There were strong seasonal differences in wild herbivore densities between the reserve and the ranches during 1977–2010. Individual species responded differentially to pastoralism and protection. Three distinct patterns were apparent, all of which could be explained in terms of distinctions in body size and feeding guild and their consequences for nutritional quality and quantity of forage, predation risk and competition with livestock.

Small sized herbivores Small species that are constrained by food quality and predation tend to prefer short grass areas (Fryxell 1991; Illius and Gordon 1992) and were thus Buparlisib more abundant in the ranches than the reserve regardless of season or feeding guild as revealed by the significant differences between their densities in the reserve and the ranches during 1977–2010. Repeated livestock grazing in the same areas of the ranches probably increased the crude protein

production of grasses (Anderson et al. 2010; Augustine et al. 2010), enabling the small grazers to derive sufficient energy by selecting high-quality forage from the low-biomass areas (Fryxell et al. 2005). Reduced predation risk as a result of lower CB-5083 nmr vegetation cover on the ranches (Ogutu et al. 2005) is yet another advantage of concentrating in the short grass plains, since tall grasses conceal ambush predators and significantly increase their efficiency at catching prey animals (Hopcraft et al. 2005). The distribution patterns we observed for small herbivores are therefore concordant with the initial expectation that small herbivores (except eltoprazine warthog) should concentrate in areas of relatively fewer predators (safer) and shorter grasses maintained by heavy livestock grazing in the ranches. This outcome also concurs with findings of studies encompassing

a variety of spatial scales and species (Olff et al. 2002; Cromsigt and Olff 2006) besides reinforcing the notion that both predation and resource limitation act simultaneously in limiting herbivore populations (Sinclair et al. 2003). Medium sized herbivores The second pattern was expressed by species that moved between the ranches and the reserve seasonally, suggesting that they preferred either the reserve or the ranches PF-02341066 datasheet depending on season. Specifically, the medium-sized topi, wildebeest and zebra moved seasonally between the reserve and the ranches, thus supporting our second prediction. As a result, medium herbivores had higher densities in the ranches in the wet season but higher densities in the reserve in the dry season. This pattern suggests that medium herbivores tend to utilize the ranches when water and short, nutritious grasses, created and maintained by heavy livestock grazing (Rannestad et al. 2006), are widely available, enabling them to enhance their total protein consumption (McNaughton 1976).