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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 Management of tree cover, either to curb bush encroachment or to mitigate losses of woody cover to over‐browsing, is a major concern in savanna ecosystems. Once established, trees are often “trapped” as saplings, since interactions among disturbance, plant competition, and precipitation delay sapling recruitment into adult size classes. Saplings can be directly suppressed by wildlife browsing and competition from adjacent plants, and indirectly facilitated by grazers, such as cattle, which feed on neighboring grasses. Yet few experimental studies have simultaneously quantified the effects of cattle and wildlife on sapling growth, particularly over long time scales. We used a series of replicated 4‐ha herbivore‐manipulation plots to investigate the net effects of wildlife and moderate cattle grazing onAcacia drepanolobiumsapling growth over 10 years that encompassed extended wet and dry periods. We also simulated more intense cattle grazing using grass removal treatments (0.5‐m radius around saplings), and we quantified the role of intraspecific tree competition using neighborhood tree surveys (trees within a 3‐m radius). Wildlife, which included elephants, had a positive effect on sapling growth. Wildlife also reduced neighbor tree density during the 10‐yr study, which likely caused the positive effect of wildlife on saplings. Although moderate cattle grazing did not affect sapling growth, grass removal treatments simulating heavy grazing increased sapling growth. Both grass removal and neighbor tree effects on saplings were strongest during above‐average rainfall years following drought. This highlights that livestock‐driven reductions in grass cover and catastrophic wildlife damage to trees during droughts present a need, or an opportunity, for targeted management of sapling growth and woody plant cover during ensuing wet periods. 

Abstract Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long‐lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remain relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape‐scale experimental manipulations—the first a multi‐guild large herbivore exclosure experiment (20 years after establishment) and the second a tree removal experiment (8 years after establishment)—to demonstrate that patterns inOdontotermestermite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15–1,000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that are significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover are intimately tied to herbivore‐driven resource availability. 

Abstract Nearly every terrestrial ecosystem hosts invasive ant species, and many of those ant species construct underground nests near roots and/or tend phloem‐feeding hemipterans on plants. We have a limited understanding of how these invasive ant behaviours change photosynthesis, carbohydrate availability and growth of woody plants.We measured photosynthesis, water relations, carbohydrate concentrations and growth for screenhouse‐rearedAcacia drepanolobiumsaplings on which we had manipulated invasivePheidole megacephalaants and nativeCeroplastessp. hemipterans to determine whether and how soil nesting and hemipteran tending by ants affect plant carbon dynamics. In a field study, we also compared leaf counts of vertebrate herbivore‐excluded and ‐exposed saplings in invaded and non‐invaded savannas to examine how ant invasion and vertebrate herbivory are associated with differences in sapling photosynthetic crown size.Though hemipteran infestations are often linked to declines in plant performance, our screenhouse experiment did not find an association between hemipteran presence and differences in plant physiology. However, we did find that soil nesting byP. megacephalaaround screenhouse plants was associated with >58% lower whole‐crown photosynthesis, >31% lower pre‐dawn leaf water potential, >29% lower sucrose concentrations in woody tissues and >29% smaller leaf areas. In the field, sapling crowns were 29% smaller in invaded savannas than in non‐invaded savannas, mimicking screenhouse results.Synthesis. We demonstrate that soil nesting near roots, a common behaviour byPheidole megacephalaand other invasive ants, can directly reduce carbon fixation and storage ofAcacia drepanolobiumsaplings. This mechanism is distinct from the disruption of a native ant mutualism byP. megacephala, which causes similar large declines in carbon fixation for matureA. drepanolobiumtrees.Acacia drepanolobiumalready has extremely low natural rates of recruitment from the sapling to mature stage, and we infer that these negative effects of invasion on saplings potentially curtail recruitment and reduce population growth in invaded areas. Our results suggest that direct interactions between invasive ants and plant roots in other ecosystems may strongly influence plant carbon fixation and storage.