Grazing herbivores reduce herbaceous biomass and fire activity across African savannas.

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/km2 of 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.

Determinants of microbiome composition: Insights from free-ranging hybrid zebras (Equus quagga × grevyi).

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 the trnL-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.

Environmental context shapes the relationship between grass consumption and body size in African herbivore communities.

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.

Impacts of large herbivores on terrestrial ecosystems.

Large herbivores play unique ecological roles and are disproportionately imperiled by human activity. As many wild populations dwindle towards extinction, and as interest grows in restoring lost biodiversity, research on large herbivores and their ecological impacts has intensified. Yet, results are often conflicting or contingent on local conditions, and new findings have challenged conventional wisdom, making it hard to discern general principles. Here, we review what is known about the ecosystem impacts of large herbivores globally, identify key uncertainties, and suggest priorities to guide research. Many findings are generalizable across ecosystems: large herbivores consistently exert top-down control of plant demography, species composition, and biomass, thereby suppressing fires and the abundance of smaller animals. Other general patterns do not have clearly defined impacts: large herbivores respond to predation risk but the strength of trophic cascades is variable; large herbivores move vast quantities of seeds and nutrients but with poorly understood effects on vegetation and biogeochemistry. Questions of the greatest relevance for conservation and management are among the least certain, including effects on carbon storage and other ecosystem functions and the ability to predict outcomes of extinctions and reintroductions. A unifying theme is the role of body size in regulating ecological impact. Small herbivores cannot fully substitute for large ones, and large-herbivore species are not functionally redundant - losing any, especially the largest, will alter net impact, helping to explain why livestock are poor surrogates for wild species. We advocate leveraging a broad spectrum of techniques to mechanistically explain how large-herbivore traits and environmental context interactively govern the ecological impacts of these animals.

Dynamic landscapes of fear: understanding spatiotemporal risk.

The landscape of fear (LOF) concept posits that prey navigate spatial heterogeneity in perceived predation risk, balancing risk mitigation against other activities necessary for survival and reproduction. These proactive behavioral responses to risk can affect individual fitness, population dynamics, species interactions, and coexistence. Yet, antipredator responses in free-ranging prey often contradict expectations, raising questions about the generality and scalability of the LOF framework and suggesting that a purely spatial, static LOF conceptualization may be inadequate. Here, we outline a 'dynamic' LOF framework that explicitly incorporates time to account for predictable spatiotemporal variation in risk-resource trade-offs. This integrated approach suggests novel predictions about predator effects on prey behaviors to refine understanding of the role predators play in ecological communities.

Evolutionary causes and consequences of ungulate migration.

Ungulate migrations are crucial for maintaining abundant populations and functional ecosystems. However, little is known about how or why migratory behaviour evolved in ungulates. To investigate the evolutionary origins of ungulate migration, we employed phylogenetic path analysis using a comprehensive species-level phylogeny of mammals. We found that 95 of 207 extant ungulate species are at least partially migratory, with migratory behaviour originating independently in 17 lineages. The evolution of migratory behaviour is associated with reliance on grass forage and living at higher latitudes wherein seasonal resource waves are most prevalent. Indeed, originations coincide with mid-Miocene cooling and the subsequent rise of C4 grasslands. Also, evolving migratory behaviour supported the evolution of larger bodies, allowing ungulates to exploit new ecological space. Reconstructions of migratory behaviour further revealed that seven of ten recently extinct species were probably migratory, suggesting that contemporary migrations are important models for understanding the ecology of the past.

Seasonal strategies differ between tropical and extratropical herbivores.

Seasonal diet shifts and migration are key components of large herbivore population dynamics, but we lack a systematic understanding of how these behaviours are distributed on a macroecological scale. The prevalence of seasonal strategies is likely related to herbivore body size and feeding guild, and may also be influenced by properties of the environment, such as soil nutrient availability and climate seasonality. We evaluated the distribution of seasonal dietary shifts and migration across large-bodied mammalian herbivores and determined how these behaviours related to diet, body size and environment. We found that herbivore strategies were consistently correlated with their traits: seasonal diet shifts were most prevalent among mixed feeding herbivores and migration among grazers and larger herbivores. Seasonality also played a role, particularly for migration, which was more common at higher latitudes. Both dietary shifts and migration were more widespread among extratropical herbivores, which also exhibited more intermediate diets and body sizes. Our findings suggest that strong seasonality in extratropical systems imposes pressure on herbivores, necessitating widespread behavioural responses to navigate seasonal resource bottlenecks. It follows that tropical and extratropical herbivores may have divergent responses to global change, with intensifying herbivore pressure in extratropical systems contrasting with diminishing herbivore pressure in tropical systems.

Elevated wildlife-vehicle collision rates during the COVID-19 pandemic.

