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The metamicrobiome is an integrated concept to study carbon and nutrient recycling in ecosystems. Decomposition of plant-derived matter by free-living microbes and fire – two key recycling pathways – are highly sensitive to global change. Mutualistic associations of microbes with plants and animals strongly reduce this sensitivity. By solving a fundamental allometric trade-off between metabolic and homeostatic capacity, these mutualisms enable continued recycling of plant matter where and when conditions are unfavourable for the free-living microbiome. A diverse metamicrobiome – where multiple plant- and animal-associated microbiomes complement the free-living microbiome – thus enhances homeostasis of ecosystem recycling rates in variable environments. Research into metamicrobiome structure and functioning in ecosystems is therefore important for progress towards understanding environmental change.

Variation in defense traits likely depends on access to different resources and risk from herbivory. Plant defense theories have predicted both positive and negative associations between defense traits and access to resources, but relatively few studies have explored intraspecific variation in defense traits along multiple resource and mammalian herbivory risk gradients. We assessed relationships between herbivory intensity, multiple resources, and plant defense traits using a widely distributed tropical savanna herb,Solanum incanum. As independent measures of risk from large mammal herbivores are rare, we used a satellite-based vegetation index to predict herbivory intensity at the landscape scale. We found that the satellite-based estimate of herbivory intensity was positively associated with browser abundance and total soil P, but negatively associated with rainfall. Intraspecific defense traits too varied substantially across sites (n=43) but only variation in spine density was associated with herbivory intensity and plant resources, such that spine density was positively associated with both rainfall and soil P, but bimodally associated with herbivory intensity. Taken together, it suggests that defenses maybe favored either where resources for defense are abundant under low but still present risk (i.e, at high rainfall sites) or where resource-expensive plant tissue is at high risk (i.e, at high soil P sites). This hints at the possibility of a shift from a resource-associated (bottom-up) to an herbivory-associated (top-down) control of allocation to defenses along an environmental gradient. Additionally, the independent effect of soil P on a carbon-based defense, spine density, suggests potential for resources that are not components of defenses to also influence allocation to defense traits. Thus, our study provides evidence for the influence of multiple drivers, resources, and herbivory intensity, on anti-herbivore defenses and their shifting relative importance on allocation to defenses along an environmental gradient.

International audience; Regular vegetation patterns have been predicted to indicate a system slowing down and possibly desertification of drylands. However, these predictions have not yet been observed in dryland vegetation due to the inherent logistic difficulty to gather longerterm in situ data. Here, we evaluate the theoretical prediction that regular vegetation patterns are associated with empirically derived temporal indicators (autocorrelation, variance, responsiveness) of critical slowing down in a dryland ecosystem in Sudan using different remote sensing products. We use recently developed methods using remote-sensing EVI time-series in combination with classified regular vegetation patterns along a rainfall gradient in Sudan to test the predicted slowing down. We tested our empirical findings against theoretical predictions from a stochastic version of a spatial explicit model that has been used to describe vegetation dynamics in drylands under aridity stress. Overall, three temporal indicators (responsiveness, temporal autocorrelation, variance) show slowing down as vegetation patterns change from gaps to labyrinths to spots towards more arid conditions, confirming predictions. However, this transition exhibits non-linearities, specifically when patterns change configuration. Model simulations reveal that the transition between patterns temporarily slows down the system affecting the temporal indicators. These transient states when vegetation patterns reorganize thus affect the systems resilience indicators in a non-linear way. Our findings suggest that spatial self-organization of dryland vegetation is associated with critical slowing down, but this transition towards reduced resilience happens in a non-linear way. Future work should aim to better understand transient dynamics in regular vegetation patterns in dryland ecosystems, because long transients make regular vegetation patterns of limited use for management in anticipating critical transitions.

