Mechanisms and consequences of shell fouling in the kelp snail,Norrisia norrisi (Sowerby) (Trochidae): Indirect effects of octopus drilling

The kelp snail, Norrisia norrisi (Sowerby), dwells on the large kelps Macrocystis pyrifera (L.) and Eisenia arborea Areschoug, and is rare on benthic substrata. Approximately 4% of the snail population is dislodged from plants each day. Per capita mortality of snails on the benthos is an order of magnitude greater than individuals on kelp. Even though snails displaced to the bottom actively seek a plant to climb, we found that between 20–40% of marked snails released on the bottom were killed over a 24-day period. While most N. norrisi are free of epibionts on their shells, ≈30% of the population are encrusted by the barnacle Megabalanus californicus (Pilsbry). The majority of barnacle fouling was the result of failure by Octopusbimaculatus (Verrill) to kill Norrisia norrisi; barnacle cyprids settled in octopus drill holes in the shells of live kelp snails. N. norrisi encrusted by barnacles were less effective in exploiting kelp compared with unfouled snails. Moderately to heavily fouled snails were dislodged from kelp twice as frequently and remained on the benthos much longer than snails without epibionts. Laboratory tests revealed that escape velocities of N. norrisi from asteroid predators were much reduced by shell fouling, and that snails carrying barnacles were most likely to be captured and consumed. Hence octopus increase mortality of kelp snails in two ways. First, octopus can kill N. norrisi encountered on the bottom. Second, kelp snails drilled by octopus, but not killed outright, will become fouled by barnacles. This results in a subsequent risk of mortality from all benthic predators that is at least 8-fold greater than for unfouled kelp snails.