Sea otters homogenize mussel beds and reduce habitat provisioning in a rocky intertidal ecosystem

Sea otters (Enhydra lutris) are keystone predators that consume a variety of benthic invertebrates, including the intertidal mussel, Mytilus californianus. By virtue of their competitive dominance, large size, and longevity, M. californianus are ecosystem engineers that form structurally complex beds that provide habitat for diverse invertebrate communities. We investigated whether otters affect mussel bed characteristics (i.e. mussel length distributions, mussel bed depth, and biomass) and associated community structure (i.e. biomass, alpha and beta diversity) by comparing four regions that varied in their histories of sea otter occupancy on the west coast of British Columbia and northern Washington. Mussel bed depth and average mussel lengths were 1.5 times lower in regions occupied by otters for >20 years than those occupied for <5 yrs. Diversity of mussel bed associated communities did not differ between regions; however, the total biomass of species associated with mussel beds was more than three-times higher where sea otters were absent. We examined alternative explanations for differences in mussel bed community structure, including among-region variation in oceanographic conditions and abundance of the predatory sea star Pisaster ochraceus. We cannot discount multiple drivers shaping mussel beds, but our findings indicate the sea otters are an important one. We conclude that, similar to their effects on subtidal benthic invertebrates, sea otters reduce the size distributions of intertidal mussels and, thereby, habitat available to support associated communities. Our study indicates that by reducing populations of habitat-providing intertidal mussels, sea otters may have substantial indirect effects on associated communities.