Interactions between large herbivores and litter removal by termites across a rainfall gradient in a South African savanna

Litter-feeding termites influence key aspects of the structure and functioning of semi-arid ecosystems around the world by altering nutrient and material fluxes, affecting primary production, foodweb dynamics and modifying vegetation composition. Understanding these complex effects depends on quantifying spatial heterogeneity in termite foraging activities, yet such information is scarce for semi-arid savannas. Here, the amount of litter that was removed from 800 litterbags in eight plots (100 litterbags per plot) was measured in Hluhluwe-iMfolozi Park (HiP) South Africa. These data were used to quantify variation in litter removal at two spatial scales: the local scale (within 450-m(2) plots) and the landscape scale (among sites separated by 8-25 km). Subsequently, we attempted to understand the possible determinants of termites' foraging patterns by testing various ecological correlates, such as plant biomass and bare ground at small scales and rainfall and fences that excluded large mammalian herbivores at larger scales. No strong predictors for heterogeneity in termite foraging intensity were found at the local scale. At the landscape scale termite consumption depended on an interaction between rainfall and the presence of large mammalian herbivores: litter removal by termites was greater in the presence of large herbivores at the drier sites but lower in the presence of large herbivores at the wetter sites. The effect of herbivores on termite foraging intensity may indicate a switch between termites and large herbivore facilitation and competition across a productivity gradient. In general, litter removal decreased with increasing mean annual rainfall, which is in contrast to current understanding of termite consumption across rainfall and productivity gradients. These results generate novel insights into termite ecology and interactions among consumers of vastly different body sizes across spatial scales