In the Greater Serengeti-Mara ecosystem, with the Serengeti National Park (SNP) at its core, people and wildlife are strongly dependent on water supply that has a strong seasonal and inter-annual variability. The Mara River, the only perennial river in SNP, and a number of small streams originate from outside SNP before flowing through it. In those watersheds increasing grazing pressure from livestock, deforestation, irrigation and other land uses affect river flows in SNP that subsequently have impacts on wildlife. We quantified the changes since the 1970s of river discharge dynamics. We found that the baseflow recession period for the Mbalageti River has remained unchanged at 70 days, which is a natural system inside SNP. By contrast it has decreased from 100 days in the 1970s to 16 days at present for the Mara River, coinciding with increased commercial-scale irrigation in Kenya that extract Mara River water before it reaches SNP. This irrigation will result in zero flow in the river in SNP if the proposed dams in the river in Kenya are built. We observed high flash floods and prolonged periods of zero flows in streams draining livestock grazed watersheds, where severe major erosion prevails that results in gully formation. This eroded sediment is expected to silt and dry out the scattered dry season water holes in SNP, which are an important source of drinkable water for wildlife during the dry season. It appears likely that the future water supply of SNP is at risk, and this has major consequences for its people and wildlife. Ecohydrology-based solutions at the catchment scale are urgently needed to reduce catchment degradation while ensuring sustainable water provision.

Global warming compels larger endothermic animals to adapt either physiologically or behaviourally to avoid thermal stress, especially in tropical ecosystems. Their adaptive responses may however be compromised by other constraints, such as predation risk or starvation. Using an exceptional camera-trap dataset spanning 32 protected areas across southern Africa, we find that intermediate-sized herbivores (100–550 kg) switch activity to hotter times of the day when exposed to predation by lions. These herbivores face a tight window for foraging activity being exposed to nocturnal predation and to heat during the day, suggesting a trade-off between predation risk and thermoregulation mediated by body size. These findings stress the importance of incorporating trophic interactions into climate change predictions. Rising temperatures and predator avoidance constrain herbivore activity on the southern African savannas, forcing them into ever-tightening windows of activity, in a ‘timescape of fear’.

Climate change predictions indicate that summer droughts will become more severe and frequent. Yet, the impact of soil communities on the response of plant communities to drought remains unclear. Here, we report the results of a novel field experiment, in which we manipulated soil communities by adding soil inocula originating from different successional stages of coastal dune ecosystems to a plant community established from seeds on bare dune sand. We tested if and how the added soil biota from later-successional ecosystems influenced the sensitivity (resistance and recovery) of plant communities to drought. In contrast to our expectations, soil biota from later-successional soil inocula did not improve the resistance and recovery of plant communities subjected to drought. Instead, inoculation with soil biota from later successional stages reduced the post-drought recovery of plant communities, suggesting that competition for limited nutrients between plant community and soil biota may exacerbate the post-drought recovery of plant communities. Moreover, soil pathogens present in later-successional soil inocula may have impeded plant growth after drought. Soil inocula had differential impacts on the drought sensitivity of specific plant functional groups and individual species. However, the sensitivity of individual species and functional groups to drought was idiosyncratic and did not explain the overall composition of the plant community. Based on the field experimental evidence, our results highlight the adverse role soil biota can play on plant community responses to environmental stresses. These outcomes indicate that impacts of soil biota on the stability of plant communities subjected to drought are highly context-dependent and suggest that in some cases the soil biota activity can even destabilize plant community biomass responses to drought.

Regular vegetation patterns have been predicted to indicate a system slowing down and possibly desertification of drylands. However, these predictions have not yet been observed in dryland vegetation due to the inherent logistic difficulty to gather longer-term in situ data. Here, we use recently developed methods using remote-sensing EVI time-series in combination with classified regular vegetation patterns along a rainfall gradient in Sudan to test these predictions. Overall, three temporal indicators (responsiveness, temporal autocorrelation, variance) show slowing down as vegetation patterns change from gaps to labyrinths to spots towards more arid conditions, confirming predictions. However, this transition exhibits non-linearities, specifically when patterns change configuration. Model simulations reveal that the transition between patterns temporarily slows down the system affecting the temporal indicators. These transient states when vegetation patterns reorganize thus affect the systems resilience indicators in a non-linear way. Our findings suggest that spatial self-organization of dryland vegetation is associated with critical slowing down, but this transition towards reduced resilience happens in a non-linear way. Future work should aim to better understand transient dynamics in regular vegetation patterns in dryland ecosystems, because long transients make regular vegetation patterns of limited use for management in anticipating critical transitions.

