Positive indirect effects of reef fishes on kelp performance: the importance of mesograzers.

It has been suggested that microcarnivorous reef fishes may play an important role in giant kelp forest communities by preventing infestations of mesograzers that could severely impact or potentially destroy recovering kelp forests after extreme disturbance events. However, these trophic linkages, specifically the direct and indirect effects of fishes on the biomass of mesograzers, grazing intensity, and the performance of giant kelp, have not been sufficiently quantified and evaluated as to their importance and in the absence of such disturbance events. We examined experimentally the effects of mesograzers on the growth and performance of giant kelp in the presence and absence of their fish predators near Santa Catalina Island, California (U.S.A.). Mesograzer biomass and grazing intensity were significantly higher when fishes were excluded from giant kelp, which in turn, lowered kelp performance. This pattern was consistent both on experimental plots of kelp as habitat isolates, and on a continuous reef. Moreover, the abundance of mesograzers was inversely related to the abundance of kelp perch among several kelp-forested reefs, suggesting that these effects can occur at larger spatial scales. Because of differences in the diet and behavior of two microcarnivorous fishes, the kelp perch and señorita, we conducted an experiment manipulating each species and its density independently to determine their separate effects on mesograzers and kelp performance. Concurrently we examined the growth and mortality of juvenile kelp. Grazing intensity decreased, estimates of kelp performance increased, and the growth of juvenile kelp increased with increasing densities of fish but with no detectable effects between fishes. Our results demonstrate that these microcarnivorous fishes have positive indirect effects on kelp performance by reducing mesograzer biomass and grazing intensity, and the early life stages of other fishes also may be important. More specifically, these fishes have a positive effect on the density of fronds of giant kelp that can result in greater recruitment success and the abundance of kelp-associated invertebrates and fishes. Indeed, this study suggests that mesograzers have the potential to be one of the most important herbivores in kelp forest ecosystems.