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Competition, facilitation, and predation offer alternative explanations for successional patterns of migratory herbivores. However, these interactions are difficult to measure, leaving uncertainty about the mechanisms underlying body-size-dependent grazing-and even whether succession occurs at all. We used data from an 8-year camera-trap survey, GPS-collared herbivores, and fecal DNA metabarcoding to analyze the timing, arrival order, and interactions among migratory grazers in Serengeti National Park. Temporal grazing succession is characterized by a "push-pull" dynamic: Competitive grazing nudges zebra ahead of co-migrating wildebeest, whereas grass consumption by these large-bodied migrants attracts trailing, small-bodied gazelle that benefit from facilitation. "Natural experiments" involving intense wildfires and rainfall respectively disrupted and strengthened these effects. Our results highlight a balance between facilitative and competitive forces in co-regulating large-scale ungulate migrations. more...

Vertebrate herbivores and fire are known to be important drivers of vegetation dynamics in African savannas. It is of particular importance to understand how changes in herbivore population density, especially of elephants, and fire frequency will affect the amount of tree cover in savanna ecosystems, given the critical importance of tree cover for biodiversity, ecosystem function, and human welfare. We developed a spatially realistic simulation model of vegetation, fire, and dominant herbivore dynamics, tailored to the Serengeti ecosystem of east Africa. The model includes key processes such as tree-grass competition, fire, and resource-based density dependence and adaptive movement by herbivores. We used the model to project the ecosystem 100 years into the future from its present state under different fire, browsing (determined by elephant population density), and grazing (with and without wildebeest present) regimes. The model produced the following key results: (1) elephants and fire exert synergistic negative effects on woody cover; when grazers are excluded, the impact of fire and the strength of the elephant-fire interaction increase; (2) at present population densities of 0.15 elephants/km2, the total amount of woody cover is predicted to remain stable in the absence of fire, but the mature tree population is predicted to decline regardless of the fire regime; without grazers present to mitigate the effects of fire, the size structure of the tree population will become dominated by seedlings and mature trees; (3) spatial heterogeneity in tree cover varies unimodally with elephant population density; fire increases heterogeneity in the presence of grazers and decreases it in their absence; (4) the marked rainfall gradient in the Serengeti directly affects the pattern of tree cover in the absence of fire; with fire, the woody cover is determined by the grazing patterns of the migratory wildebeest, which are partly rainfall driven. Our results show that, in open migratory ecosystems such as the Serengeti, grazers can modulate the impact of fire and the strength of the interaction between fire and browsers by altering fuel loads and responding to the distribution of grass across the landscape, and thus exert strong effects on spatial patterns of tree cover. more...