Adding teeth to wave action: the destructive effects of wave-borne rocks on intertidal organisms.

Observations in rocky intertidal areas demonstrate that breaking waves 'throw' rocks and cobbles and that these missiles can damage and kill organisms. Targets in the intertidal were dented by impacts from wave-borne rocks. New dents/day in these targets was positively correlated with the daily maximum significant wave height and with new patches/day in aggregations of the barnacle Chthamalus fissus. Impact frequency was highest in the upper intertidal and varied dramatically between microhabitats on individual boulders (edges, tops and faces). These patterns were reflected in the microhabitat abundances of 'old' and 'young' barnacles. Comparisons were made of the survivorship and the frequency of shell damage in two populations of the limpet Lottia gigantea living in habitats which differed primarily in the number of moveable rocks (i.e. potential projectiles). The mortality rate and frequency of shell damage were significantly higher in the projectilerich habitat. In addition only in this habitat did the frequency of shell damage covary significantly with seasonal periods of high surf. Investigation of the response of limpet shells to impacts suggests that shell strength varies between species and increases with shell size. Species-specific patterns of non-fatal shell breakage may have evolved to absorb the energy of impacts. In two of the intertidal habitats studied, wave-borne rock damage was chronic and, at least in part, may have governed the faunal makeup of the community by contributing to the physical 'boundaries" of the environment within which the inhabitants must survive.