Protected areas provide major benefits for humans in the form of ecosystem services, but landscape degradation by human activity at their edges may compromise their ecological functioning. Using multiple lines of evidence from 40 years of research in the Serengeti-Mara ecosystem, we find that such edge degradation has effectively “squeezed” wildlife into the core protected area and has altered the ecosystem’s dynamics even within this 40,000-square-kilometer ecosystem. This spatial cascade reduced resilience in the core and was mediated by the movement of grazers, which reduced grass fuel and fires, weakened the capacity of soils to sequester nutrients and carbon, and decreased the responsiveness of primary production to rainfall. Similar effects in other protected ecosystems worldwide may require rethinking of natural resource management outside protected areas.

Summary1. Large herbivores play a key role in creating spatial heterogeneity through the formation of grazing lawns. Recent research suggests that the currently accepted nutrient-based theory on the formation of these grazing lawns cannot universally explain their formation in all ecosystems where they are found. 2. We developed and investigated an alternative hypothesis on grazing lawn formation and maintenance based on herbivore effects on the plant-soil water balance. We propose that large herbivores change the soil water balance in grazing lawns through defoliation and soil compaction, causing a shift in vegetation composition towards a drought-tolerant plant community. 3. Investigating this idea in a tropical savanna, we indeed found profound differences in grazing lawn soil properties and water balance. In particular, defoliation increased soil temperatures and potential evaporation rates while soil compaction increased bulk density and decreased water infiltration rates, especially on fine-textured soils. Soil moisture was therefore generally much lower in grazing lawns than in adjacent bunch grass areas. 4. Furthermore, we found that grazing lawn species show drought-tolerant traits, with higher leaf sodium levels, suggesting evolutionary adaptation to these herbivore-induced dry conditions. However, leaf water potentials did not differ between grazing lawn and bunch grass species. 5. Synthesis. This study shows that large herbivores might form grazing lawns through previously underestimated effects on water balance. Thus, future studies on large herbivore effects on vegetation should increasingly focus on additional pathways of soil compaction and defoliation. While nutrient-based processes driving grazing lawn formation may operate during the wet season in savannas, we suggest that water balance-based processes are additionally important during the dry season.

It is widely recognized that the release of nutrients by herbivores via their waste products strongly impacts nutrient availability for autotrophs. The ratios of nitrogen (N) and phosphorus (P) recycled through herbivore release (i.e., waste N:P) are mainly determined by the stoichiometric composition of the herbivore's food (food N:P) and its body nutrient content (body N:P). Waste N:P can in turn impact autotroph nutrient limitation and productivity. Herbivore-driven nutrient recycling based on stoichiometric principles is dominated by theoretical and experimental research in freshwater systems, in particular interactions between algae and invertebrate herbivores. In terrestrial ecosystems, the impact of herbivores on nutrient cycling and availability is often limited to studying carbon (C):N and C:P ratios, while the role of terrestrial herbivores in mediating N:P ratios is also likely to influence herbivore-driven nutrient recycling. In this review, we use rules and predictions on the stoichiometry of nutrient release originating from algal-based aquatic systems to identify the factors that determine the stoichiometry of nutrient release by herbivores. We then explore how these rules can be used to understand the stoichiometry of nutrient release by terrestrial herbivores, ranging from invertebrates to mammals, and its impact on plant nutrient limitation and productivity. Future studies should focus on measuring both N and P when investigating herbivore-driven nutrient recycling in terrestrial ecosystems, while also taking the form of waste product (urine or feces) and other pathways by which herbivores change nutrients into account, to be able to quantify the impact of waste stoichiometry on plant communities.

Life sciences study designAll studies must disclose on these points even when the disclosure is negative. Sample sizeNo sample-size calculation was performed. We used camera trap data from 32 different protected areas in which about half lions were present. This is far beyond the sample sizes used in any study so far on herbivore behavior in response to predation risk. To calculate activitity patterns for each herbivore species, we set the sample size threshold at 100 captures per species per survey to increase the robustness of our estimates. Decreasing this threshold did not qualitatively change our results.Data exclusions We set the sample size threshold at 100 captures per species per survey to increase the robustness of our estimates. Decreasing this threshold did not qualitatively change our results. Thus, species with less than 100 captures were excluded. ReplicationThe design included replication of predator presence and absence, with 16 areas (replications) of lion presence and 16 areas (replications) of lion absence.Randomization Allocation of predator or herbivore presence was not done randomly, but based on natural occurence of herbivores and reintroduction of lions in protected area across South Africa. Nevertheless, this comes very close to a "natural experiment" as lions used to be present in all 32 protected areas and there is no bias in environmental characteristics (temperature, rainfall) between areas with and without lions present. BlindingData collection was done via camera traps so this could be seen as "blind" data collection for the purpose of this study. Reporting for specific materials, systems and methodsWe require information from authors about some types of materials, experimental systems and methods used in many studies. Here, indicate whether each material, system or method listed is relevant to your study. If you are not sure if a list item applies to your research, read the appropriate section before selecting a response.

