Termites and large herbivores influence seed removal rates in an African savanna

Seed removal can influence plant community dynamics, composition, and resulting vegetation characteristics. In the African savanna, termites and large herbivores influence vegetation in various ways, likely including indirect effects on seed predators and secondary dispersers. However, the intensity and variation of seed removal rates in African savannas has seldom been studied. We experimentally investigated whether termites and large herbivores were important factors in the mechanisms contributing to observed patterns in tree species composition on and off mounds, in Lake Mburo National Park, Uganda. Within fenced (excluding large herbivores) and unfenced termite mound and adjacent savanna plots, we placed seeds of nine native tree species within small open "cages," accessed by all animals, roofed cages that only allowed access to small vertebrates and invertebrates, and closed cages that permitted access by smaller invertebrates only (5 mm wire mesh). We found that mean seed removal rate was high (up to 87.3% per 3 d). Mound habitats experienced significantly higher removal rates than off-mound habitats. The mean removal rate of native seeds from closed cages was 11.1% per 3 d compared with 19.4% and 23.3% removed per 3 d in the roofed and open cages, respectively. Smaller seeds experienced higher removal rates than larger seeds. Large herbivore exclusion on mounds reduced native seed removal rates by a mean of 8.8% in the open cages, but increased removal rates by 1.7% in the open cages when off-mound habitats were fenced. While removal rates from open cages were higher on active mounds (30.9%) than on inactive mounds (26.7%), the removal rates from closed cages were lower on active vs. inactive mounds (6.1% vs. 11.6%, respectively). Thus, we conclude that large herbivores and Macrotermes mounds influence seed removal rates, though these effects appear indirect.