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.

Integrating defense and leaf economic spectrum traits in a tropical savanna plant.

Introduction: Allocation to plant defense traits likely depends on resource supply, herbivory, and other plant functional traits such as the leaf economic spectrum (LES) traits. Yet, attempts to integrate defense and resource acquisitive traits remain elusive. Methods: We assessed intraspecific covariation between different defense and LES traits in a widely distributed tropical savanna herb, Solanum incanum, a unique model species for studying allocations to physical, chemical, and structural defenses to mammalian herbivory. Results: We found that in a multivariate trait space, the structural defenses - lignin and cellulose - were positively related to the resource conservative traits - low SLA and low leaf N. Phenolic content, a chemical defense, was positively associated with resource acquisitive traits - high SLA and high leaf N - while also being associated with an independent third component axis. Both principal components 1 and 3 were not associated with resource supply and herbivory intensity. In contrast, spine density - a physical defense - was orthogonal to the LES axis and positively associated with soil P and herbivory intensity. Discussion: These results suggest a hypothesized "pyramid" of trade-offs in allocation to defense along the LES and herbivory intensity axes. Therefore, future attempts to integrate defense traits with the broader plant functional trait framework, such as the LES, needs a multifaceted approach that accounts for unique influences of resource acquisitive traits and herbivory risk.

Episodic herbivory, plant density dependence, and stimulation of aboveground plant production.

Herbivory is a major energy transfer within ecosystems; an open question is under what circumstances it can stimulate aboveground seasonal primary production. Despite multiple field demonstrations, past theory considered herbivory as a continuous process and found stimulation of seasonal production to be unlikely. Here, we report a new theoretical model that explores the consequences of discrete herbivory events, or episodes, separated in time. We discovered that negative density (biomass) dependence of plant growth, such as might be expected from resource limitation of plant growth, favors stimulation of seasonal production by infrequent herbivory events under a wide range of herbivory intensities and maximum plant relative growth rates. Results converge to those of previous models under repeated, short-interval herbivory, which generally reduces seasonal production. Model parameters were estimated with new and previous data from the Serengeti ecosystem. Patterns of observed frequent and large magnitude stimulated production in these data agreed generally with those predicted by the episodic herbivory model. The model thus may provide a new framework for evaluating the sustainability and impact of herbivory.

Cross-boundary human impacts compromise the Serengeti-Mara ecosystem.

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.

Reaction and diffusion thermodynamics explain optimal temperatures of biochemical reactions.

Ubiquitous declines in biochemical reaction rates above optimal temperatures (Topt) are normally attributed to enzyme state changes, but such mechanisms appear inadequate to explain pervasive Topt well below enzyme deactivation temperatures (Tden). Here, a meta-analysis of 92 experimental studies shows that product formation responds twice as strongly to increased temperature than diffusion or transport. This response difference has multiple consequences for biochemical reactions, such as potential shifts in the factors limiting reactions as temperature increases and reaction-diffusion dynamics that predict potential product inhibition and limitation of the reaction by entropy production at temperatures below Tden. Maximizing entropy production by the reaction predicts Topt that depend on enzyme concentration and efficiency as well as reaction favorability, which are patterns not predicted by mechanisms of enzyme state change. However, these predictions are strongly supported by patterns in a meta-analysis of 121 enzyme kinetic studies. Consequently, reaction-diffusion thermodynamics and entropy production may constrain organism performance at higher temperatures, yielding temperature optima of life that may depend on reaction characteristics and environmental features rather than just enzyme state changes.

Genetic variation and population structure in a threatened species, the Utah prairie dog Cynomys parvidens: the use of genetic data to inform conservation actions.

