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.

Baboons are dietary generalists, consuming a wide range of food items in varying proportions. It is thus difficult to quantify and explain the dietary behavior of these primates. We present stable carbon (delta(13)C) and nitrogen (delta(15)N) isotopic data, and percentage nitrogen (%N), of feces from chacma baboons (Papio ursinus) living in two savanna environments of South Africa: the mountainous Waterberg region and the low-lying Kruger National Park. Baboons living in the more homogeneous landscapes of the Waterberg consume a more isotopically heterogeneous diet than their counterparts living in Kruger Park. Grasses and other C(4)-based foods comprise between approximately 10-20% (on average) of the bulk diet of Kruger Park baboons. Carbon isotopic data from the Waterberg suggest diets of approximately 30-50% grass, which is higher than generally reported for baboons across the African savanna. Based on observations of succulent-feeding, we propose that baboons in the Waterberg consume a mix of C(4) grasses and CAM-photosynthesizing succulents in combined proportions varying between approximately 5-75% (average, approximately 35%). Fecal delta(15)N of baboons is lower than that of sympatric ungulates, which may be due to a combination of low levels of faunivory, foraging on subterranean plant parts, or the use of human foods in the case of Kruger Park populations. Fecal N levels in baboons are consistently higher than those of sympatric ungulate herbivores, indicating that baboons consume a greater proportion of protein-rich foods than do other savanna mammals. These data suggest that chacma baboons adapt their dietary behavior so as to maximize protein intake, regardless of their environment.

The African elephant (Loxodonta africana) is a large-bodied, generalist herbivore that eats both browse and grass. The proportions of browse and grass consumed are largely expected to reflect the relative availability of these resources. We investigated variations in browse (C(3) biomass) and grass (C(4)) intake of the African elephant across seasons and habitats by stable carbon isotope analysis of elephant feces collected from Kruger National Park, South Africa. The results reflect a shift in diet from higher C(4) grass intake during wet season months to more C(3) browse-dominated diets in the dry season. Seasonal trends were correlated with changes in rainfall and with nitrogen (%N) content of available grasses, supporting predictions that grass is favored when its availability and nutritional value increase. However, switches to dry season browsing were significantly smaller in woodland and grassland habitats where tree communities are dominated by mopane (Colophospermum mopane), suggesting that grasses were favored here even in the dry season. Regional differences in diet did not reflect differences in grass biomass, tree density, or canopy cover. There was a consistent relationship between %C(4) intake and tree species diversity, implying that extensive browsing is avoided in habitats characterized by low tree species diversity and strong dominance patterns, i.e., mopane-dominated habitats. Although mopane is known to be a preferred species, maintaining dietary diversity appears to be a constraint to elephants, which they can overcome by supplementing their diets with less abundant resources (dry season grass). Such variations in feeding behavior likely influence the degree of impact on plant communities and can therefore provide key information for managing elephants over large, spatially diverse, areas.

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic-and hence dietary-niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C 4 grass consumption through the twentieth century. We suggest that switching between C 3 browsing and C 4 grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.

Stable carbon isotope analyses of vertebrate hard tissues such as bones, teeth, and tusks provide information about animal diets in ecological, archeological, and paleontological contexts. There is debate about how carbon isotope compositions of collagen and apatite carbonate differ in terms of their relationship to diet, and to each other. We evaluated relationships between δ13Ccollagen and δ13Ccarbonate among free‐ranging southern African mammals to test predictions about the influences of dietary and physiological differences between species. Whereas the slopes of δ13Ccollagen–δ13Ccarbonate relationships among carnivores are ≤1, herbivore δ13Ccollagen increases with increasing dietary δ13C at a slower rate than does δ13Ccarbonate, resulting in regression slopes >1. This outcome is consistent with predictions that herbivore δ13Ccollagen is biased against low protein diet components (13C‐enriched C4 grasses in these environments), and δ13Ccarbonate is 13C‐enriched due to release of 13C‐depleted methane as a by‐product of microbial fermentation in the digestive tract. As methane emission is constrained by plant secondary metabolites in browse, the latter effect becomes more pronounced with higher levels of C4 grass in the diet. Increases in δ13Ccarbonate are also larger in ruminants than nonruminants. Accordingly, we show that Δ13Ccollagen‐carbonate spacing is not constant within herbivores, but increases by up to 5 ‰ across species with different diets and physiologies. Such large variation, often assumed to be negligible within trophic levels, clearly cannot be ignored in carbon isotope‐based diet reconstructions.

