The importance of marshes providing soil stabilization to resist fast-flow erosion in case of a dike breach.

Salt marshes provide valuable ecosystem services including coastal protection by reducing wave loading on dikes and seawalls. If the topsoil is erosion resistant to fast-flowing water, it may also reduce breach depth if a dike fails. In this experiment, we quantified the topsoil erosion resistance from marshes and bare tidal flats with different soil types to understand the extent to which they can help reduce breach depth. Intact soil samples were collected from 11 locations in the Netherlands at different tidal elevations and then exposed for 3 h to 2.3 m/s currents. To the samples that remained stable after flow exposure, an artificial crack was made to test their stability following soil disturbance. All samples from the tidal flats were completely eroded, regardless of sediment type. In contrast, all samples from well-established marsh plateaus were stable as long as no disturbances were made, including those with sandy subsoils. After creating artificial cracks, samples with a thin cohesive top layer on top of sandy subsoil collapsed, while marshes with silty subsoils remained stable. Pioneer marshes on sandy substrate without a cohesive top layer were the only vegetated soils that completely eroded. The lower erosion of marshes with either sandy or silty soils compared to bare tidal flats was best explained by the presence of a top layer with belowground biomass, high organic content, high water content, and low bulk density. When analyzing the erodibility of marshes only, fine root density was the best predictor of erosion resistance. This study demonstrates the importance of preserving, restoring, or creating salt marshes, to obtain a topsoil that is erosion resistant under fast-flowing water, which helps reduce breach dimensions if a dike fails. The probability of topsoil erosion in established marshes with sandy subsoil is higher than in silty marshes. A silty layer of cohesive sediment on top of the sand provides extra erosion resistance as long as it does not break. Pioneer marshes that have not developed a cohesive top layer are erosion sensitive, especially in sandy soils. For future marsh creations, using fine-grained sediments or a mixture of sand with silt or clay is recommended.

Ontogenetic niche shifts as a driver of seasonal migration.

Ontogenetic niche shifts have helped to understand population dynamics. Here we show that ontogenetic niche shifts also offer an explanation, complementary to traditional concepts, as to why certain species show seasonal migration. We describe how demographic processes (survival, reproduction and migration) and associated ecological requirements of species may change with ontogenetic stage (juvenile, adult) and across the migratory range (breeding, non-breeding). We apply this concept to widely different species (dark-bellied brent geese (Branta b. bernicla), humpback whales (Megaptera novaeangliae) and migratory Pacific salmon (Oncorhynchus gorbuscha) to check the generality of this hypothesis. Consistent with the idea that ontogenetic niche shifts are an important driver of seasonal migration, we find that growth and survival of juvenile life stages profit most from ecological conditions that are specific to breeding areas. We suggest that matrix population modelling techniques are promising to detect the importance of the ontogenetic niche shifts in maintaining migratory strategies. As a proof of concept, we applied a first analysis to resident, partial migratory and fully migratory populations of barnacle geese (Branta leucopsis). We argue that recognition of the costs and benefits of migration, and how these vary with life stages, is important to understand and conserve migration under global environmental change.

Termites promote resistance of decomposition to spatiotemporal variability in rainfall.

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.

Herbivore exclusion promotes a more stochastic plant community assembly in a natural grassland.

