Biodiversity and food web indicators of community recovery in intertidal shellfish reefs

In conservation strategies of marine ecosystems, priority is given to habitat-structuring foundation species (e.g. seagrasses, mangroves and reef-building corals, shellfish) with the implicit goal to protect or restore associated communities and their interactions. However, the number and accuracy of community level metrics to measure the success of these strategies are limited. Using intertidal shellfish reefs as a model, we tested to what extent foundation species alter community and food web structure, and explored whether basic metrics of food web structure are useful indicators of ecosystem complexity compared to other often-used indices. We found that shellfish reefs strongly modified community and food web structure by modifying habitat conditions (e.g. hydrodynamics, sediment grain size). Stable isotope-based food web reconstruction captured important differences between communities from bare mudflat and shellfish reefs that did not emerge from classic abundance or diversity measures. On shellfish reefs, link density and the number of top predators were consistently higher, while both connectance and the richness of intermediate species was lower. Species richness (+42%), species density (+79%) and total biomass of benthos, fish and birds (+41%) was also higher on shellfish reefs, but this did not affect the Shannon diversity or Evenness. Hence, our results showed that basic food web metrics such as link density and number of top consumers and intermediate species combined with traditional measures of species richness can provide a robust tool to measure conservation and restoration success. We therefore suggest that these metrics are included as Essential Biodiversity Variables (EBV), and implemented as ecosystem health indicators in legislative frameworks such as the Marine Strategy Framework Directive (MSFD).