Protective skin structures in shell-less marine gastropods

Skin and stomach epithelia of the four suborders of nudibranch gastropods (Mollusca, Opisthobranchia) are characterized by large intracellular ovoid disks, here called spindles. These spindles are an evolutionary novelty in the nudibranchs; in shell-less sacoglossan species they are missing. We here examined whether the distribution and occurrence of the spindles is consistent with the assumption of a protective role against discharging nematocysts of co-habiting and prey Cnidaria. Spindles were abundant in epidermal cells of regions exposed to nematocysts, such as the cerata, the lips, the edges of lamellate rhinophores, the surfaces of non-retractile gill-like organs, as well as in the stomach epithelium of eolid and dendronotacean species which feed on Cnidaria. While cells packed with spindles almost exclusively formed the epidermis of eolid and arminacean species, they were less numerous in the skin of dendronotacean and dorid species, where glandular cells predominated. The preponderance of either glandular or spindle cells suggests a dual complementary defense strategy, on the one side the production of mucus coats and aversive glandular secretions, on the other structural defensive devices that are cushion-like entities filled with inert grains.