The ecological impact of rapid environmental change will depend on the resistance of key ecosystems processes, which may be promoted by species that exert strong control over local environmental conditions. Recent theoretical work suggests that macrodetritivores increase the resistance of African savanna ecosystems to changing climatic conditions, but experimental evidence is lacking. We examined the effect of large fungus-growing termites and other non-fungus-growing macrodetritivores on decomposition rates empirically with strong spatiotemporal variability in rainfall and temperature. Non-fungus-growing larger macrodetritivores (earthworms, woodlice, millipedes) promoted decomposition rates relative to microbes and small soil fauna (+34%) but both groups reduced their activities with decreasing rainfall. However, fungus-growing termites increased decomposition rates strongest (+123%) under the most water-limited conditions, making overall decomposition rates mostly independent from rainfall. We conclude that fungus-growing termites are of special importance in decoupling decomposition rates from spatiotemporal variability in rainfall due to the buffered environment they create within their extended phenotype (mounds), that allows decomposition to continue when abiotic conditions outside are less favorable. This points at a wider class of possibly important ecological processes, where soil-plant-animal interactions decouple ecosystem processes from large-scale climatic gradients. This may strongly alter predictions from current climate change models.

The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land‐use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two‐dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water‐dependent grazers as wildebeest, zebra and buffalo–dominant species in savanna ecosystems – to be replaced by smaller, less water‐dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two‐dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change.

Citation: Kihwele, E. S., V. Mchomvu, N. Owen-Smith, R. S. Hetem, M. C. Hutchinson, A. B. Potter, H. Olff, and M. P. Veldhuis. 2020. Quantifying water requirements of African ungulates through a combination of functional traits. Ecological Monographs 90(2):Abstract. Climate and land use change modify surface water availability in African savannas. Surface water is a key resource for both wildlife and livestock and its spatial and temporal distribution is important for understanding the composition of large herbivore assemblages in savannas. Yet, the extent to which ungulate species differ in their water requirements remains poorly quantified. Here, we infer the water requirements of 48 African ungulates by combining six different functional traits related to physiological adaptations to reduce water loss, namely minimum dung moisture, relative dung pellet size, relative surface area of the distal colon, urine osmolality, relative medullary thickness, and evaporation rate. In addition, we investigated how these differences in water requirements relate to differences in dietary water intake. We observed strong correlations between traits related to water loss through dung, urine and evaporation, suggesting that ungulates minimize water loss through multiple pathways simultaneously, which suggests that each trait can thus be used independently to predict water requirements. Furthermore, we found that browsers and grazers had similar water requirements, but browsers are expected to be less dependent on surface water because they acquire more water through their diet. We conclude that these key functional traits are a useful way to determine differences in water requirements and an important tool for predicting changes in herbivore community assembly resulting from changes in surface water availability.

Abstract1. Territorial or resting behaviour of large herbivores can cause strong local deposits of dung, in different places than where they graze. Additionally, dung beetles and other macrodetritivores can subsequently affect local nutrient budgets through post-depositional re-dispersion of dung and accompanying nutrients. Such horizontal displacement of nutrients by animals has strong implications for savanna ecosystem functioning, but remains poorly studied as it is notoriously difficult to accurately map these flows and incredibly time-consuming.2. In an African savanna, with alternating patches of lawn, bunch grasses and trees/ shrubs, we undertook such effort and studied nutrient aggregation and redistribution by different large herbivore functional groups and dung beetles for a full growing season. We used movable cages to quantify herbivore consumption rates and measured nutrient return through biweekly dung counts. Furthermore, we estimated the offtake of nitrogen (N) and phosphorus (P) by the dominant megagrazer (white rhinoceros) to middens (dung deposition hotspots). Last, we experimentally measured the removal amount and movement paths of telocoprid dung beetles to quantify their nutrient redistribution effects.3. Our estimates suggest white rhinoceros to cause a large export of nutrients from grazing areas towards middens resulting in negative nutrient budgets for both lawn and bunch grassland types. Mesograzers (50-600 kg) realized a net nitrogen input towards high forage quality lawn vegetation at the expense of lower quality bunch grasslands. Browsers caused a net flow from trees/shrubs towards grassland patches.4. Interestingly, while the magnitude of our estimated flows of N consumption and return by large herbivores were rather similar, the P returns were about half of what has been consumed. This is in agreement with ecological stoichiometry theory that predicts that large herbivores should recycle more N than P, due to their relatively high P demand. Furthermore, dung-rolling beetles had a small, but significant, directed movement from lawn to bunch grassland vegetation.5. Synthesis. We conclude that within-ecosystem nutrient redistributions by animals are important and approximately of the same order of magnitude as regional atmospheric nutrient in and outputs (e.g. fire emissions, atmospheric N deposition, biological N fixation), and hence are important for understanding savanna ecosystem functioning.