The functional anatomy of the mantle complex and columellar muscle of tectibranch molluscs (Gastropoda: Opisthobranchia), and its bearing on the evolution of opisthobranch organization.

Abstract

An account is given of the anatomy of a series of opisthobranch molluscs principally to assess the change in importance and functioning of the mantle cavity and columellar muscle throughout the transition from prosobranch to opisthobranch organization. Intermediate steps are represented by living tectibranchs, of which Philine and Scaphander are investigated in detail, Acteon, Bulla, Haminoea, Akera, Aglaja and Gastropteron more briefly. Though an opisthobranch, Acteon has an organization typical of a monotocardian prosobranch; the remainder show trends affecting the shell and visceral mass, mantle cavity and head-foot, which resulted finally in the production of nudibranch types. It is confirmed that the adaptations exhibited by primitive tectibranchs relate to the assumption of a burrowing mode of life. Initial changes were the reduction of the nuchal area and sealing of the mantle cavity anteriorly so that it opened on the right, where it became restricted, the first perhaps prompting the sealing. A broadening and an anterior elongation of the head-foot produced a wedge to facilitate burrowing. Change in disposition of the mantle edge, incurred by differential growth, produced an involute shell with a large body whorl, alignment changing from erect to horizontal. The resultant streamlining eased infaunal progression; no vertical insinking of the viscera was involved. Subsequently the shell became reduced and finally lost. A section of the mantle edge enlarged to produce a posterior mantle lobe upon which sit both the shell and viscera, and which later became redundant as posterior elongation of the head-foot produced a slug-like form, the viscera being incorporated within the head-foot. As the nuchal area became reduced, mechanical needs prompted alteration to both the form and attachment of the columellar muscle. In Acteon the muscle is like that of a prosobranch, but the proximal region has broadened, a change of proportion required by primitive tectibranchs in order to support the floor of the mantle cavity formed from the section of mantle skirt which in prosobranchs lies on the right. This was followed by reduction and re-alignment of the entire muscle along an anteroposterior axis as emphasis changed from the muscle effecting retraction into a shell to producing contorsions of the head-foot. The shell, similarly reduced, instead of providing anchorage, became itself anchored by additional anterior and posterior attachment zones with, in more advanced forms, dorsoventral muscles of the body wall rather than longitudinal muscles fastening to the former. Importance was placed on the mutual stabilization of constituent parts of the posterior body region. Re-alignments of the muscle induced breaking up of the longitudinal muscle sheet of the head-foot to produce muscle tracts, best exhibited in those tectibranchs which swim; they are derived from both the columellar muscle and intrinsic body wall muscles...