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.

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.

1.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. This article is protected by copyright. All rights reserved.

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.

How is woody vegetation patchiness affected by rainfall, fire and large herbivore biomass? Can we predict woody patchiness and cover over large-scale environmental gradients? We quantified variation in local patchiness as the lacunarity of woody cover on satellite-derived images. Using Random Forest regression we analysed how both average woody cover and its patchiness depend on annual rainfall, fire frequency and grazer and browser metabolic biomass densities. Fire frequency and rainfall were the clearest predictors, whereas effects of large herbivores on woody vegetation were smaller and more complex. Under low rainfall conditions (500 mm·yr−1) trees had less total cover and were more regularly spaced across the landscape (lower patchiness). Woody cover and vegetation patchiness increased with rainfall whereas fire frequency decreased woody cover and further increased patchiness. These results suggest a switch from competition between neighbouring trees under low rainfall conditions towards benefits among neighbours with increasing fire frequencies. Whereas overall woody cover and its patchiness are two independent aspects of savanna woody vegetation patterns, both need to be investigated to obtain a good understanding of the functioning and diversity of savanna ecosystems.

Global warming compels larger endothermic animals to adapt either physiologically or behaviourally to avoid thermal stress, especially in tropical ecosystems. Their adaptive responses may however be compromised by other constraints, such as predation risk or starvation. Using an exceptional camera-trap dataset spanning 32 protected areas across southern Africa, we find that intermediate-sized herbivores (100-550 kg) switch activity to hotter times of the day when exposed to predation by lions. These herbivores face a tight window for foraging activity being exposed to nocturnal predation and to heat during the day, suggesting a trade-off between predation risk and thermoregulation mediated by body size. These findings stress the importance of incorporating trophic interactions into climate change predictions.

Nutrient availability in terrestrial ecosystems has been found to vary along regional climatic and soil gradients and drive variation in plant community composition and vegetation structure. However, more local biotic feedbacks also affect nutrient availability, but their importance in determining vegetation structure relative to regional drivers is yet unclear. Mesic African savannas form a transition zone between the dry grasslands with relatively low nitrogen availability (indicated by low plant N:P ratios) and the wet woodlands of the continent characterized by relatively low phosphorous availability (high plant N:P ratios). They host strong feedback mechanisms of both vegetation and consumers, where large grazers can create short grazing lawns that alternate over short distances with tall fire-dominated bunch grasslands, and patches dominated by woody species with abundant macrodetritivores. Here, we test if such local biotic interactions can overrule regional, landscape-level drivers of plant nutrient availability. In a South African savanna, we find that plant N:P ratios, N and P resorption efficiencies and proficiencies, were indeed much stronger affected by small-scale biotic-created heterogeneity than by a regional rainfall gradient (530-830mmyr(-1)). Furthermore, we observe that differences in N:P ratios between vegetation structural types are not caused by simple differences in inherent plant traits. This suggests that availability of N and P to plants is strongly contingent on the local biotic interplay between different vegetation structural types and within-system feedbacks linked to each vegetation type. The regional rainfall gradient did affect nutrient availability, but mainly through its effect on the relative distribution of the different vegetation structural types thus setting the range of possible outcomes of biotic interactions.Synthesis. Regional and global studies that model savanna carbon and nutrient cycling only on the basis of regional gradients in soil and climatic conditions may insufficiently capture the dominant ecosystem processes involved.

