Additive effects of physical stress and herbivores on intertidal seaweed biodiversity

Patterns in rocky intertidal seaweed biodiversity influence the resilience and functioning of these important primary producer communities. In turn, seaweed biodiversity patterns are the result of many ecological factors. We determined the influences of thermal and desiccation stress, herbivory, and nutrients on seaweed biodiversity on a northern California rocky shoreline. In a fully crossed design at two tidal heights at wave-protected and exposed sites, we deployed screens to reduce stress, removed herbivores, and added nutrients for 18 months. The treatments reduced temperature, increased relative humidity, decreased herbivore abundances, and increased nitrogen in both seawater and seaweeds. Seaweed abundance and biodiversity (cover, biomass, species richness, diversity, evenness, and community composition) were influenced by tidal height, physical stress, and herbivores. Wave exposure affected all response variables except biomass and evenness. Stress and herbivores had independent additive effects on seaweed abundance and diversity. Physical stress did not make the community as a whole more susceptible to herbivores, and screens had overarching positive effects on seaweed biodiversity even though they also had positive effects on herbivore abundance. Nutrients had virtually no effect on seaweed biodiversity, and we observed no bottom-up effects of nutrient addition on herbivore density or biomass. Small green algae and diatoms were important contributors to overall algal cover and to changes in composition across treatments, but larger macroalgae dominated the species richness response. The striking absence of interactions between stress and herbivory highlights how seaweed communities can respond independently to important drivers of biodiversity. Thus, nonadditive, potentially synergistic effects do not necessarily complicate the understanding of how seaweed biodiversity responds to environmental change.