The Utah prairie dog (Cynomys parvidens), listed as threatened under the United States Endangered Species Act, was the subject of an extensive eradication program throughout its range during the 20th century. Eradication campaigns, habitat destruction/fragmentation/conversion, and epizootic outbreaks (e.g., sylvatic plague) have reduced prairie dog numbers from an estimated 95,000 individuals in the 1920s to approximately 14,000 (estimated adult spring count) today. As a result of these anthropogenic actions, the species is now found in small isolated sets of subpopulations. We characterized the levels of genetic diversity and population genetic structure using 10 neutral nuclear microsatellite loci for twelve populations (native and transplanted) representative of the three management designated "recovery units," found in three distinct biogeographic regions, sampled across the species' range. The results indicate (1) low levels of genetic diversity within colonies (H e = 0.109-0.357; H o = 0.106- 0.313), (2) high levels of genetic differentiation among colonies (global F ST = 0.296), (3) very small genetic effective population sizes, and (4) evidence of genetic bottlenecks. The genetic data reveal additional subdivision such that colonies within recovery units do not form single genotype clusters consistent with recovery unit boundaries. Genotype cluster membership support historical gene flow among colonies in the easternmost West Desert Recovery Unit with the westernmost Pausaugunt colonies and among the eastern Pausaugunt colonies and the Awapa Recovery unit to the north. In order to maintain the long-term viability of the species, there needs to be an increased focus on maintaining suitable habitat between groups of existing populations that can act as connective corridors. The location of future translocation sites should be located in areas that will maximize connectivity, leading to maintenance of genetic variation and evolutionary potential.

Intraspecific trait variation drives functional responses of old-field plant communities to nutrient enrichment.

Environmental changes are expected to shift the distribution of functional trait values in plant communities through a combination of species turnover and intraspecific variation. The strength of these shifts may depend on the availability of individuals with trait values adapted to new environmental conditions, represented by the functional diversity (FD) of existing community residents or dispersal from the regional species pool. We conducted a 3-year nutrient- and seed-addition experiment in old-field plant communities to examine the contributions of species turnover and intraspecific variation to community trait shifts, focusing on four key plant functional traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We further examined the influence of initial FD and seed availability on the strength of these shifts. Community mean height, leaf area, and SLA increased in response to fertilization, and these shifts were driven almost entirely by intraspecific variation. The strength of intraspecific shifts in height and leaf area was positively related to initial intraspecific FD in these traits. Intraspecific trait responses to fertilization varied among species, with species of short stature displaying stronger shifts in SLA and LDMC but weaker shifts in leaf area. Trait shifts due to species turnover were generally weak and opposed intraspecific responses. Seed addition altered community taxonomic composition but had little effect on community trait shifts. These results highlight the importance of intraspecific variation for short-term community functional responses and demonstrate that the strength of these responses may be mediated by community FD.

Contrasting effects of different mammalian herbivores on sagebrush plant communities.

Herbivory by both grazing and browsing ungulates shapes the structure and functioning of terrestrial ecosystems worldwide, and both types of herbivory have been implicated in major ecosystem state changes. Despite the ecological consequences of differences in diets and feeding habits among herbivores, studies that experimentally distinguish effects of grazing from spatially co-occurring, but temporally segregated browsing are extremely rare. Here we use a set of long-term exclosures in northern Utah, USA, to determine how domestic grazers vs. wild ungulate herbivores (including browsers and mixed feeders) affect sagebrush-dominated plant communities that historically covered ~62 million ha in North America. We sampled plant community properties and found that after 22 years grazing and browsing elicited perceptible changes in overall plant community composition and distinct responses by individual plant species. In the woody layer of the plant community, release from winter and spring wild ungulate herbivory increased densities of larger Wyoming big sagebrush (Artemisia tridentata, ssp. wyomingensis) at the expense of small sagebrush, while disturbance associated with either cattle or wild ungulate activity alone was sufficient to increase bare ground and reduce cover of biological soil crusts. The perennial bunchgrass, bottlebrush squirretail (Elymus elymoides), responded positively to release from summer cattle grazing, and in turn appeared to competitively suppress another more grazing tolerant perennial grass, Sandberg's blue grass (Poa secunda). Grazing by domestic cattle also was associated with increased non-native species biomass. Together, these results illustrate that ungulate herbivory has not caused sagebrush plant communities to undergo dramatic state shifts; however clear, herbivore-driven shifts are evident. In a dry, perennial-dominated system where plant community changes can occur very slowly, our results provide insights into potential long-term trajectories of these plant communities under different large herbivore regimes. Our results can be used to guide long-term management strategies for sagebrush systems and improve habitat for endemic wildlife species such as sage-grouse (Centrocercus spp.).

Community functional responses to soil and climate at multiple spatial scales: when does intraspecific variation matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.

The effect of fire on habitat selection of mammalian herbivores: the role of body size and vegetation characteristics.

