Jacqueline Codron

Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces

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 (C3 biomass) and grass (C4) 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 C4 grass intake during wet season months to more C3 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 %C4 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.

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

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 C4 grass consumption through the twentieth century. We suggest that switching between C3 browsing and C4 grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.

Within-Population Isotopic Niche Variability in Savanna Mammals: Disparity between Carnivores and Herbivores

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 (13C-depleted C3 vegetation) are the primary resource (browsers) and those for which grasses (13C-enriched C4 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 behaviour 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.

Predator–prey interactions amongst Permo‐Triassic terrestrial vertebrates as a deterministic factor influencing faunal collapse and turnover

Unlike modern mammalian communities, terrestrial Paleozoic and Mesozoic vertebrate systems were characterized by carnivore faunas that were as diverse as their herbivore faunas. The comparatively narrow food base available to carnivores in these paleosystems raises the possibility that predator–prey interactions contributed to unstable ecosystems by driving populations to extinction. Here, we develop a model of predator–prey interactions based on diversity, abundance and body size patterns observed in the Permo‐Triassic vertebrate fossil record of the Karoo Basin, South Africa. Our simulations reflect empirical evidence that despite relatively high carnivore: herbivore species ratios, herbivore abundances were sufficient for carnivores to maintain required intake levels through most of the Karoo sequence. However, high mortality rates amongst herbivore populations, even accounting for birth rates of different‐sized species, are predicted for assemblages immediately preceding the end‐Guadalupian and end‐Permian mass extinctions, as well as in the Middle Triassic when archosaurs replaced therapsids as the dominant terrestrial fauna. These results suggest that high rates of herbivore mortality could have played an important role in biodiversity declines leading up to each of these turnover events. Such declines would have made the systems especially vulnerable to subsequent stochastic events and environmental perturbations, culminating in large‐scale extinctions.

Meso-Carnivore Niche Expansion in Response to an Apex Predator's Reintroduction - a Stable Isotope Approach

Apex predators can have considerable impacts on meso-carnivore diets, through competition or facilitation. Facilitation occurs when smaller predators consume carrion created by larger predators, especially large-bodied prey species normally inaccessible to meso-carnivores. In contrast, apex predators can also negatively affect meso-carnivore consumption of important resources through competitive interactions. Thus, predicting meso-carnivore responses to trophic structure changes (i.e. apex predator extirpation or reintroduction) is often difficult. We investigated stable carbon and nitrogen isotope niche breadths of black-backed jackal (Canis mesomelas) in response to the reintroduction of an apex predator, the African lion (Panthera leo), to the Karoo National Park, South Africa. Jackal faecal isotopic niche widths were larger in post-lion than pre-lion samples, indicating a niche expansion to include pure C3- and C4-based food sources when lions were present. Most prey items of this nature in the study area are large-bodied ungulates. Our results agree with results of traditional scat analysis, which showed that prey species >92 kg were consumed more often after the lion reintroduction. Stable isotope data from carnivore faeces are effective for tracking responses of wildlife to changing ecological conditions, providing an alternative source of information about changes in community structure brought about by management interventions.

Within trophic level shifts in collagen-carbonate stable carbon isotope spacing are propagated by diet and digestive physiology in large mammal herbivores

Abstract 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.