Ungulate community structure and ecological processes: body size, hoof area and trampling in African savannas.

A wide range of bioenergetic, production, life history and ecological traits scale with body size in vertebrates. However, the consequences of differences in community body-size structure for ecological processes have not been explored. We studied the scaling relationships between body mass, shoulder height, hoof area, stride length and daily ranging distance in African ungulates ranging in size from the 5 kg dik-dik to the 5,000 kg African elephant, and the implications of these relationships on the area trampled by single and multispecies herbivore communities of differing structure. Hoof area, shoulder height and stride length were strongly correlated with body mass (Pearson's r >0.98, 0.95 and 0.90, respectively). Hoof area scaled linearly to body mass with a slope of unity, implying that the pressures exerted on the ground per unit area by a small antelope and an elephant are identical. Shoulder height and stride length scaled to body mass with similar slopes of 0.32 and 0.26, respectively; larger herbivores have relatively shorter legs and take relatively shorter steps than small herbivores, and so trample a greater area of ground per unit distance travelled. We compared several real and hypothetical single- and multi-species ungulate communities using exponents of between 0.1 and 0.5 for the body mass to daily ranging distance relationship and found that the estimated area trampled was greater in communities dominated by larger animals. The impacts of large herbivores are not limited to trampling. Questions about the ecological implications of community body-size structure for such variables as foraging and food intake, dung quality and deposition rates, methane production, and daily travelling distances remain clear research priorities.