Factors determining the distribution of the sea pansy, Renilla kollikeri, in a subtidal sand-bottom habitat.

The biological and physical factors that affect the distribution of the subtidal, epibenthic, sand-dwelling cnidarian Renilla kollikeri along a depth gradient from 2 to 13 m were determined by manipulative field and laboratory experiments. The sea pansy's nearshore distributional limit (2.5 m in depth) was set by the animal's inability to remain anchored on the bottom in the face of increasing wave surge associated with shallow water. Renilla was outcompeted for space by the sand dollar Dendraster excentricus in an area between 6 and 9 m in depth and was largely excluded from the substrate when sand dollar densities exceeded 350/m2. Predator-prey interactions also influence the Renilla distribution. The asteroid Astropecten brasiliensis feeds on and removes Renilla 40 mm and less in diameter from areas seaward of the sand dollar bed (9 m in depth). Astropecten did not forage in the shallow area where Renilla was most abundant because of its intolerance to wave surge and because of its difficulty in crossing the sand dollar bed. The nudibranch Armina californica fed on Renilla throughout the depth gradient but did not remove colonies greater than 20 mm in diameter because of the Armina- specific escape behavior of Renilla. The foraging behavior of Armina was also curtailed by strong surge conditions. The nearshore area (3-6 m in depth) of high Renilla density was a refuge in which animals that had settled from the plankton could escape predation. As the animals grow they lose their ability to anchor in the substrate in shallow water and thus move into deeper water. The results of multiway interactions among sets of three species, particularly the beneficial results of the competitive interaction with Dendraster in light of the activities of the predators are discussed with reference to the abundance and distribution of Renilla.