Changes in Alaskan soft-bottom prey communities along a gradient in sea otter predation

Sea Otter (Enhydra lutris), well documented as "keystone" predators in rocky marine communities, were found to exert a strong influence on infaunal prey communities in soft-sediment habitats. Direct and indirect effects of sea otter predation on subtidal soft-bottom prey communities were evaluated along a temporal gradient of sea otter occupancy around the Kodiak Archipelago. The results indicate that Kodiak otters forage primarily on bivalve prey and dramatically reduce infaunal bivalve and green sea urchin (Strongylocentrotus droebachiensis) prey populations. Bivalve prey abundance, biomass, and size were inversely related to duration of sea otter occupancy. The relative conditions of shells discarded by otters in shallow (<10 m) vs. deep (> 20 m) water at the same sites indicate that otters first exploited Saxidomus in shallow-water feeding areas, and later switched to Macoma spp. in deeper water. Otter-cracked shells of the deep-burrowing clam Tresus capax were rarely found, even at otter foraging sites where the clam accounted for the majority of available prey biomass, suggesting that it has a partial depth refuge from otter predation. The indirect effects of otter predation included substratum disturbance and the facilitation of sea star predation on infaunal prey. Sea stars, Pycnopodia helianthoides, were attracted to experimentally dug excavations as well as natural sea otter foraging pits, where the sea stars foraged on smaller size classes of infaunal bivalves than those eaten by otters. Otters also discard clam shells on the sediment surface and expose old, buried shells during excavation. Surface shells were found to provide attachment sites for large anemones and kelp. Our study shows that sea otters can affect soft-sediment communities, not only through predation, as in rocky habitats, but also through disturbance, and thus retain a high degree of influence in two very different habitat types.