The positive effect of coexisting ecosystem engineers: a unique seaweed-mussel association provides refuge for native mud crabs against a non-indigenous predator.

In marine sedimentary bottoms, mussels and macroalgae have long been recognized as important autogenic engineers that create habitat and modify abiotic conditions. The structural complexity added by bivalves and macroalgae may also mediate intraguild predation amongst marine decapod crustaceans. While spatial distributions of these ecosystem engineers frequently overlap, there is limited understanding of compounded effects when more than one engineer is present. Here we demonstrate that the coexistence of two ecosystem engineers may create habitat valuable for the survival of a small native species, the Atlantic mud crab (), in the presence of the invasive green crab (). Using laboratory and field habitat mimics, we measured mud crab survival rates as a proxy for refuge quality. We compared the refuge provided by a unique association between shells of blue mussels () and the giant strain of Irish moss () to that provided by bare substrate, and by each engineer alone. These experiments revealed that the association of giant Irish moss with blue mussel shells positively and non-additively increased mud crab survival compared to the other less complex habitat mimics. In contrast, parallel experiments revealed that high habitat complexity was less important for young green crabs to survive predation from large conspecifics. These results suggest that the impact of ecosystem engineers on trophic dynamics should be considered in a broader, whole-community context encompassing multiple habitat-forming species present.