International audience; Ecosystems comprise flows of energy and materials, structured by organisms and their interactions. Important generalizations have emerged in recent decades about conversions by organisms of energy (metabolic theory of ecology) and materials (ecological stoichiometry). However, these new insights leave a key question about ecosystems inadequately addressed: are there basic organizational principles that explain how the interaction structure among species in ecosystems arises? Here we integrate recent contributions to the understanding of how ecosystem organization emerges through ecological autocatalysis (EA), in which species mutually benefit through self-reinforcing circular interaction structures. We seek to generalize the concept of EA by integrating principles from community and ecosystem ecology. We discuss evidence suggesting that ecological autocatalysis is facilitated by resource competition and natural selection, both central principles in community ecology. Furthermore, we suggest that pre-emptive resource competition by consumers and plant resource diversity drive the emergence of autocatalytic loops at the ecosystem level. Subsequently, we describe how interactions between such autocatalytic loops can explain pattern and processes observed at the ecosystem scale, and summarize efforts to model different aspect of the phenomenon. We conclude that EA is a central principle that forms the backbone of the organization in systems ecology, analogous to autocatalytic loops in systems chemistry.

1. 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 insavannas, we suggest that water balance-based processes are additionally important during the dryseason.

Threats to the Serengeti Protected areas are an important tool for conserving biodiversity and ecosystem functioning. But how well do these areas withstand pressure from human activity in surrounding landscapes? Veldhuis et al. studied long-term data from the Serengeti-Mara ecosystem in East Africa. Human activities at boundary regions cause animals to concentrate in the core of the protected area, which eventually reduces soil carbon storage and nitrogen fixation rates and increases vulnerability to extreme droughts. Similar patterns are likely for many, if not all, large protected areas. Science , this issue p. 1424

The grass layer of African savannas consists of two main vegetation types: grazing lawns, dominated by short, mostly clonally reproducing grasses, and bunch grasslands, dominated by tall bunch grasses. This patchy distribution of vegetation types is mostly created by large herbivores, which selectively feed on the more nutritious lawn grass species. Besides grazing, herbivores trample the soil, thereby causing soil compaction, with possible consequences for water infiltration. This raises two questions: (i) is water more limiting in grazing lawns than in bunch grasslands and (ii) are lawn grasses more drought tolerant than bunch grasses? To study these questions, we compared drought conditions in both lawn and bunch grasslands in a South African savanna. Additionally, in a climate room, we compared the performance of three lawn and three bunch grass species under a control and a water limitation treatment. Thirdly, we investigated whether there are differences between lawn and bunch grasses in traits related to drought tolerance. Our results show that despite large differences in water availability in the field, lawn and bunch grasses did not differ in their growth response to drought. Drought reduced growth of both growth forms equally. However, we found strong intrinsic trait differences between growth forms, with lawn grasses having higher specific root length and relative growth rate and bunch grasses having a higher root:shoot ratio. These results suggest that after drought-induced plant death, lawn grasses might be more capable of recolonizing patches of bare soil.

Savanna grasslands are characterized by high spatial heterogeneity in vegetation structure, aboveground biomass and nutritional quality, with high quality short-grass grazing lawns forming mosaics with patches of tall bunch grasses of lower quality. This heterogeneity can arise because of local differences in consumption, because of differences in productivity, or because both processes enforce each other (more production and consumption). However, the relative importance of both processes in maintaining mosaics of lawn and bunch grassland types has not been measured. Also their interplay been not been assessed across landscape gradients. In a South African savanna, we, therefore, measured the seasonal changes in primary production, nutritional quality and herbivore consumption (amount and percentage) of grazing lawns and adjacent bunch grass patches across a rainfall gradient. We found both higher amounts of primary production and, to a smaller extent, consumption for bunch grass patches. In addition, for bunch grasses primary production increased towards higher rainfall while foliar nitrogen concentrations decreased. Foliar nitrogen concentrations of lawn grasses decreased much less with increasing rainfall. Consequently, large herbivores targeted the biomass produced on grazing lawns with on average 75 % of the produced biomass consumed. We conclude that heterogeneity in vegetation structure in this savanna ecosystem is better explained by small-scale differences in productivity between lawn and bunch grass vegetation types than by local differences in consumption rates. Nevertheless, the high nutritional quality of grazing lawns is highly attractive and, therefore, important for the maintenance of the heterogeneity in species composition (i.e. grazing lawn maintenance). Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3698-y) contains supplementary material, which is available to authorized users.