Abstract
In marine ecosystems, fishing often targets predators, which can drive direct and indirect effects on entire food webs. Marine reserves can induce trophic cascades by increasing predator density and body size, thereby increasing predation pressure on populations of herbivores, such as sea urchins. In California's northern Channel Islands, two species of sea urchins are abundant: the red urchin Mesocentrotus franciscanus, which is targeted by an economically valuable fishery, and the virtually unfished purple urchin Strongylocentrotus purpuratus. We hypothesized that urchin populations inside marine reserves would be depressed by higher predation, but that red urchins would be less affected due to fishing outside reserves. Instead, our analyses revealed that purple urchin populations were unaffected by reserves, and red urchin biomass significantly increased in response to protection. Therefore, urchin biomass overall has increased inside reserves, and we found no evidence that giant kelp is positively affected by reserves. Our results reveal the overwhelming direct effect of protecting fished species in marine reserves over indirect effects that are often predicted but seldom clearly documented. Indirect effects due to marine reserves may eventually occur in some cases, but very effective predators, large reserves or extended time periods may be needed to induce them.