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Abstract In Earth history, our understanding of how large‐bodied herbivores shape a variety of ecosystem processes is limited by the quality of paleoecological proxies for herbivore composition and abundance. Fecal stanols are lipids that can be produced by microbes within animal digestive systems and that could remedy this dearth of proxies. We used two multi‐decadal herbivore exclosures in Kruger National Park, South Africa, to constrain whether and how biomarker signatures preserve signals of herbivore abundance. Soil samples and dung counts were collected along transects across crests, mid‐slopes, and sodic sites inside and outside exclosures. Soils were analyzed for steroid (sterols and stanols) concentrations and distributions. We found that stanol concentrations were significantly greater in sodic soils outside exclosures, where herbivore dung densities were greatest. In contrast, sterol concentrations did not differ between treatments. Ratios of stanol isomers to sterols, which account for both compound degradation and source, increased strongly with herbivore dung counts. Finally, while herbivore species compositions influenced steroid distributions, total herbivore abundance was their strongest predictor. Further calibration is needed, but this work provides strong preliminary evidence that wild herbivore populations are quantitatively recorded by fecal biomarker distributions. 

Though herbivore grass dependence has been shown to increase with body size across herbivore species, it is unclear whether this relationship holds at the community level. Here we evaluate whether grass consumption scales positively with body size within African large mammalian herbivore communities and how this relationship varies with environmental context. We used stable carbon isotope and community occurrence data to investigate how grass dependence scales with body size within 23 savanna herbivore communities throughout eastern and central Africa. We found that dietary grass fraction increased with body size for the majority of herbivore communities considered, especially when complete community data were available. However, the slope of this relationship varied, and rainfall seasonality and elephant presence were key drivers of the variation—grass dependence increased less strongly with body size where rainfall was more seasonal and where elephants were present. We found also that the dependence of the herbivore community as a whole on grass peaked at intermediate woody cover. Intraspecific diet variation contributed to these community‐level patterns: common hippopotamus (Hippopotamus amphibius) and giraffe (Giraffa camelopardalis) ate less grass where rainfall was more seasonal, whereas Cape buffalo (Syncerus caffer) and savanna elephant (Loxodonta africana) grass consumption were parabolically related to woody cover. Our results indicate that general rules appear to govern herbivore community assembly, though some aspects of herbivore foraging behavior depend upon local environmental context. 

Abstract The composition of mammalian gut microbiomes is highly conserved within species, yet the mechanisms by which microbiome composition is transmitted and maintained within lineages of wild animals remain unclear. Mutually compatible hypotheses exist, including that microbiome fidelity results from inherited dietary habits, shared environmental exposure, morphophysiological filtering and/or maternal effects. Interspecific hybrids are a promising system in which to interrogate the determinants of microbiome composition because hybrids can decouple traits and processes that are otherwise co‐inherited in their parent species. We used a population of free‐living hybrid zebras (Equus quagga×grevyi) in Kenya to evaluate the roles of these four mechanisms in regulating microbiome composition. We analysed faecal DNA for both thetrnL‐P6 and the 16S rRNA V4 region to characterize the diets and microbiomes of the hybrid zebra and of their parent species, plains zebra (E. quagga) and Grevy's zebra (E. grevyi). We found that both diet and microbiome composition clustered by species, and that hybrid diets and microbiomes were largely nested within those of the maternal species, plains zebra. Hybrid microbiomes were less variable than those of either parent species where they co‐occurred. Diet and microbiome composition were strongly correlated, although the strength of this correlation varied between species. These patterns are most consistent with the maternal‐effects hypothesis, somewhat consistent with the diet hypothesis, and largely inconsistent with the environmental‐sourcing and morphophysiological‐filtering hypotheses. Maternal transmittance likely operates in conjunction with inherited feeding habits to conserve microbiome composition within species. 

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. 

1. Herbivory is a key process structuring vegetation in savannas, especially in Africa where large mammal herbivore communities remain intact. Exclusion experiments consistently show that herbivores impact savanna vegetation, but effect size variation has resisted explanation, limiting our understanding of the past, present and future roles of herbivory in savanna ecosystems. 2. Synthesis of vegetation responses to herbivore exclusion shows that herbivory decreased grass abundance by 57.0% and tree abundance by 30.6% across African savannas. 3. The magnitude of herbivore exclusion effects scaled with herbivore abundance: more grazing herbivores resulted in larger grass responses and more browsing herbivores in larger tree responses. However, existing experiments are concentrated in semi-arid savannas (400–800-mm rainfall) and soils data are mostly lack- ing, which makes disentangling environmental constraints a challenge and priority for future research. 4. Observed herbivore impacts were ~2.1× larger than existing estimates modelled based on consumption. Wildlife metabolic rates may be higher than are usually used for estimating consumption, which offers one clear avenue for reconciling estimated herbivore consumption with observed herbivore impacts. Plant-soil feedbacks, plant community composition, and the phenological or demographic timing of herbivory may also influence vegetation productivity, thereby magnify- ing herbivore impacts. 5. Because herbivore abundance so closely predicts vegetation impact, changes in herbivore abundance through time are likely predictive of the past and future of their impacts. Grazer diversity in Africa has declined from its peak 1 million years ago and wild grazer abundance has declined historically, suggesting that grazing likely had larger impacts in the past than it does today. 6. Current wildlife impacts are dominated by small-bodied mixed feeders, which will likely continue into the future, but the magnitude of top-down control may also depend on changing climate, fire and atmospheric CO2. 7. Synthesis. Herbivore biomass determines the magnitude of their impacts on savanna vegetation, with effect sizes based on direct observation that outstrip existing modelled estimates across African savannas. Findings suggest substantial ecosystem impacts of herbivory and allow us to generate evidence-based hypotheses of the past and future impacts of herbivores on savanna vegetation.