Both bottom-up (e.g., nutrients) and top-down (e.g., herbivory) forces structure plant communities, but it remains unclear how they affect the relative importance of stochastic and deterministic processes in plant community assembly. Moreover, different-sized herbivores have been shown to have contrasting effects on community structure and function, but their effects on the processes governing community assembly (i.e., how they generate the impacts on structure) remain largely unknown. We evaluated the influence of bottom-up and top-down forces on the relative importance of deterministic and stochastic processes during plant community assembly. We used the data of a 7-yr factorial experiment manipulating nutrient availability (ambient and increased) and the presence of vertebrate herbivores (>1 kg) of different body size in a floodplain grassland in The Netherlands. We used a null model that describes a community composition expected by chance (i.e., stochastic assembly) and compared the plant community composition in the different treatments with this null model (the larger the difference, the more deterministically assembled). Our results showed that herbivore exclusion promoted a more stochastic plant community assembly, whereas increased nutrients played a relatively minor role in determining the relative importance of stochasticity in community assembly. Large herbivores facilitated intermediate-sized mammal herbivores, resulting in synergistic effects of enhanced grazing pressure and a more deterministic and convergent plant community assembly. We conclude that herbivores can act as strong deterministic forces during community assembly in natural systems. Our results also reveal that although large- and intermediate-sized mammal herbivores often have contrasting effects on many community and ecosystem properties, they can also synergistically homogenize plant communities.

Decoupled diversity dynamics in green and brown webs during primary succession in a saltmarsh.

Terrestrial ecosystems are characterized by a strong functional connection between the green (plant-herbivore-based) and brown (detritus-detritivore-based) parts of the food web, which both develop over successional time. However, the interlinked changes in green and brown food web diversity patterns in relation to key ecosystem processes are rarely studied. Here, we demonstrate changes in species richness, diversity and evenness over a wide range of invertebrate green and brown trophic groups during 100 years of primary succession in a saltmarsh ecosystem, using a well-calibrated chronosequence. We contrast two hypotheses on the relationship between green and brown food web diversity across succession: (i) 'coupled diversity hypothesis', which predicts that all trophic groups covary similarly with the main drivers of successional ecosystem assembly vs. (ii) the 'decoupled diversity hypothesis', where green and brown trophic groups diversity respond to different drivers during succession. We found that, while species richness for plants and invertebrate herbivores (green web groups) both peaked at intermediate productivity and successional age, the diversity of macrodetritivores, microarthropod microbivores and secondary consumers (brown web groups) continuously increased towards the latest successional stages. These results suggest that green web trophic groups are mainly driven by vegetation parameters, such as the amount of bare soil, vegetation biomass production and vegetation height, while brown web trophic groups are mostly driven by the production and standing stock of dead organic material and soil development. Our results show that plant diversity cannot simply be used as a proxy for the diversity of all other species groups that drive ecosystem functioning, as brown and green diversity components in our ecosystem responded differently to successional gradients.

How habitat-modifying organisms structure the food web of two coastal ecosystems.

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.

Grassland structural heterogeneity in a savanna is driven more by productivity differences than by consumption differences between lawn and bunch grasses.

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).

Indirect interactions among tropical tree species through shared rodent seed predators: a novel mechanism of tree species coexistence.

The coexistence of numerous tree species in tropical forests is commonly explained by negative dependence of recruitment on the conspecific seed and tree density due to specialist natural enemies that attack seeds and seedlings ('Janzen-Connell' effects). Less known is whether guilds of shared seed predators can induce a negative dependence of recruitment on the density of different species of the same plant functional group. We studied 54 plots in tropical forest on Barro Colorado Island, Panama, with contrasting mature tree densities of three coexisting large seeded tree species with shared seed predators. Levels of seed predation were far better explained by incorporating seed densities of all three focal species than by conspecific seed density alone. Both positive and negative density dependencies were observed for different species combinations. Thus, indirect interactions via shared seed predators can either promote or reduce the coexistence of different plant functional groups in tropical forest.

Cyclical succession in grazed ecosystems: the importance of interactions between different-sized herbivores and different-sized predators.

