Termite mounds create heterogeneity in invertebrate communities across a savanna rainfall gradient

Termite mounds create nutrient hotspots that serve as key resource areas for savanna vegetation and mammalian herbivores. However, despite the key ecological roles performed by termite mounds, few studies have investigated their influence on invertebrate communities, and none have examined such effects across environmental gradients. We hypothesised that termite mounds would support greater numbers of invertebrates than the surrounding savanna matrix and that assemblages would differ in composition due to the enhanced nutritional quality of vegetation on mounds. We also predicted that the differences between on-mound and off-mound invertebrate diversity would be more pronounced in areas where the difference in nutritional value between mounds and the savanna matrix vegetation was most prominent. We tested these hypotheses in Kruger National Park, South Africa, by sampling ground- and grass-dwelling invertebrate herbivores, omnivores and detritivores on and at various distances away from termite mounds at three savanna sites of varying vegetation quality across a rainfall gradient. All invertebrate groups sampled responded to termite mounds, but mound influence varied across trophic groups (Orthoptera showed the clearest patterns), diversity measures (changes in abundance rather than species richness) and with mean annual rainfall (strongest effects at the highest rainfall site). Orthoptera were more abundant on mounds, particularly at the wettest site, and there was a positive relationship between mound size and Orthoptera species richness. Ant assemblage composition on mounds differed from that off mounds and beetle abundance was greater on mounds, possibly as a result of concentrated mammalian herbivore activity and faecal deposition on mounds. Our results suggest that termite mounds are not only important nutrient and foraging hotspots for vertebrates, but that they also introduce heterogeneity in invertebrate communities, especially in nutrient-poor savannas.