Fire, grazers, and browsers interact with grass competition to determine tree establishment in an African savanna

In savanna ecosystems, fire and herbivory alter the competitive relationship between trees and grasses. Mechanistically, grazing herbivores favor trees by removing grass, which reduces tree-grass competition and limits fire. Conversely, browsing herbivores consume trees and limit their recovery from fire. Herbivore feeding decisions are in turn shaped by risk-resource trade-offs that potentially determine the spatial patterns of herbivory. Identifying the dominant mechanistic pathways by which fire and herbivores control tree cover remains challenging, but is essential for understanding savanna dynamics. We used an experiment in the Serengeti ecosystem and a simple simulation driven by experimental results to address two main aims: (1) determine the importance of direct and indirect effects of grass, fire, and herbivory on seedling establishment and (2) establish whether predators determine the spatial pattern of successful seedling establishment via effects on mesoherbivore distribution. We transplanted tree seedlings into plots with a factorial combination of grass and herbivores (present/absent) across a lion-kill-risk gradient in the Serengeti, burning half of the plots near the end of the experiment. Ungrazed grass limited tree seedling survival directly via competition, indirectly via fire, and by slowing seedling growth, which drove higher seedling mortality during fires. These effects restricted seedling establishment to below 18% and, in conjunction with browsing, resulted in seedling establishment dropping below 5%. In the absence of browsing and fire, grazing drove a 7.5-fold increase in seedling establishment. Lion predation risk had no observable impact on herbivore effects on seedling establishment. The severe negative effects of grass on seedling mortality suggests that regional patterns of tree cover and fire may overestimate the role of fire in limiting tree cover, with regular fires representing a proxy for the competitive effects of grass.