Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree-covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two-month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.