Indirect effects of sea otters on rockfish (Sebastes spp.) in giant kelp forests.

Sea otters are a classic example of a predator controlling ecosystem productivity through cascading effects on basal, habitat-forming kelp species. However, their indirect effects on other kelp-associated taxa like fishes are poorly understood. We examined the effects of sea otter (Enhydra lutris) reintroduction along the west coast of Vancouver Island, Canada on giant kelp (Macrocystis pyrifera) distributions and the trophic niches and growth of two common kelp forest fishes, black (Sebastes melanops) and copper (S. caurinus) rockfishes. We sampled 47 kelp forests, and found that red sea urchins (Strongylocentrotus franciscanus) were eliminated in the presence of otters, and that kelp forests were 3.7 times deeper and 18.8 times larger. Despite order-of-magnitude differences in kelp forest size, adult black and copper rockfishes contained less kelp-derived carbon in their tissues (as measured by stable isotopes of C and N) in regions with otters. Adults of both species had higher mean trophic positions in the presence of otters, indicating more frequent consumption of higher trophic level prey such as fishes. Smaller trophic niche space of rockfishes in the presence of otters indicated a higher degree of trophic specialization. Juvenile black rockfishes rapidly shifted to higher kelp-carbon contents, trophic positions, and body condition factors after settling in kelp forests. The relationships of growth to length, percentage of kelp carbon, and trophic position varied between the two regions, indicating that potential effects of kelp forest size on trophic ontogeny may also affect individual performance. Our results provide evidence that the indirect effects of otters on rockfishes arise largely through the creation of habitat for fishes and other prey rather than a direct trophic connection through invertebrates or other consumers of kelp productivity