Given the role of fire in shaping ecosystems, especially grasslands and savannas, it is important to understand its broader impact on these systems. Post-fire stimulation of plant nutrients is thought to benefit grazing mammals and explain their preference for burned areas. However, fire also reduces vegetation height and increases visibility, thereby potentially reducing predation risk. Consequently, fire may be more beneficial to smaller herbivores, with higher nutritional needs and greater risks of predation. We tested the impacts of burning on different sized herbivores' habitat preference in Serengeti National Park, as mediated by burning's effects on vegetation height, live : dead biomass ratio and leaf nutrients. Burning caused a less than 4 month increase in leaf nitrogen (N), and leaf non-N nutrients [copper (Cu), potassium (K), and magnesium (Mg)] and a decrease in vegetation height and live : dead biomass. During this period, total herbivore counts were higher on burned areas. Generally, smaller herbivores preferred burned areas more strongly than larger herbivores. Unfortunately, it was not possible to determine the vegetation characteristics that explained burned area preference for each of the herbivore species observed. However, total herbivore abundance and impala (Aepyceros melampus) preference for burned areas was due to the increases in non-N nutrients caused by burning. These findings suggest that burned area attractiveness to herbivores is mainly driven by changes to forage quality and not potential decreases in predation risk caused by reductions in vegetation height.

Body size mediated coexistence in swans.

Differences in body sizes may create a trade-off between foraging efficiency (foraging gains/costs) and access to resources. Such a trade-off provides a potential mechanism for ecologically similar species to coexist on one resource. We explored this hypothesis for tundra (Cygnus columbianus) and trumpeter swans (Cygnus buccinator), a federally protected species, feeding solely on sago pondweed (Stuckenia pectinata) tubers during fall staging and wintering in northern Utah. Foraging efficiency was higher for tundra swans because this species experienced lower foraging and metabolic costs relative to foraging gains; however, trumpeter swans (a) had longer necks and therefore had access to exclusive resources buried deep in wetland sediments and (b) were more aggressive and could therefore displace tundra swans from lucrative foraging locations. We conclude that body size differentiation is an important feature of coexistence among ecologically similar species feeding on one resource. In situations where resources are limiting and competition for resources is strong, conservation managers will need to consider the trade-off between foraging efficiency and access to resources to ensure ecologically similar species can coexist on a shared resource.

Plant compensation to grazing and soil carbon dynamics in a tropical grassland.

The effects of grazing on soil organic carbon (SOC) dynamics, particularly in the tropics, are still poorly understood. Plant compensation to grazing, whereby plants maintain leaf area (C input capacity) despite consumption (C removal) by grazers, has been demonstrated in tropical grasslands but its influence on SOC is largely unexplored. Here, the effect of grazing on plant leaf area index (LAI) was measured in a field experiment in Serengeti National Park, Tanzania. LAI changed little for grazing intensities up to 70%. The response curve of LAI versus grazing intensity was used in a mass balance model, called SNAP, of SOC dynamics based on previous data from the Serengeti. The model predicted SOC to increase at intermediate grazing intensity, but then to decline rapidly at the highest grazing intensities. The SNAP model predictions were compared with observed SOC stocks in the 24 grazed plots of a 10-year grazing exclosure experiment at eight sites across the park that varied in mean annual rainfall, soil texture, grazing intensity and plant lignin and cellulose. The model predicted current SOC stocks very well (R (2) > 0.75), and suggests that compensatory plant responses to grazing are an important means of how herbivores might maintain or increase SOC in tropical grasslands.

Effects of grazing on grassland soil carbon: a global review.

Soils of grasslands represent a large potential reservoir for storing CO2 , but this potential likely depends on how grasslands are managed for large mammal grazing. Previous studies found both strong positive and negative grazing effects on soil organic carbon (SOC) but explanations for this variation are poorly developed. Expanding on previous reviews, we performed a multifactorial meta-analysis of grazer effects on SOC density on 47 independent experimental contrasts from 17 studies. We explicitly tested hypotheses that grazer effects would shift from negative to positive with decreasing precipitation, increasing fineness of soil texture, transition from dominant grass species with C3 to C4 photosynthesis, and decreasing grazing intensity, after controlling for study duration and sampling depth. The six variables of soil texture, precipitation, grass type, grazing intensity, study duration, and sampling depth explained 85% of a large variation (±150 g m(-2)  yr(-1) ) in grazing effects, and the best model included significant interactions between precipitation and soil texture (P = 0.002), grass type, and grazing intensity (P = 0.012), and study duration and soil sampling depth (P = 0.020). Specifically, an increase in mean annual precipitation of 600 mm resulted in a 24% decrease in grazer effect size on finer textured soils, while on sandy soils the same increase in precipitation produced a 22% increase in grazer effect on SOC. Increasing grazing intensity increased SOC by 6-7% on C4 -dominated and C4 -C3 mixed grasslands, but decreased SOC by an average 18% in C3 -dominated grasslands. We discovered these patterns despite a lack of studies in natural, wildlife-dominated ecosystems, and tropical grasslands. Our results, which suggest a future focus on why C3 vs. C4 -dominated grasslands differ so strongly in their response of SOC to grazing, show that grazer effects on SOC are highly context-specific and imply that grazers in different regions might be managed differently to help mitigate greenhouse gas emissions.

