Intertidal community structure : Experimental studies on the relationship between a dominant competitor and its principal predator

Abstract

Along exposed rocky intertidal shorelines of western North America the mussel Mytilus californianus exists as a characteristic, well-defined band. Measurements at Mukkaw Bay and Tatoosh Island, Washington State, suggest that the upper limit to distribution is constant. The lower limit is also predictably constant, as judged by photographs of the same areas taken up to 9 years apart. The band of mussels is formed by larval recruitment to a variety of substrates, especially the filamentous red alga Endocladia muricata. From the settlement site, if the mussels survive a series of predators including the starfish Pisaster ochraceus and a variety of carnivorous gastropods (Thais spp.), the mussles may be washed inward or migrate (be pushed) downward.When Pisaster was removed manually, the zonation pattern changed rapidly. Mussels advanced downward at Mukkaw Bay a vertical distance of 0.85 m in 5 years. No movement was observed on 2 adjacent control sites. At Tatoosh Island a maximum displacement of 1.93 m has been observed in 3 years; the slope there is 40°. Again, there was no change at control sites with Pisaster. At Mukkaw Bay over 25 species of invertebrates and benthic algae are excluded from occupancy of the primary substratum by mussels. The ecological dominance of mussels is discussed; predation is shown to enhance coexistence among potential competitors. A survival curve for Pollicipes polymerus indicates that the time course for interspecific competitive exclusion may be long (76 months). The clarity of the biological interrelationships and the constancy of pattern through time provide no support for the contention that intertidal communities are physically-controlled.