Resource availability and plant age drive defense against herbivory in salt marshes

Abstract

The resource‐availability hypothesis (RAH) and the intraspecific RAH (RAHintra), posit that resources, (i.e. nutrients) control plant antiherbivore defenses. Both hypotheses predict that in low‐resource environments, plant growth is slow, and constitutive defense is high. In high‐resource environments, however, the RAH predicts that plant growth is fast, and constitutive defense is low, whereas the RAHintra predicts that increased resources attract more herbivores, and this intensified grazing pressure leads to high constitutive defense. Salt marshes are nutrient‐limited ecosystems threatened by eutrophication and chronic herbivory, yet we know little about how these stressors shape saltmarsh plant antiherbivore defenses, which influence trophic interactions and ecosystem resilience. We manipulated resource availability via nutrient addition and herbivory via the marsh periwinkle Littoraria irrorata, on the saltmarsh foundation species Spartina alterniflora, in mesocosms. Because plant age can also influence trait variation, we measured traits in both original and clonally‐grown new stems. Feeding assays then evaluated how treatments and plant age affected subsequent Littoraria consumption of Spartina. Nutrient addition stimulated growth, while decreasing defensive traits (e.g. fiber and silica content), following the RAH. Herbivory enhanced belowground production and increased stem diameter, yet did not induce defensive traits, contrary to our expectations. Herbivory plus nutrients increased Spartina biomass and reduced phenolics, a defensive trait, further supporting the RAH. Regardless of treatment, clonally‐grown new stems had greater variation in measured traits. Despite altered traits, however, treatments and plant age did not affect Littoraria consumption. Our results support the RAH and part of the RAHintra and suggest: 1) nutrient availability is a primary driver of plant trait change and 2) plant age controls the magnitude of trait variation in Spartina. Further, our findings indicate that eutrophic conditions may not always increase top‒down control by herbivores, and in some instances can enhance saltmarsh resilience against sea‐level rise via stimulated Spartina biomass production.