Predator-avoidance behavior extends trophic cascades to refuge habitats

Consideration of how trait-mediated indirect interactions (TMIIs) affect community dynamics is recognized as an important focus for ecological research. Although these indirect effects have been shown to mediate trophic cascades in ecological communities, our understanding of how habitat refuge influences the strength and direction of cascading effects is limited. We examined whether or not oyster toadfish (top predator) affect mud crab (intermediate predator) foraging on juvenile hard clams (infaunal prey) in oyster reefs, a physically complex habitat that can provide refuge for both intermediate predators and basal prey. In particular, we manipulated toadfish presence in mesocosms containing experimental oyster reefs and quantified both mud crab and juvenile clam mortality. Toadfish significantly reduced mud crab foraging on clams and increased clam survivorship even though mud crabs foraging on the surface of the reef sought refuge from toadfish deeper within the oyster-shell matrix where they were more proximal to clams. This counterintuitive result suggests, that toadfish suppression of mud crab foraging activity is far stronger than toadfish-avoidance behavior that potentially increases crab-clam encounter rates. Therefore, TMIIs can reinforce trophic cascades even in refuge habitats where intermediate predators and their prey are physically isolated from top predators. Determining the generality of cascading effects on lower trophic levels within refugia will require investigating how habitat refuge affects the relative importance of TMIIs.