Abstract. At a landscape scale, the combined influence of biotic and abiotic factors may determine the distribution patterns of large herbivores in African savanna ecosystems. Herbivores foraging in these ecosystems may become nutritionally stressed during an annual dry season when both forage quality and quantity are reduced. Additionally, the locations of water sources may impose a landscape-scale constraint on dry-season herbivore distributions. We used logistic regression to analyze 13 years of aerial census data collected in the Kruger National Park (KNP), South Africa, and evaluated hypotheses regarding the relative influences that surface water, forage quality, and forage quantity exert on the dryseason, landscape-scale distribution patterns of eight herbivore species. Hypotheses regarding the degree of correlation between species' distributions and distance to water were developed using previous observations of species' relative water dependence. We also developed hypotheses regarding species' responses to the trade-off that may occur between surface-water constraints and nutritional requirements when either forage quality or quantity is reduced. In general, we expect an increase in species' mean distance to water as a result of individuals mitigating limitations in nutritional requirements (i.e., intake quality or quantity) by foraging farther from water. Our analyses suggest that the trade-off between nutritional requirements and surface-water constraints that species face varies according to the species' water dependence, size, and gut morphology. Of the four grazers considered in our analyses, waterbuck distributions appear to be constrained primarily by surfacewater availability. Distributions of buffalo, a large ruminant grazer, suggest that individuals face a trade-off between nutritional requirements and surface-water constraints when forage quantity is reduced. Alternatively, distributions of wildebeest, a smaller ruminant grazer, suggest that individuals face this trade-off when access to high-quality forage is limited. In comparison to buffalo and wildebeest, the strength of this trade-off is moderate for zebra, a nonruminant similar in size to wildebeest, when either forage quality or quantity is reduced. Distribution patterns for browsers are characterized by a weak relationship with distance to water, as expected for these relatively water-independent species. Population densities relative to forage quality confound exploration of this trade-off for mixed feeders.

Hypotheses to explain diversity among African ungulates focus largely on niche separation along a browser/grazer continuum. However, a number of studies advocate that the browser/grazer distinction insufficiently describes the full extent of dietary variation that occurs within and between taxa. Disparate classification schemes exist because of a lack of uniform and reliable data for many taxa, and failure to incorporate spatio-temporal variations into broader assessments of diet.In this study, we tested predictions for diet and dietary niche separation of African savanna ungulates using stable carbon isotope evidence from faeces for proportions of C 3 (browse) to C 4 (grass) intake among 19 species from the Kruger National Park, South Africa. Dietary predictions from the literature are confirmed in the case of browsers (black rhinoceros Diceros bicornis, giraffe Giraffa camelopardalis, bushbuck Tragelaphus scriptus, kudu Tragelaphus strepsiceros), mixed-feeders (impala Aepyceros melampus, nyala Tragelaphus angasii), and most grazers (white rhinoceros Ceratotherium simum, Burchell's zebra Equus burchellii, warthog Phacochoerus africanus, hippopotamus Hippopotamus amphibius, blue wildebeest Connochaetes taurinus, tsessebe Damaliscus lunatus, waterbuck Kobus ellipsiprymnus). In contrast, several species showed results differing from most expectations derived from the available literature, including eland Taurotragus oryx, steenbok Raphicerus campestris, grey duiker Sylvicapra grimmia, buffalo Syncerus caffer, roan antelope Hippotragus equinus and sable antelope Hippotragus niger. Many of these discrepancies can be accounted for by seasonal and/or regional dietary differences. Cluster analysis based on a data matrix that incorporates the extent of spatio-temporal dietary variation among Kruger Park ungulates reveals several distinct categories of feeding preferences that extend beyond a two-edged browser/grazer dichotomy, such as mixed-feeders with a preference for either forage class, and spatial/seasonal shifts between uniform and mixed-feeding styles among variable browsers (e.g. grey duiker) and variable grazers (e.g. buffalo). These results highlight the need for approaches that are sensitive to spatio-temporal variations and the continuity of diet.

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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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In savannas, the tree–grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.