Wildlife-vehicle collisions threaten both humans and wildlife, but we still lack information about the relationship between traffic volume and wildlife-vehicle collisions. The COVID-19 pandemic allowed us to investigate the effects of traffic volume on wildlife-vehicle collisions in the United States. We observed decreased traffic nationwide, particularly in densely populated states with low or high disease burdens. Despite reduced traffic, total collisions were unchanged; wildlife-vehicle collisions did decline at the start of the pandemic, but increased as the pandemic progressed, ultimately exceeding collisions in the previous year. As a result, nationwide collision rates were higher during the pandemic. We suggest that increased wildlife road use offsets the effects of decreased traffic volume on wildlife-vehicle collisions. Thus, decreased traffic volume will not always reduce wildlife-vehicle collisions.

Heterogeneity in African savanna elephant distributions and their impacts on trees in Kruger National Park, South Africa.

Though elephants are a major cause of savanna tree mortality and threaten vulnerable tree species, managing their impact remains difficult, in part because relatively little is known about how elephant impacts are distributed throughout space.This is exacerbated by uncertainty about what determines the distribution of elephants themselves, as well as whether the distribution of elephants is even informative for understanding the distribution of their impacts.To better understand the factors that underlie elephant impacts, we modeled elephant distributions and their damage to trees with respect to soil properties, water availability, and vegetation in Kruger National Park, South Africa, using structural equation modeling.We found that bull elephants and mixed herds differed markedly in their distributions, with bull elephants concentrating in sparsely treed basaltic sites close to artificial waterholes and mixed herds aggregating around permanent rivers, particularly in areas with little grass.Surprisingly, we also found that the distribution of elephant impacts, while highly heterogeneous, was largely unrelated to the distribution of elephants themselves, with damage concentrated instead in densely treed areas and particularly on basaltic soils.Results underscore the importance of surface water for elephants but suggest that elephant water dependence operates together with other landscape factors, particularly vegetation community composition and historical management interventions, to influence elephant distributions.

Public parks and the pandemic: How park usage has been affected by COVID-19 policies.

Public parks serve an important societal function as recreational spaces for diverse communities of people, with well documented physical and mental health benefits. As such, parks may be crucial for how people have handled effects of the COVID-19 pandemic, particularly the increasingly limited recreational opportunities, widespread financial uncertainty, and consequent heightened anxiety. Despite the documented benefits of parks, however, many states have instituted park shutdown orders due to fears that public parks could facilitate SARS-CoV-2 transmission. Here we use geotagged social media data from state, county, and local parks throughout New Jersey to examine whether park visitation increased when the COVID-19 pandemic began and whether park shutdown orders were effective at deterring park usage. We compare park usage during four discrete stages of spring 2020: (1) before the pandemic began, (2) during the beginning of the pandemic, (3) during the New Jersey governor's state-wide park shutdown order, and (4) following the lifting of the shutdown. We find that park visitation increased by 63.4% with the onset of the pandemic. The subsequent park shutdown order caused visitation in closed parks to decline by 76.1% while parks that remained open continued to experience elevated visitation levels. Visitation then returned to elevated pre-shutdown levels when closed parks were allowed to reopen. Altogether, our results indicate that parks continue to provide crucial services to society, particularly in stressful times when opportunities for recreation are limited. Furthermore, our results suggest that policies targeting human behavior can be effective and are largely reversible. As such, we should continue to invest in public parks and to explore the role of parks in managing public health and psychological well-being.

Drought-response strategies of savanna herbivores.

Climate models predict increases in drought frequency and severity worldwide, with potential impacts on diverse systems, including African savannas. These droughts pose a concern for the conservation of savanna mammal communities, such that understanding how different species respond to drought is vital.Because grass decreases so consistently during droughts, we predict that grass-dependent species (grazers and mixed feeders) will respond strongly to drought, whether by changing diets, seeking drought refugia, or suffering mortality.A recent severe but heterogeneous drought in Kruger National Park, South Africa, afforded a rare opportunity to test these hypotheses in situ-crucial, given the central role of landscape-scale movement as a potential herbivore strategy. We used herbivore dung as a proxy, integrating spatial distributions (dung counts) with diet composition (carbon isotope analysis of dung).As predicted, browsers showed little response to drought. However, mixed feeders switched their diets to incorporate more C3 trees/forbs, but did not move. Meanwhile, grazers and megaherbivores instead moved toward drought refugia. Synthesis and applications: The responses we observed by savanna herbivores are largely amplifications of typical dry season strategies and reflect constraints imposed by body size and feeding ecology. Grazers may be at particular risk from increased drought frequency and spatial extent if drought refugia become decreasingly available. Conservation strategies should recognize these constraints and work to facilitate the diverse responses of herbivores to drought.