Species’ partitioning of resources remains one of the most integral components for understanding community assembly. Analysis of stable carbon and nitrogen isotopes in animal tissues has the potential to help resolve patterns of partitioning because these proxies represent the individual’s diet and trophic niche, respectively. Using free-ranging rodents in a southern African savanna as a model community, we find that syntopic species within habitats occupy distinct isotope niches. Moreover, species with strongly overlapping isotope niches did not overlap in their spatial distribution patterns, suggesting an underlying effect of competitive exclusion. Niche conservatism appears to characterize the behaviour of most species in our sample - with little or no observed changes across habitats - with the exception of one species, Mastomys coucha. This species displayed a generalist distribution, being found in similar abundances across a variety of habitats. This spatial pattern was coupled with a generalist isotope niche that shifted across habitats, likely in response to changes in species composition over the same spatial gradient. The case for M. coucha supports contentions that past competition effects played a significant evolutionary role in shaping community structures of today, including the absence of strong interspecific niche overlaps within particular habitats. Our study highlights the value of stable isotope approaches to help resolve key questions in community ecology, and moreover introduces novel analytical approaches to quantifying isotope niche breadths and niche overlaps that are easily comparable with traditional metrices.

Large mammal ecosystems have relatively simple food webs, usually comprising three-and sometimes only two-trophic links. Since many syntopic species from the same trophic level therefore share resources, dietary niche partitioning features prominently within these systems. In African and other subtropical savannas, stable carbon isotopes readily distinguish between herbivore species for which foliage and other parts of dicot plants (13 C-depleted C 3 vegetation) are the primary resource (browsers) and those for which grasses (13 C-enriched C 4 vegetation) are staples (grazers). Similarly, carbon isotopes distinguish between carnivore diets that may be richer in either browser, grazer, or intermediate-feeding prey. Here, we investigate levels of carbon and nitrogen isotopic niche variation and niche partitioning within populations (or species) of carnivores and herbivores from South African savannas. We emphasize predictable differences in within-population trends across trophic levels: we expect that herbivore populations, which require more foraging effort due to higher intake requirements, are far less likely to display within-population resource partitioning than carnivore populations. Our results reveal generally narrower isotopic niche breadths in herbivore than carnivore populations, but more importantly we find lower levels of isotopic differentiation across individuals within herbivore species. While these results offer some support for our general hypothesis, the current paucity of isotopic data for African carnivores limits our ability to test the complete set of predictions arising from our hypothesis. Nevertheless, given the different ecological and ecophysiological constraints to foraging behavior within each trophic level, comparisons across carnivores, and herbivores, which are possible within such simplified foodwebs, make these systems ideal for developing a process-based understanding of conditions underlying the evolution of intra-specific, individual-level separation of ecological niches.

Discussions about early hominin diets have generally excluded grass leaves as a staple food resource, despite their ubiquity in most early hominin habitats. In particular, stable carbon isotope studies have shown a prevalent C component in the diets of most taxa, and grass leaves are the single most abundant C resource in African savannas. Grass leaves are typically portrayed as having little nutritional value (e.g., low in protein and high in fiber) for hominins lacking specialized digestive systems. It has also been argued that they present mechanical challenges (i.e., high toughness) for hominins with bunodont dentition. Here, we compare the nutritional and mechanical properties of grass leaves with the plants growing alongside them in African savanna habitats. We also compare grass leaves to the leaves consumed by other hominoids and demonstrate that many, though by no means all, compare favorably with the nutritional and mechanical properties of known primate foods. Our data reveal that grass leaves exhibit tremendous variation and suggest that future reconstructions of hominin dietary ecology take a more nuanced approach when considering grass leaves as a potential hominin dietary resource.