A facilitation cascade enhances local biodiversity in seagrass beds

Invertebrate diversity can be a key driver of ecosystem functioning, yet understanding what factors influence local biodiversity remains uncertain. In many marine and terrestrial systems, facilitation cascades where primary foundation and/or autogenic ecosystem engineering species promote the settlement and survival of a secondary foundation/engineering species have been shown to enhance local biodiversity and ecosystem functioning. We experimentally tested if a facilitation cascade occurs among eelgrass (Zostera marina), pen clams (Atrina rigida), and community diversity in temperate seagrass beds in North Carolina, U.S.A., and if this sequence of direct positive interactions created feedbacks that affected various metrics of seagrass ecosystem function and structure. Using a combination of surveys and transplant experiments, we found that pen clam density and survivorship was significantly greater in seagrass beds, indicating that eelgrass facilitates pen clams. Pen clams in turn enhanced local diversity and increased both the abundance and species richness of organisms (specifically, macroalgae and fouling invertebrate fauna)-the effect of which scaled with increasing clam density. However, we failed to detect an impact of pen clams on other seagrass functions and hypothesize that functioning may more likely be enhanced in scenarios where secondary foundation species specifically increase the diversity of key functional groups such as epiphyte grazers and/or when bivalves are infaunal rather than epifaunal. Our findings add to the growing amount of literature that demonstrates that secondary foundation species are important drivers of local biodiversity in marine ecosystems. Further experimentation is needed that directly examines (i) the role of functional versus overall diversity on seagrass functions and (ii) the relative importance of life-history strategy in determining when and where engineering bivalves increase biodiversity and/or functioning of seagrass beds.