Development of Cerata in the Cladobranchian Sea Slug Pteraeolidia semperi (Mollusca: Gastropoda: Nudibranchia).

Abstract

Cladobranchian sea slugs are characterized by a number of dorsal projections, called "cerata," which are presumably involved in such biological functions as kleptocnidal defense, gas exchange, and symbiotic photosynthesis. Here, we investigated the developmental pattern of ceras formation in a cladobranchian, Pteraeolidia semperi, using field-collected individuals at various postembryonic developmental stages. As the body length increased, the total number of cerata increased in a logistic manner, up to 280 per individual. On the dorsal side of the body, the cerata exhibited a conspicuous formation of repeated, laterally-paired clusters, or rows, along the antero-posterior axis of the animals. As the body length increased, the number of ceras rows increased in a logistic manner, reaching a plateau at around 15 rows per individual. Two types of ceras clusters were observed: well-developed ceras clusters forming a glove-like structure with a basal bulge, which tended to be found in larger animals and at the anterior body region, and less-developed ceras clusters without the bulge, which tended to be found in smaller animals and at the posterior body region. Statistical and simulation analyses suggested that bulge formation underlies increased ceras number, even after the plateaued formation of new ceras rows. These results indicate that, in the postembryonic development of P. semperi, the increase of dorsal cerata entails the following processes: (i) increase of the number of ceras rows, (ii) formation of the basal bulge in each ceras cluster, and (iii) increase of the number of cerata per ceras cluster.