Body size of vertebrate herbivores is strongly linked to other life history traits, most notably (1) tolerance of low quality forage and (2) vulnerability to predation, which both impact the composition and dynamics of natural communities. However, no study has thus far explored how the combination of these two body-size related traits affects the long-term composition and dynamics of the herbivore and plant communities. We made a simple model of ordinary differential equations and simulated a grassland system with three herbivore species (small, medium, large) and two predator species (small, large) to investigate how the combination of low-quality tolerance and predation-vulnerability structure the herbivore and plant community. We found that facilitation and competition between different-sized herbivores and predation by especially small predators stimulate coexistence among herbivore species. Furthermore, the interaction between different-sized herbivores and predators generated cyclical succession in the plant community, i.e. alternating periods of short vegetation dominated by high-quality plants, with periods of tall vegetation dominated by low-quality plants. Our results suggest that cyclical succession in plant communities is more likely to occur when a predator predominantly preys on small herbivore species. Large predators also play an important role, as their addition relaxed the set of conditions under which cyclical succession occurred. Consequently, our model predictions suggest that a diverse predator community plays an important role in the long-term dynamics and maintenance of diversity in both the herbivore and plant community.

Mesoherbivores affect grasshopper communities in a megaherbivore-dominated South African savannah.

African savannahs are among the few places on earth where diverse communities of mega- and meso-sized ungulate grazers dominate ecosystem functioning. Less conspicuous, but even more diverse, are the communities of herbivorous insects such as grasshoppers, which share the same food. Various studies investigated the community assembly of these groups separately, but it is poorly known how ungulate communities shape grasshopper communities. Here, we investigated how ungulate species of different body size alter grasshopper communities in a South African savannah. White rhino is the most abundant vertebrate herbivore in our study site. Other common mesoherbivores include buffalo, zebra and impala. We hypothesized that white rhinos would have greater impact than mesoherbivores on grasshopper communities. Using 10-year-old exclosures, at eight sites we compared the effects of ungulates on grasshopper communities in three nested treatments: (i) unfenced plots ('control plots') with all vertebrate herbivores present, (ii) plots with a low cable fence, excluding white rhino ('megaherbivore exclosures'), and (iii) plots with tall fences, excluding all herbivores larger than rodents ('complete ungulate exclosures'). In each plot, we collected data of vegetation structure, grass and grasshopper community composition. Complete ungulate exclosures contained 30% taller vegetation than megaherbivore exclosures and they were dominated by different grass and grasshopper species. Grasshoppers in complete ungulate exclosures were on average 3.5 mm longer than grasshoppers in megaherbivore exclosures, possibly due to changes in plant communities or vegetation structure. We conclude that surprisingly, in this megaherbivore hotspot, mesoherbivores, instead of megaherbivores, most strongly affect grasshopper communities.

Generalities in grazing and browsing ecology: using across-guild comparisons to control contingencies.

In community ecology, broad-scale spatial replication can accommodate contingencies in patterns within species groups, but contingencies in processes across species groups remain problematic. Here, based on a focused review of grazing and browsing by large mammals, we use one trophic guild as a "control" for the other to identify generalities that are not contingent upon specific consumer-resource interactions. An example of such a generality is the Jarman-Bell principle, which explains how allometries of metabolism and digestion influence dietary tolerance and thereby enable resource partitioning within both guilds at multiple scales. By comparing the grazing succession with browsing stratification we show how competition from smaller herbivores, rather than facilitation from larger ones, is the underlying process structuring ungulate assemblages when shared resources become limiting. Also, grazing lawns and browsing hedges are functionally similar. In each case, plants expressing tolerance traits can withstand chronic grazing or browsing in sites where the nutritive value of the local food resource is enhanced in positive feedback to the actions of its consumers. The debate over whether ungulates accelerate or decelerate nutrient cycling can be resolved by comparing grazing and browsing effects in the same ecosystem type. Evidence from African savannas points to the rate of nutrient cycling being controlled by the mix of tolerance and resistance traits in plants; not the relative dominance of grazing or browsing by local herbivores. We recommend this across-guild comparative approach as a novel solution with widespread utility for resolving contingencies in community processes.

Cross-habitat interactions among bivalve species control community structure on intertidal flats.

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

Herbivore trampling as an alternative pathway for explaining differences in nitrogen mineralization in moist grasslands.

