Large herbivore assemblages in a changing climate: incorporating water dependence and thermoregulation.

The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land-use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two-dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water-dependent grazers as wildebeest, zebra and buffalo-dominant species in savanna ecosystems - to be replaced by smaller, less water-dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two-dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change.

Foraging theory upscaled: the behavioural ecology of herbivore movement.

We outline how principles of optimal foraging developed for diet and food patch selection might be applied to movement behaviour expressed over larger spatial and temporal scales. Our focus is on large mammalian herbivores, capable of carrying global positioning system (GPS) collars operating through the seasonal cycle and dependent on vegetation resources that are fixed in space but seasonally variable in availability and nutritional value. The concept of intermittent movement leads to the recognition of distinct movement modes over a hierarchy of spatio-temporal scales. Over larger scales, periods with relatively low displacement may indicate settlement within foraging areas, habitat units or seasonal ranges. Directed movements connect these patches or places used for other activities. Selection is expressed by switches in movement mode and the intensity of utilization by the settlement period relative to the area covered. The type of benefit obtained during settlement periods may be inferred from movement patterns, local environmental features, or the diel activity schedule. Rates of movement indicate changing costs in time and energy over the seasonal cycle, between years and among regions. GPS telemetry potentially enables large-scale movement responses to changing environmental conditions to be linked to population performance.

Megafaunal extinctions: the conservation message from 11,000 years B.p.

At the end of the Pleistocene, the Americas, northern Eurasia: and Australia experienced a vast decline in large mammal diversity, while Africa and tropical Asia were hardly affected. The elimination of the megaberbivores (animals weighing >1000 kg, probably by human predation, removed the vegetation impact of these species. The resultant reduction in habitat mosaic diversity and in forage quality probably precipitated the extinctions of lesser large mammalian species. Surviving megaherbivores in the form of elephants and rhinoceroses are currently being exterminated from many African conservation areas. African savanna ecosystems could prove more resistant to species losses than north temperate ecosystems, because geomorphic factors plus low and erratic rainfall enhance spatial heterogeneity and vegetation quality independently of large herbivore impact Nevertheless, the history of the Hluhluwe Game Reserve in South Africa suggests that certain African ecosystems may become susceptible to an inexorable decline in populations of some large herbivores following the extermination of elephants. If elephants and rhinoceroses cannot be conserved active habitat manipulation will be needed to retain a diverse fauna of large mammals in such regions.

Condensed tannins deter feeding by browsing ruminants in a South African savanna.

The palatability of 14 species of woody plant was assessed for three species of browsing ruminant, namely kudus, impalas and goats. Results show that palatability was most clearly related to leaf contents of condensed tannins. The effect was a threshold one, with all plants containing more than 5% condensed tannins being rejected as food during the wet season period. In contrast palatability was not influenced by concentrations of protein-precipitating polyphenols, and only weakly related to contents of nitrogen, phosphorus, cations, fibre components and other secondary metabolites. Insect herbivory shows a different pattern. These findings support the hypotheses that (i) condensed tannins function to protect plant cell walls against microbial attack; (ii) hydrolyzable tannins function to inactivate the digestive enzymes of insect herbivores. Large mammalian herbivores are influenced by condensed tannins due to their dependance upon microbial fermentation of plant cell walls for part of their energy needs.