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  1. null (Ed.)
  2. 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.

     
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  3. 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.

     
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  4. 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.

     
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  5. Abstract

    Invasive ants shape assemblages and interactions of native species, but their effect on fundamental ecological processes is poorly understood. In East Africa,Pheidole megacephalaants have invaded monodominant stands of the ant‐treeAcacia drepanolobium, extirpating native ant defenders and rendering trees vulnerable to canopy damage by vertebrate herbivores. We used experiments and observations to quantify direct and interactive effects of invasive ants and large herbivores onA. drepanolobiumphotosynthesis over a 2‐year period. Trees that had been invaded for ≥ 5 years exhibited 69% lower whole‐tree photosynthesis during key growing seasons, resulting from interaction between invasive ants and vertebrate herbivores that caused leaf‐ and canopy‐level photosynthesis declines. We also surveyed trees shortly before and after invasion, finding that recent invasion induced only minor changes in leaf physiology. Our results from individual trees likely scale up, highlighting the potential of invasive species to alter ecosystem‐level carbon fixation and other biogeochemical cycles.

     
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  6. Abstract

    Biological invasions can lead to the reassembly of communities and understanding and predicting the impacts of exotic species on community structure and functioning are a key challenge in ecology. We investigated the impact of a predatory species of invasive ant,Pheidole megacephala, on the structure and function of a foundational mutualism betweenAcacia drepanolobiumand its associated acacia‐ant community in an East African savanna. Invasion byP. megacephalawas associated with the extirpation of three extrafloral nectar‐dependentCrematogasteracacia ant species and strong increases in the abundance of a competitively subordinate and locally rare acacia ant species,Tetraponera penzigi, which does not depend on host plant nectar. Using a combination of long‐term monitoring of invasion dynamics, observations and experiments, we demonstrate thatP. megacephaladirectly and indirectly facilitatesT. penzigiby reducing the abundance ofT. penzigi’s competitors (Crematogasterspp.), imposing recruitment limitation on these competitors, and generating a landscape of low‐reward host plants that favor colonization and establishment by the strongly dispersingT. penzigi. Seasonal variation in use of host plants byP. megacephalamay further increase the persistence ofT. penzigicolonies in invaded habitat. The persistence of theT. penzigi–A. drepanolobiumsymbiosis in invaded areas afforded host plants some protection against herbivory by elephants (Loxodonta africana), a key browser that reduces tree cover. However, elephant damage onT. penzigi‐occupied trees was higher in invaded than in uninvaded areas, likely owing to reducedT. penzigicolony size in invaded habitats. Our results reveal the mechanisms underlying the disruption of this mutualism and suggest thatP. megacephalainvasion may drive long‐term declines in tree cover, despite the partial persistence of the ant–acacia symbiosis in invaded areas.

     
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  7. Abstract

    Savanna tree cover is dynamic due to disturbances such as fire and herbivory. Frequent fires can limit a key demographic transition from sapling to adult height classes in savanna trees. Saplings may be caught in a ‘fire trap’, wherein individuals repeatedly resprout following fire top‐kill events. Saplings only rarely escape the cycle by attaining a fire‐resistant height (e.g. taller than the minimum scorch height) during fire‐free intervals.

    Large mammalian herbivores also may trap trees in shorter size classes. Browsing herbivores directly limit sapling height, while grazing herbivores such as cattle facilitate sapling growth indirectly via grass removal. Experimental studies investigating how meso‐wildlife, megaherbivores and domestic livestock affect height of resprouts following fire are rare, but necessary for fully understanding how herbivory may reinforce (or counteract) the fire trap. In our study system, interactive fire–herbivore effects on transitions from sapling (<1 m) to adult tree (>1 m) height classes may be further influenced by plant defences, such as symbiotic ants.

    We used the Kenya Long‐term Exclosure Experiment (KLEE) to investigate how post‐fire resprout size of a widespread monodominant East African tree,Acacia drepanolobiumwas influenced by (a) herbivory by different combinations of cattle, meso‐wildlife (15–1,000 kg) and megaherbivores (>1,000 kg) and (b) the presence of acacia–ant mutualists that confer tree defences. We sampled height, stem length and ant occupancy of resprouts exposed to different herbivore combinations before and after controlled burns.

    Resprout height of saplings that were short prior to fire (<1 m) was reduced primarily by meso‐wildlife. Negative effects of elephants on post‐fire resprout height increased with pre‐fire tree size, suggesting that resprouts of the tallest trees (with the greatest potential to escape the fire trap cycle) were preferentially browsed and reduced in height by elephants. There were no significant cattle effects.

    Synthesis. We provide experimental evidence for two potential pathways through which large herbivores exert control over sapling escape from the fire trap: (a) post‐fire meso‐wildlife browsing of short (<1 m) resprouts and (b) elephant browsing of the largest size class of resprouts, which would otherwise be most likely to escape the fire trap.

     
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