The cost of safety: refuges increase the impact of predation risk in aquatic systems.

Although use of refuge habitats by prey can reduce their risk of predation, refuge use may also involve costs such as increased within-refuge competition for resources. Despite the ubiquity of refuge use by prey, it is unknown whether predator-induced use of refuges has widespread, negative nonconsumptive effects on prey growth, survival, and fecundity. We performed a meta-analysis of 204 studies of aquatic taxa containing data on 271 distinct predator--prey pairs and found strong evidence that the negative effect of predation risk on prey activity, growth, and fecundity increases when prey have access to refuge habitats. Moreover, the effect of refuge habitats on growth and activity depends upon whether the refuge provides partial or total protection from predators. These results suggest that prey choosing whether to use refuges face a trade-off between lowering the immediate risk of being consumed and increased nonconsumptive costs of refuge use. Our results suggest that changes in nonconsumptive effects in the presence of refuge habitats may alter prey population dynamics, coexistence, and metapopulation dynamics. Moreover, our results reveal key pragmatic considerations: the magnitude and direction of nonconsumptive effects may depend on the presence of refuge habitat and whether the refuge provides partial or total protection from predators.