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    Tree resprout dynamics following fire depend on herbivory by wild ungulate herbivores
    (Journal of Ecology, 2019-04) Kimuyu, Duncan M.; LaMalfa, Eric M.; Sensenig, Ryan L.; Young, Truman P.; Riginos, Corinna; Veblen, Kari E.
    1. 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. 2. 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. 3. We used the Kenya Long-term Exclosure Experiment (KLEE) to investigate how post‐fire resprout size of a widespread monodominant East African tree, Acacia drepanolobium was 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. 4. 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. 5. 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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    Relationships Between Cattle and Biodiversity in Multiuse Landscape Revealed by Kenya Long-Term Exclosure Experiment
    (Elsevier, 2018-05) Kimuyu, Duncan M.; Young, Truman P.; Porensky, Lauren M.; Riginos, Corinna; Veblen, Kari E.; Odadi, Wilfred O.; Charles, Grace K.; Young, Hillary S.
    On rangelands worldwide, cattle interact with many forms of biodiversity, most obviously with vegetation and other large herbivores. Since 1995, we have been manipulating the presence of cattle, mesoherbivores, and megaherbivores (elephants and giraffes) in a series of eighteen 4-ha (10-acre) plots at the Kenya Long-term Exclosure Experiment. We recently (2013) crossed these treatments with small-scale controlled burns. These replicated experimental treatments simulate different land management practices. We seek to disentangle the complex relationships between livestock and biodiversity in a biome where worldwide, uneasy coexistence is the norm. Here, we synthesize more than 20 yr of data to address three central questions about the potentially unique role of cattle in savanna ecology: 1) To what extent do cattle and wild herbivores compete with or facilitate each other? 2) Are the effects of cattle on vegetation similar to those of wildlife, or do cattle have unique effects? 3) What effects do cattle and commercial cattle management have on other savanna organisms? We found that 1) Cattle compete at least as strongly with browsers as grazers, and wildlife compete with cattle, although these negative effects are mitigated by cryptic herbivores (rodents), rainfall, fire, and elephants. 2) Cattle effects on herbaceous vegetation (composition, productivity) are similar to those of the rich mixture of ungulates they replace, differing mainly due to the greater densities of cattle. In contrast, cattle, wild mesoherbivores, and megaherbivores have strongly guild-specific effects on woody vegetation. 3) Both cattle and wild ungulates regulate cascades to other consumers, notably termites, rodents, and disease vectors (ticks and fleas) and pathogens. Overall, cattle management, at moderate stocking densities, can be compatible with the maintenance of considerable native biodiversity, although reducing livestock to these densities in African rangelands is a major challenge.
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    Fire disturbance disrupts an acacia ant–plant mutualism in favor of a subordinate ant species
    (Wiley and sons, 2017-03) Kimuyu, Duncan K.; Sensenig, Ryan L.; Guajardo, Juan C. Ruiz; Veblen, Kari E.; Riginos, Corinna; Young, Truman P.
    Although disturbance theory has been recognized as a useful framework in examining the stability of ant–plant mutualisms, very few studies have examined the effects of fire disturbance on these mutualisms. In myrmecophyte-dominated savannas, fire and herbivory are key drivers that could influence ant–plant mutualisms by causing complete colony mortality and/or decreasing colony size, which potentially could alter dominance hierarchies if subordinate species are more fire resilient. We used a large-scale, replicated fire experiment to examine long-term effects of fire on acacia–ant community composition. To determine if fire shifted ant occupancy from a competitive dominant to a subordinate ant species, we surveyed the acacia–ant community in 6–7 yr old burn sites and examined how the spatial scale of these burns influenced ant community responses. We then used two short-term fire experiments to explore possible mechanisms for the shifts in community patterns observed. Because survival of ant colonies is largelydependent on their ability to detect and escape an approaching fire, we first tested the evacuation response of all four ant species when exposed to smoke (fire signal). Then to better understand how fire and its interaction with large mammal herbivory affect the density of ants per tree, we quantified ant worker density in small prescribed burns within herbivore exclusion plots. We found clear evidence suggesting that fire disturbance favored the subordinate ant Crematogaster nigriceps more than the dominant and strong mutualist ant C. mimosae, whereby C. nigriceps (1) was the only species to occupy a greater proportion of trees in 6–7 yr old burn sites compared to unburned sites, (2) had higher burn/unburn tree ratios with increasing burn size, and (3) evacuated significantly faster than C. mimosae in the presence of smoke. Fire and herbivory had opposite effects on ant density per meter of branch for both C. nigriceps and C. mimosae, with fire decreasing ant densities per meter of branch and the presence of large herbivores increasing ant density. Taken together, these experiments suggest that major ecosystem disturbances like fire can disrupt mutualistic associations and maintain diversity in partner qualityand identitydespite a clear dominance hierarchy.
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    Fire disturbance disrupts an acacia ant–plant mutualism in favor of a subordinate ant species
    (EcologicalSocietyof America, 2017) Kimuyu, Duncan K.; Sensenig, Ryan L.; Guajardo, Juan C. Ruiz; Veblen, Kari E.; Riginos, Corinna; YOUNG, Truman P.
    Although disturbance theory has been recognized as a useful framework in examining the stability of ant–plant mutualisms, very few studies have examined the effects of fire disturbance on these mutualisms. In myrmecophyte-dominated savannas, fire and herbivory are key drivers that could influence ant–plant mutualisms by causing complete colony mortality and/or decreasing colony size, which potentially could alter dominance hierarchies if subordinate species are more fire resilient. We used a large-scale, replicated fire experiment to examine long-term effects of fire on acacia–ant community composition. To determine if fire shifted ant occupancy from a competitive dominant to a subordinate ant species, we surveyed the acacia–ant community in 6–7 yr old burn sites and examined how the spatial scale of these burns influenced ant community responses. We then used two short-term fire experiments to explore possible mechanisms for the shifts in community patterns observed. Because survival of ant colonies is largelydependent on their ability to detect and escape an approaching fire, we first tested the evacuation response of all four ant species when exposed to smoke (fire signal). Then to better understand how fire and its interaction with large mammal herbivory affect the density of ants per tree, we quantified ant worker density in small prescribed burns within herbivore exclusion plots. We found clear evidence suggesting that fire disturbance favored the subordinate ant Crematogaster nigriceps more than the dominant and strong mutualist ant C. mimosae, whereby C. nigriceps (1) was the only species to occupy a greater proportion of trees in 6–7 yr old burn sites compared to unburned sites, (2) had higher burn/unburn tree ratios with increasing burn size, and (3) evacuated significantly faster than C. mimosae in the presence of smoke. Fire and herbivory had opposite effects on ant density per meter of branch for both C. nigriceps and C. mimosae, with fire decreasing ant densities per meter of branch and the presence of large herbivores increasing ant density. Taken together, these experiments suggest that major ecosystem disturbances like fire can disrupt mutualistic associations and maintain diversity in partner qualityand identitydespite a clear dominance hierarchy
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