Does a shell matter for defence? Chemical deterrence in two cephalaspidean gastropods with calcified shells

Opisthobranch molluscs show an evolutionary trend to reduce, internalize and lose the shell. Many of them base their defensive strategies on natural deterrent products and current evolutionary theory suggests that the acquisition of chemical defences preceded shell reduction and loss, which has characterized the evolution of this group. Here we show that basal, shelled opisthobranch molluscs are defended against sympatric predators even if their protective shell is removed. The cephalaspideans Bulla striata and Haminoea orbignyana, both with distinct shell calcification, significantly deterred feeding by sympatric crab and fish predators, both in laboratory and field assays. However, our results argue against a progressive increment of chemical defences associated with shell reduction, because the cephalaspidean with the more fully calcified shell, Bulla striata, was also the more deterrent. These findings suggest that effective chemical defences might have evolved independently from shell loss, at least in basal opisthobranchs such as cephalaspideans