Studies addressing the role of large herbivores on nitrogen cycling in grasslands have suggested that the direction of effects depends on soil fertility. Via selection for high quality plant species and input of dung and urine, large herbivores have been shown to speed up nitrogen cycling in fertile grassland soils while slowing down nitrogen cycling in unfertile soils. However, recent studies show that large herbivores can reduce nitrogen mineralization in some temperate fertile soils, but not in others. To explain this, we hypothesize that large herbivores can reduce nitrogen mineralization in loamy or clay soils through soil compaction, but not in sandy soils. Especially under wet conditions, strong compaction in clay soils can lead to periods of soil anoxia, which reduces decomposition of soil organic matter and, hence, N mineralization. In this study, we use a long-term (37-year) field experiment on a salt marsh to investigate the hypothesis that the effect of large herbivores on nitrogen mineralization depends on soil texture. Our results confirm that the presence of large herbivores decreased nitrogen mineralization rate in a clay soil, but not in a sandy soil. By comparing a hand-mown treatment with a herbivore-grazed treatment, we show that these differences can be attributed to herbivore-induced changes in soil physical properties rather than to above-ground biomass removal. On clay soil, we find that large herbivores increase the soil water-filled porosity, induce more negative soil redox potentials, reduce soil macrofauna abundance, and reduce decomposition activity. On sandy soil, we observe no changes in these variables in response to grazing. We conclude that effects of large herbivores on nitrogen mineralization cannot be understood without taking soil texture, soil moisture, and feedbacks through soil macrofauna into account.

The metamicrobiome: key determinant of the homeostasis of nutrient recycling.

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.

The seafloor from a trait perspective. A comprehensive life history dataset of soft sediment macrozoobenthos.

Biological trait analysis (BTA) is a valuable tool for evaluating changes in community diversity and its link to ecosystem processes as well as environmental and anthropogenic perturbations. Trait-based analytical techniques like BTA rely on standardised datasets of species traits. However, there are currently only a limited number of datasets available for marine macrobenthos that contain trait data across multiple taxonomic groups. Here, we present an open-access dataset of 16 traits for 235 macrozoobenthic species recorded throughout multiple sampling campaigns of the Dutch Wadden Sea; a dynamic soft bottom system where humans have long played a substantial role in shaping the coastal environment. The trait categories included in this dataset cover a variety of life history strategies that are tightly linked to ecosystem functioning and the resilience of communities to (anthropogenic) perturbations and can advance our understanding of environmental changes and human impacts on the functioning of soft bottom systems.

Testing the metabolic theory of ecology.

The metabolic theory of ecology (MTE) predicts the effects of body size and temperature on metabolism through considerations of vascular distribution networks and biochemical kinetics. MTE has also been extended to characterise processes from cellular to global levels. MTE has generated both enthusiasm and controversy across a broad range of research areas. However, most efforts that claim to validate or invalidate MTE have focused on testing predictions. We argue that critical evaluation of MTE also requires strong tests of both its theoretical foundations and simplifying assumptions. To this end, we synthesise available information and find that MTE's original derivations require additional assumptions to obtain the full scope of attendant predictions. Moreover, although some of MTE's simplifying assumptions are well supported by data, others are inconsistent with empirical tests and even more remain untested. Further, although many predictions are empirically supported on average, work remains to explain the often large variability in data. We suggest that greater effort be focused on evaluating MTE's underlying theory and simplifying assumptions to help delineate the scope of MTE, generate new theory and shed light on fundamental aspects of biological form and function.

Alternative mechanisms alter the emergent properties of self-organization in mussel beds.

