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Abstract Aboveground ecosystem structure moderates and even confers essential ecosystem functions. This includes an ecosystem’s carbon dynamics, which are strongly influenced by its structure: for example, tropical savannas like those in central Kenya store substantial amounts of carbon in soil. Savannas’ belowground allocation of carbon makes them important for global carbon sequestration, but difficult to monitor. However, the labile soil carbon pool is responsive to changes in ecosystem structure and is thus a good indicator of overall soil organic carbon dynamics. Kenya’s savanna structure is controlled by belowground ecosystem engineers (termites), ambient weather conditions, and the aboveground engineering influences of large-bodied, mammalian consumers. As a result, climate change and biodiversity loss are likely to change savannas’ aboveground structure. To predict likely outcomes of these threats on savanna soil carbon, it is critical to explore the relationships between labile soil carbon and ecosystem structure, local climate, and mammalian consumer community composition. In a large-scale, long-term herbivore exclosure experiment in central Kenya, we sampled labile carbon from surface soils at three distinct savanna structural elements: termite mounds, beneath tree canopies, and the grassland matrix. In one sampling year, we measured total extractable organic carbon (TEOC), total extractable nitrogen (TEN), and extractable microbial biomass for each sample. Across three sampling years with varying weather conditions, we measured rate of labile soil carbon mineralization. We quantified areal coverage of each structural element across herbivore community treatments to estimate pool sizes and mineralization dynamics at the plot scale. Concentrations and stocks of soil TEOC, TEN, and microbial biomass were driven by the structural element from which they were sampled (soils collected under tree canopies generally had the highest of each). Large-bodied herbivore community composition interacted variably with concentrations, stocks, and carbon mineralization, resulting in apparently compensatory effects of herbivore treatment and structural element with no net effects of large herbivore community composition on plot-scale labile carbon dynamics. We confirmed engineering of structural heterogeneity by consumers and identified distinct labile carbon dynamics in each structural element. However, carbon and nitrogen were also influenced by consumer community composition, indicating potentially compensatory interacting effects of herbivore treatment and structural element. These results suggest that one pathway by which consumers influence savanna carbon is by altering its structural heterogeneity and thus the heterogeneity of its plot-scale labile carbon. 

Abstract Irruptions in plant and animal populations are not uncommon, but the factors underlying irruptions are rarely explored quantitatively. In addition, it has been suggested that these irruptions may be reduced by predators or herbivores, but there is a paucity of controlled experimental evidence. Using data from the Kenya Long‐term Exclosure Experiment (KLEE), we show that populations of perennialHibiscusspp. (primarilyHibiscus flavifolius) show multiple short‐term irruptions a year after rainy periods, increasing in abundance in some cases by more than an order of magnitude before declining in ensuing months and years. We demonstrate that these irruptions are largely limited to experimental plots from which large mammalian herbivores have been excluded, particularly megaherbivores (elephants, mostly). This represents a rare controlled, replicated experimental demonstration of top‐down regulation of irruptions. African elephants and giraffes are often at greater risk of local extirpation than other large mammals, and their absence appears to destabilize this African savanna ecosystem. 

Abstract Fire and herbivory have profound effects on vegetation in savanna ecosystems, but little is known about how different herbivore groups influence vegetation dynamics after fire. We assessed the separate and combined effects of herbivory by cattle and wild meso‐ and megaherbivores on postfire herbaceous vegetation cover, species richness, and species turnover in a savanna ecosystem in central Kenya. We measured these vegetation attributes for five sampling periods (from 2013 to 2017) in prescribed burns and unburned areas located within a series of replicated long‐term herbivore exclosures that allow six different combinations of cattle and wild meso‐ and megaherbivores (elephants and giraffes). Vegetation cover (grasses, mainly) and species richness were initially reduced by burning but recovered by 15–27 months after fire, suggesting strong resilience to infrequent fire. However, the rates of recovery differed in plots accessible by different wild and domestic herbivore guilds. Wildlife (but not cattle) delayed postfire recovery of grasses, and the absence of wildlife (with or without cattle) delayed recovery of forbs. Herbivory by only cattle increased grass species richness in burned relative to unburned areas. Herbivory by cattle (with or without wildlife), however, reduced forb species richness in burned relative to unburned areas. Herbivory by wild ungulates (but not cattle) increased herbaceous species turnover in burned relative to unburned areas. Megaherbivores had negligible modifying effects on these results. This study demonstrates that savanna ecosystems are remarkably resilient to infrequent fires, but postfire grazing by cattle and wild mesoherbivores exerts different effects on recovery trajectories of herbaceous vegetation. 

Abstract Climate models predict increases in the frequency and intensity of extreme‐weather events. The impacts of these events may be modulated by biotic agents in unpredictable ways, yet few experiments cover sufficient spatiotemporal scales to measure the interactive effects of multiple extreme events.We used 15 years of a 28‐year experiment spanning several significant droughts to investigate how rainfall, large herbivores, and soil‐engineering termites affect understorey vegetation in a semi‐arid savanna.Herbivory was the dominant influence on community structure—decreasing cover, increasing species richness, and favouring occurrence of annuals relative to perennials—but these effects were contingent on rainfall and termitaria in non‐additive (hence unpredictable) ways.A separate experiment showed that resource enrichment, mimicking the effects of termitaria, does not straightforwardly compensate for top‐down effects of herbivory.Synthesis. Our study highlights the potency of top‐down forcing in African savannas. It suggests impressive robustness to drought and underscores the value of multi‐decadal experiments for studying interactions among multiple drivers of ecosystem dynamics. 

Abstract Fire and herbivory interact to alter ecosystems and carbon cycling. In savannas, herbivores can reduce fire activity by removing grass biomass, but the size of these effects and what regulates them remain uncertain. To examine grazing effects on fuels and fire regimes across African savannas, we combined data from herbivore exclosure experiments with remotely sensed data on fire activity and herbivore density. We show that, broadly across African savannas, grazing herbivores substantially reduce both herbaceous biomass and fire activity. The size of these effects was strongly associated with grazing herbivore densities, and surprisingly, was mostly consistent across different environments. A one‐zebra increase in herbivore biomass density (~100 kg/km²of metabolic biomass) resulted in a ~53 kg/ha reduction in standing herbaceous biomass and a ~0.43 percentage point reduction in burned area. Our results indicate that fire models can be improved by incorporating grazing effects on grass biomass.