Savanna tree-grass competition is modified by substrate type and herbivory

Question Woody plant and grass interactions in savannas have frequently been studied from the perspective of the response of one growth form on the other but seldom evaluated as two-way interactions. What causes woody plant encroachment in semi-arid savannas and what are the competitive responses of tree seedlings and grasses on rocky and sandy substrates? Methods In this greenhouse study, we investigated the influence of substrate and grazing on responses to interspecific competition by tree seedlings and grasses. We measured competitive/facilitative responses on biomass and nutrient status of tree seedlings and grasses grown together. Results Interspecific competition suppressed growth of trees and grasses. Tree seedlings and uncut grass accumulated double the biomass when grown without competition relative to when they competed. Competitive responses varied on different substrates. Grass biomass on rocky substrate showed no response to tree competition, but appeared to be facilitated by trees on sandy substrate. Grass clipping resulted in higher tree seedling biomass on rocky substrate, but not on sandy substrate. There was a positive response of grass nutrient status to competition from tree seedlings. Conclusion Selective grass herbivory in the absence of browsing or suppression of shade-intolerant grasses by trees are commonly cited reasons behind bush encroachment in savannas. We show that grazing may confer a competitive advantage to tree seedlings and promote bush encroachment more readily on rocky substrates. This may be due to the imposed sharing of the soil depth niche on rocky substrates, whereas possible niche separation on sandy substrates minimizes the advantage conferred by reduced competition.