Theoretical models predict that spatial self-organization can have important, unexpected implications by affecting the functioning of ecosystems in terms of resilience and productivity. Whether and how these emergent effects depend on specific formulations of the underlying mechanisms are questions that are often ignored. Here, we compare two alternative models of regular spatial pattern formation in mussel beds that have different mechanistic descriptions of the facilitative interactions between mussels. The first mechanism involves a reduced mussel loss rate at high density owing to mutual protection between the mussels, which is the basis of prior studies on the pattern formation in mussels. The second mechanism assumes, based on novel experimental evidence, that mussels feed more efficiently on top of mussel-generated hummocks. Model simulations point out that the second mechanism produces very similar types of spatial patterns in mussel beds. Yet the mechanisms predict a strikingly contrasting effect of these spatial patterns on ecosystem functioning, in terms of productivity and resilience. In the first model, where high mussel densities reduce mussel loss rates, patterns are predicted to strongly increase productivity and decrease the recovery time of the bed following a disturbance. When pattern formation is generated by increased feeding efficiency on hummocks, only minor emergent effects of pattern formation on ecosystem functioning are predicted. Our results provide a warning against predictions of the implications and emergent properties of spatial self-organization, when the mechanisms that underlie self-organization are incompletely understood and not based on the experimental study.

Ecosystem assembly rules: the interplay of green and brown webs during salt marsh succession.

Current theories about vegetation succession and food web assembly are poorly compatible, as food webs are generally viewed to be static, and succession is usually analyzed without the inclusion of higher trophic levels. In this study we present results from a detailed analysis of ecosystem assembly rules over a chronosequence of 100 years of salt marsh succession. First, using 13 yearlong observations on vegetation and soil parameters in different successional stages, we show that the space-for-time substitution is valid for this chronosequence. We then quantify biomass changes for all dominant invertebrate and vertebrate species across all main trophic groups of plants and animals. All invertebrate and vertebrate species were assigned to a trophic group according to feeding preference, and changes in trophic group abundance were quantified for seven different successional stages of the ecosystem. We found changes from a marine-fueled, decomposer-based (brown) food web in early stages to a more terrestrial, plant-based, herbivore-driven (green) food web in intermediate succession stages, and finally to a decomposer-based, terrestrial-driven food web in the latest stages. These changes were accompanied not only by an increase in live plant biomass and a leveling toward late succession but also by a constant increase in the amount of dead plant biomass over succession. Our results show that the structure and dynamics of salt marsh food webs cannot be understood except in light of vegetation succession, and vice versa.

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers.

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.

The distribution of large herbivore hotspots in relation to environmental and anthropogenic correlates in the Mara region of Kenya.

1. The distributions of large herbivores in protected areas and their surroundings are becoming increasingly restricted by changing land use, with adverse consequences for wildlife populations. 2. We analyse changes in distributions of herbivore hotspots to understand their environmental and anthropogenic correlates using 50 aerial surveys conducted at a spatial resolution of 5 × 5 km(2) (n = 289 cells) in the Mara region of Kenya during 1977-2010. We compare the distributions across seasons, land use types (protection, pastoralism and agro-pastoralism) and 10 species with different body sizes and feeding styles. 3. Small herbivores that are the most susceptible to predation and dependent on high-quality forage concentrate in the greenest and wet areas and close to rivers in Masai pastoral ranches in both seasons. Livestock grazing creates conditions favouring small herbivores in these ranches, including high-quality short grasses and better visibility, implying facilitation. But in the reserve, they concentrate in browner, drier and flatter areas and farther from rivers, suggesting facilitation by large grazers in the wet season, or little competition with migratory herbivores occupying the reserve in the dry season. 4. In the wet season, medium herbivores concentrate in similar areas to small herbivores in the ranches and reserve. However, in the dry season, they stay in the reserve, and also concentrate in green and wet areas close to rivers when migrants occur in the reserve. As such areas typically have higher predation risk, this suggests facilitation by the migrants by absorbing most predation pressure or, alternatively, competitive displacement by the migrants from preferred habitats. 5. Large herbivores, which suffer the least predation, depend on bulk forage and are the most likely to engender conflicts with people, concentrate in the reserve all year. This suggests attraction to the taller and denser grass and perceived greater safety in the reserve in both seasons. 6. These results reveal how predation risk, forage quantity and quality, water, competition with and facilitation by livestock interact with individual life-history traits, seasons and land use in shaping the dynamics of herbivore hotspots in protected and human-dominated savannas.