The species-rich assemblages of tintinnids (marine planktonic protists) are structured by mouth size.

Many microbial taxa in the marine plankton appear super-saturated in species richness. Here, we provide a partial explanation by analyzing how species are organized, species packing, in terms of both taxonomy and morphology. We focused on a well-studied group, tintinnid ciliates of the microzooplankton, in which feeding ecology is closely linked to morphology. Populations in three distinct systems were examined: an Eastern Mediterranean Gyre, a Western Mediterranean Gyre and the California Current. We found that species abundance distributions exhibited the long-tailed, log distributions typical of most natural assemblages of microbial and other organisms. In contrast, grouping in oral size-classes, which corresponds with prey-size exploited, revealed a geometric distribution consistent with a dominant role of a single resource in structuring an assemblage. The number of species found in a particular oral size-class increases with the numerical importance of the size-class in the overall population. We suggest that high species diversity reflects the fact that accompanying each dominant species are many ecologically similar species, presumably able to replace the dominant species, at least with regard to the size of prey exploited. Such redundancy suggests that species diversity greatly exceeds ecological diversity in the plankton.

Herbivore effects on above- and belowground plant production and soil nitrogen availability in the Trans-Himalayan shrub-steppes.

Large mammalian herbivores may have positive, neutral, or negative effects on annual net aboveground plant production (NAP) in different ecosystems, depending on their indirect effects on availability of key nutrients such as soil N. In comparison, less is known about the corresponding influence of grazers, and nutrient dynamics, over annual net belowground plant production (NBP). In natural multi-species plant communities, it remains uncertain how grazing influences relative allocation in the above- and belowground compartments in relation to its effects on plant nutrients. We evaluated grazer impacts on NAP, NBP, and relative investment in the above- and belowground compartments, alongside their indirect effects on soil N availability in the multiple-use Trans-Himalayan grazing ecosystem with native grazers and livestock. Data show that a prevailing grazing intensity of 51% increases NAP (+61%), but reduces NBP (-35%). Grazing also reduced C:N ratio in shoots (-16%) and litter (-50%), but not in roots, and these changes coincided with increased plant-available inorganic soil N (+23%). Areas used by livestock and native grazers showed qualitatively similar responses since NAP was promoted, and NBP was reduced, in both cases. The preferential investment in the aboveground fraction, at the expense of the belowground fraction, was correlated positively with grazing intensity and with improvement in litter quality. These results are consistent with hypothesized herbivore-mediated positive feedbacks between soil nutrients and relative investment in above- and belowground compartments. Since potentially overlapping mechanisms, such as N mineralization rate, plant N uptake, compositional turnover, and soil microbial activity, may contribute towards these feedbacks, further studies may be able to discern their respective contributions.

Introduced grazers can restrict potential soil carbon sequestration through impacts on plant community composition.

Grazing occurs over a third of the earth's land surface and may potentially influence the storage of 10(9) Mg year(-1) of greenhouse gases as soil C. Displacement of native herbivores by high densities of livestock has often led to overgrazing and soil C loss. However, it remains unknown whether matching livestock densities to those of native herbivores can yield equivalent soil C sequestration. In the Trans-Himalayas we found that, despite comparable grazing intensities, watersheds converted to pastoralism had 49% lower soil C than watersheds which retain native herbivores. Experimental grazer-exclusion within each watershed type, show that this difference appears to be driven by indirect effects of livestock diet selection, leading to vegetation shifts that lower plant production and reduce likely soil C inputs from vegetation by c. 25 gC m(-2) year(-1). Our results suggest that while accounting for direct impacts (stocking density) is a major step, managing indirect impacts on vegetation composition are equally important in influencing soil C sequestration in grazing ecosystems.

