Two species of sea urchins (Colobocentrotus atratus and Echinometra mathaei) commonly co-occur on wave-swept intertidal shores in the Indo West Pacific. E. mathaei is a typical spiny urchin and is confined to cavities in the rock. In contrast, C. atratus has an unusual morphology, in which the spines are much reduced, and is found on substrata fully exposed to wave-induced velocities and accele...