Showing papers by "Daryl Codron published in 2007"

Diets of savanna ungulates from stable carbon isotope composition of faeces

Abstract: 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 C3 (browse) to C4 (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.

Nutritional content of savanna plant foods: implications for browser/grazer models of ungulate diversification

Abstract: Models of herbivore diversification rely heavily on adaptations that reflect the nutritional quality of foods consumed. In particular, browsers and grazers are expected to show dichotomous adaptations to deal with high quality (concentrate) browse-based and poor quality grass-based diets, respectively. In this study, we test the widespread assumption that browse represents a higher quality food source than grass. We analyzed plants from a South African savanna, collected over one dry and one wet season across several habitat types, for percent nitrogen (%N), neutral detergent fiber (NDF), acid detergent fiber (ADF), and acid detergent lignin (ADL) to compare variations in nutritional value of different food types. Results show consistently higher %N and lower NDF and ADF of tree foliage and forbs compared to monocots, but the former have consistently higher ADL, implying a higher fiber digestibility in grass compared with browse. Some fruit species have a high NDF and ADL content, implying poorer nutritional value than is commonly assumed. Our findings are in agreement with several other studies depicting relatively poor digestibility of browse (tree foliage and fruit) compared to grass. Reference to browse as high quality foods is therefore misleading, and models of herbivory that rest on this assumption require revision. The more efficient fiber digestibility recorded in grazers compared to browsers cannot be treated as an adaptation to poor quality diets, but rather to maximize benefits of higher fiber digestibility of grass. Spatio-seasonal variations in plant nutritional seem to reflect seasonal and spatial diet changes expected for grazers and intermediate (mixed) feeders. We propose that future studies require further detail on variations in diet, diet quality, and digestive efficiency to properly understand mechanisms of adaptation.

Significance of diet type and diet quality for ecological diversity of African ungulates

Abstract: Summary 1 We test two nutritional hypotheses for the ecological diversity of ungulates, the browser/grazer (diet type) and diet quality models, among free-ranging herbivores in a South African savanna, the Kruger National Park Tests are based on assessment of relationships between diet type and diet quality with body mass and hypsodonty, two morphological features that have been associated with both elements 2 We use stable carbon isotope ratios of faeces to reconstruct diet in terms of proportions of C3 plants (browse) and C4 plants (grass) consumed by different species in different seasons These data are combined with proxies for diet quality (per cent nitrogen, neutral detergent fibre, acid detergent fibre, and acid detergent lignin) from faeces to track changes in diet quality 3 Two statistical approaches are used in model selection, ie tests of significant correlations based on linear regression analyses, and an information-theory approach (Akaike's Information Criterion) providing insight into strength of evidence for models 4 Results of both methods show that, contrary to many predictions, body mass and diet type are not related, but these data confirm predictions that diet quality decreases with increasing body size, especially during the dry season Hypsodonty, as expected, varies with diet type, increasing with increased grass intake 5 These findings support both a diet type and diet quality model, implying some degree of exclusivity We propose that congruence between models may be achieved through addition of diet quality proxies not included here, because hypsodonty is more likely a reflection of the abrasive properties of consumed foods, ie related to food quality, rather than food type This implies that adaptation to diets of varying quality, through changes in body size and dental features, has been the primary mechanism for diversification in ungulates 6 Our interpretation contrasts with several recent studies advocating diet type as the primary factor, exemplifying that further reconciliation between the two models is needed We discuss the implications of this study for future approaches to achieve a more cohesive understanding of the evolutionary outcomes of herbivore nutrition

Stable isotope characterization of mammalian predator–prey relationships in a South African savanna

Abstract: This paper characterizes predator–prey interactions amongst African mammals from C4 savanna environments using stable carbon and nitrogen isotope proxies for diet. Stable carbon (δ13C) and nitrogen (δ15N) isotope data from hair and faeces of large African mammal carnivores, and herbivores as potential prey, are presented for a diverse range of taxa. Carbon-isotope data imply that most carnivores from the “lowveld” savanna of South Africa form part of C4 grass-based food webs. Nitrogen isotope data show clear differences between trophic levels, although it appears that the magnitude of these differences varies between predators feeding on invertebrates and vertebrates, respectively. Whilst the number of carnivore samples for which data are available is relatively few, and data for prey are restricted mainly to large ungulate herbivores, results clearly demonstrate the potential for future applications of this technique to predator–prey food webs in African savannas. In tandem with traditional approaches, stable isotopes can help elucidate patterns of predator impacts on prey populations, domestic livestock, and resolving similar food webs in palaeoenvironmental contexts.

Stable carbon isotope reconstruction of ungulate diet changes through the seasonal cycle

Abstract: We analysed stable carbon isotope ratios (δ13C) in faeces of 11 African ungulate species from three South African savanna environments to determine whether this approach is sufficiently sensitive to record short-term seasonal diet changes in browsers (BR), mixed-feeders (IM), and grazers (GR). At monthly intervals, faecal δ13C revealed variations in proportions of C3 (browse) to C4 (grass) biomass consumed that were not detected by broader dry versus wet season comparisons, including subtle diet shifts amongst BR and GR. However, trends in faeces were influenced by changes in C3 and C4 plant isotope composition of up to 3‰. Nonetheless, faeces and plants showed strongly similar patterns of variation through the seasonal cycle, so that small diet shifts can be reliably inferred, provided that the variations in plants are controlled for. Faecal δ13C of BR may be further influenced by consumption of isotopically different plant parts such as foliage versus fruit and flowers, and GR faeces may reflect differe...

Trophic ecology of two savanna grazers, blue wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnou

Abstract: The feeding niches and trophic ecology of two South African grazers, blue wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnou, are compared using stable carbon and nitrogen isotope data from feces and tooth dentine collagen. As sympatric, closely related taxa predicted to occupy similar trophic positions, the blue and black wildebeest provide a good model for studying the mechanisms of coexistence and macroevolution in mammals. Data from feces collected from a single reserve in the Free State Province reveal different trophic behaviors between two herds of blue wildebeest and between both compared with a single herd of black wildebeest. These data suggest that sympatric coexistence of blue and black wildebeest is facilitated by differential niche occupation at family group or herd levels, rather than between species. However, such separation does not occur over longer time scales: results from dentine collagen support the hypothesis that the two species are indistinct in terms of trophic behavior, although blue wildebeest show more feeding flexibility, probably because of their wider habitat tolerance range. Similarities in premaxillary width of males and females of both species also suggest that both species are adapted to similar feeding styles. Thus, it is unlikely that changes in trophic behavior provided the trigger for divergence of the black from the blue wildebeest lineage in the Middle Pleistocene. We argue that the case of these two species represents an example of speciation that was not driven by resource competition, as is often assumed for many turnover events in mammalian evolution. We briefly discuss a previous suggestion that links black wildebeest evolution to their more territorial breeding behavior associated with Middle-to-Late Pleistocene landscape changes in southern Africa.