the FSH. Sexual dimorphism and the associated behavioural differences have important implications for the management and conservation of elephant and other dimorphic species, with the sexes effectively acting as distinct "ecological species".