Landscape variation in defense traits along gradients of multiple resources and mammalian herbivory

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.