Gastropod opercula are classified here on a new morphogenetic basis, which distinguishes three main types: (1) flexiclaudent spiral (mostly multispiral) operculum, the shape of which does not coincide with that of the aperture, (2) rigiclaudent spiral (usually paucispiral) operculum, the shape of which fits that of the aperture, and (3) rigiclaudent concentric operculum, also aperture-fitting. The first type fits by flexing into the aperture and is secreted when the soft parts are partly or wholly extended (i.e., when the operculum is not in a closed position). The other two types do not flex upon retraction (except at the very margin) and grow when the operculum closes over the aperture, with or without rotation. A study of opercular types at the family level confirms the systematic and evolutionary significance of opercula. Types 1 and 2 are the only ones present in archaeogastropods, Type 1 being predominant. Opercula (if present) in Neritopsina are always rigiclaudent. Within Caenogastropoda, Type 2 predominates; the only flexiclaudent spiral opercula are found in certain basal cerithioidean families. Concentric opercula are predominant in higher neotaenioglossans and exclusive in neogastropods. Except for one family, opercula in Heterostropha are always rigiclaudent spiral. Morphological, systematic, and histological criteria point to the flexiclaudent spiral operculum as the ancestral form. This leads us to propose the “periostracum shaving” model in prosobranchs to account for the origin of this kind of operculum. According to this model, in the earliest trochospiral gastropods the periostracum ceased to serve a shell-formation function at the band of overlap between whorls (the parietal band). The periostracal band was then extruded from the shell to constitute an incipient operculum, taking on the appearance of a spiral strip coiling opposite to the shell. The parietal segment of the periostracal groove migrated toward the epipodium and became independent from the rest of the mantle. The concomitant development of an opercular disc allowed the successive turns of periostracal strip to seal together. In this way, a spiral operculum emerged, coiling counterclockwise without matching the aperture shape. During the course of prosobranch evolution, rigiclaudent spiral opercula emerged several times from the ancestral flexiclaudent type, although they were always restricted to apertures with a spiral-shaped outer (labral) edge. Such opercula enlarged the range of shell morphologies for which the operculum constituted an efficient protective barrier to include those of neritoidean or naticoidean type. The onset of calcification in opercula took place with the rigiclaudent type. Concentric opercula also evolved independently from rigiclaudent spiral opercula in several gastropod groups, thus further broadening the spectrum of apertures and, hence, of shell morphologies using opercula for protection. From the standpoint of adaptation, the concentric type was probably the only one available to neogastropods having long and wide siphonal canals.
Origin of the holothurians (Echinodermata) derived by constructional morphology