Ciliate diversity and distribution across an environmental and depth gradient in Long Island Sound, USA.

The nature and extent of microbial biodiversity remain controversial with persistent debates over patterns of distributions (i.e. cosmopolitanism versus endemism) and the processes that structure these patterns (neutrality versus selection). We used culture-independent approaches to address these issues focusing on two groups of ciliates, the Oligotrichia (Spirotrichea) and Choreotrichia (Spirotrichea) across an environmental gradient. We assessed SSU rDNA diversity in ciliate communities at six stations in Long Island Sound spanning the frontal region that separates the fresher Connecticut River outflow plume from the open Sound. As in previous studies, we find one abundant cosmopolitan species (Strombidium biarmatum), a few moderately abundant sequences, and a long list of rare sequences. Furthermore, neither ciliate diversity nor species composition showed any clear relationship to measured environmental parameters (temperature, salinity, accessory pigment composition and chorophyll). Overall, we observed that diversity decreased moving from nearshore to offshore. We also conducted analyses to detect clustering among the sampled communities using the software Unifrac. This approach revealed three significant clusters grouping samples from nearshore, surface and deep/well mixed stations. We find no strong fit of our communities to log series, geometric or log normal distributions, though one of the 3 clusters is most consistent with a log series distribution. However, when we remove the abundant cosmopolitan species S. biarmatum, all three clusters fit to a log series distribution. These analyses suggest that, with the exception of one cosmopolitan species, the oligotrich and choreotrich communities at these stations may be distributed in a neutral manner.

A disease-mediated trophic cascade in the Serengeti and its implications for ecosystem C.

Tree cover is a fundamental structural characteristic and driver of ecosystem processes in terrestrial ecosystems, and trees are a major global carbon (C) sink. Fire and herbivores have been hypothesized to play dominant roles in regulating trees in African savannas, but the evidence for this is conflicting. Moving up a trophic scale, the factors that regulate fire occurrence and herbivores, such as disease and predation, are poorly understood for any given ecosystem. We used a Bayesian state-space model to show that the wildebeest population eruption that followed disease (rinderpest) eradication in the Serengeti ecosystem of East Africa led to a widespread reduction in the extent of fire and an ongoing recovery of the tree population. This supports the hypothesis that disease has played a key role in the regulation of this ecosystem. We then link our state-space model with theoretical and empirical results quantifying the effects of grazing and fire on soil carbon to predict that this cascade may have led to important shifts in the size of pools of C stored in soil and biomass. Our results suggest that the dynamics of herbivores and fire are tightly coupled at landscape scales, that fire exerts clear top-down effects on tree density, and that disease outbreaks in dominant herbivores can lead to complex trophic cascades in savanna ecosystems. We propose that the long-term status of the Serengeti and other intensely grazed savannas as sources or sinks for C may be fundamentally linked to the control of disease outbreaks and poaching.

Forage nutritive quality in the Serengeti ecosystem: the roles of fire and herbivory.

Fire and herbivory are important determinants of nutrient availability in savanna ecosystems. Fire and herbivory effects on the nutritive quality of savanna vegetation can occur directly, independent of changes in the plant community, or indirectly, via effects on the plant community. Indirect effects can be further subdivided into those occurring because of changes in plant species composition or plant abundance (i.e., quality versus quantity). We studied relationships between fire, herbivory, rainfall, soil fertility, and leaf nitrogen (N), phosphorus (P), and sodium (Na) at 30 sites inside and outside of Serengeti National Park. Using structural equation modeling, we asked whether fire and herbivory influences were largely direct or indirect and how their signs and strengths differed within the context of natural savanna processes. Herbivory was associated with enhanced leaf N and P through changes in plant biomass and community composition. Fire was associated with reduced leaf nutrient concentrations through changes in plant community composition. Additionally, fire had direct positive effects on Na and nonlinear direct effects on P that partially mitigated the indirect negative effects. Key mechanisms by which fire reduced plant nutritive quality were through reductions of Na-rich grasses and increased abundance of Themeda triandra, which had below-average leaf nutrients.

Rainfall and soils modify plant community response to grazing in Serengeti National Park.

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.