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Many plant species are widely distributed and consequently are exposed to multiple abiotic factors and diverse herbivores, each of which may distinctly affect the magnitude of different defense traits. Alternative theories for optimal allocation to plant defense traits predict both positive and negative associations between magnitude of defense and resource availability. These predictions may apply even within species. This suggests potential for a single species' patterns of association of defense traits and resources to vary with both the type of defense and identity of resource, but relatively few studies have explored intraspecific variation in multiple defense traits along several resource gradients simultaneously. In order to address this gap, especially in an ecosytem dominated by large mammalian herbivores, we assessed relationships between multiple resources (rainfall, soil N, and soil P) and plant defense traits (prickle density, phenolics, and lignin content) using a widely distributed tropical savanna herb, Solanum incanum, growing in naturally occuring resource gradients within the Serengeti National Park. We found substantial intraspecific variation in all three defense traits across sites (n = 43). Variation in prickle density was positively associated with rainfall and soil P, but not soil N. In contrast to prickle density, phenolics and lignin were uncorrelated with all three resource gradients. This independent association of soil P with a carbon‐based defense, prickle density, suggests potential for resources that are not components of defenses to influence allocation to defense traits. Such influence may reflect association between resource and herbivore abundance and/or preference. These varied patterns in resource–defense associations further emphasize the tremendous variation in anti‐herbivore traits which may be influenced by different plant resources and highlight the need to consider multiple resource gradients in understanding evolution of plant traits. [ABSTRACT FROM AUTHOR]

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 anaong-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. Key words: compositional similarity; congener coexistence; consumer--resource theory; functional types; grazing; plant species richness; Serengeti National Park (Tanzania); soil nitrogen; soil phosphorus.

Rapid environmental changes have fostered debates and motivated research on how to effectively preserve or restore ecosystem processes. One such debate deals with the effects of biodiversity, and the loss thereof, on ecosystem processes. Recent studies demonstrate that resource-use complementarity, now known as the 'niche-differentiation effect,' and the presence of a competitive species with strong effects on ecosystem processes, now known as the 'sampling effect,' can explain why productivity and nutrient retention are sometimes enhanced with increasing species richness. In a well-replicated outdoor mesocosm experiment, we tested these and other alternative mechanisms that could explain the effects of submersed aquatic plant (macrophyte) diversity on wetland ecosystem processes. Algal biomass increased and phosphorus loss decreased as species richness increased. This result can best be explained by an indirect sampling effect caused by one of the weakest competitors, which appeared to facilitate algal growth and thereby filtering of particles, and thus phosphorus, from the water column. The dominant competitor also appeared to decrease phosphorus loss through direct effects on phosphorus availability in the soil and water. Thus, the effects by one of the weakest and the most dominant competitors combine to produce a diversity effect on phosphorus loss. Macrophyte biomass was not enhanced, but converged toward the intermediate biomass of the most competitive species. Such an 'inverse sampling effect' may be produced when the most competitive species is not the most productive species owing to species-specific feedbacks and adaptations to the wetland environment. In summary, we reject the niche-differentiation effect as the dominant mechanism in our macrophyte communities and expand on the role of sampling effects in explaining the relationship between plant communities and ecosystem processes. In particular, indirect and inverse sampling effects combine to drive the relationship between species richness and wetland ecosystem processes. Thus, we demonstrate that plant diversity may affect wetland ecosystem processes when inferior competitors drive system productivity and nutrient retention. To ensure coexistence of such species with superior competitors, wetland systems may need to be maintained in a nonequilibrium state, such as with hydrologic disturbances, which would maintain both higher diversity and enhance ecosystem functioning. Key words: algal colonization; competitive ability; diversity; ecosystem functioning; indirect sampling effect; inverse sampling effect; nutrient retention; productivity; sampling effect; submersed aquatic macrophytes; wetland.

Author(s): John P. Haskell (corresponding author) [1]; Mark E. Ritchie [2]; Han Olff [3] Scaling laws that describe complex interactions between organisms and their environment as a function of body [...]

Author(s): Han Olff (corresponding author) [1]; Mark E. Ritchie [2]; Herbert H. T. Prins [1] Large mammalian herbivores occupy half of the earth's land surface and are important both ecologically [...]