The Scaphopoda.

Abstract

The Scaphopoda are marine infaunal carnivores that feed on foraminiferans and other microorganisms selected and manipulated by their unique feeding tentacles or captacula. Their tusk-like shell is open at both ends; the burrowing foot and captacula protrude anteriorly, while respiratory currents pass through the posterior opening. Although the scaphopods comprise one of the smallest molluskan classes in terms of species diversity, they have a worldwide distribution ranging from intertidal to depths in excess of 6000 m. Despite detailed monographic work from the late nineteenth and early twentieth centuries, however, the biology of scaphopods is today among the least understood of mollusks. To some extent, this is related to a significant departure from more typical molluscan organization inferred from studies of the larger classes. For example, the mantle cavity is elongate and restricted in space, and several associated organs are lost, including the osphradium and ctenidia; the latter is associated with the loss of auricles from the scaphopod heart. There is also little record of ontogeny in the adult shell, as the older portion is periodically decollated to maintain the posterior aperture for passage of inhalant and exhalant currents. This and other constraints on scaphopod diversification are reflected in the range limits of shell shape within the class. In contrast, the "Dentalium" egg has been a model system for experimental embryology, particularly in studies of cell lineage and early morphogenesis. Later larval development and most organogenesis is nonetheless poorly known, although recent studies have considerably enhanced our knowledge in this area. Global biogeographic patterns of scaphopod diversity have been studied only preliminarily. A general decrease in diversity with depth is broken by small diversity peaks at bathyal depths. There is also evidence for a marked latitudinal diversity gradient in the world's oceans, near equatorial in the Pacific but at approximately 20 degrees N in the Atlantic. Scaphopods have a wide diversity of ecto- and endo-symbiotic associations with other organisms, including commensal bacteria and ciliates, mutualistic anemones and corals, and parasitic algae and platyhelminths. Other documented associations include predation by naticid gastropods and ratfish, and the inhabitation of empty scaphopod shells by a variety of sipunculans and hermit crabs. Phylogeny within the class is still unresolved, although significant progress has been made recently in documenting morphological variation among families, genera, and species for application in a cladistic context. The evolutionary relationships of Scaphopoda within Mollusca are farther from resolution, with every conchiferan class a proposed sister group among modern analyses. Molecular avenues should provide significant progress in scaphopod phylogenetics, as indicated by recent comparative sequence analysis and gene expression patterns. Scaphopods comprise the most recent class of mollusks to appear in the fossil record, dating at least from the Mississippian Carboniferous (362.5 My). They appear from first-order occurrence data to have a substantially documented fossil record, with several sharp extinction and radiation events evident during their comparatively short geologic history. The unresolved position of the Scaphopoda within molluscan phylogeny, and their relatively limited range of morphological disparity but substantial derivation from other conchiferans, renders the scaphopods an enigmatic group. They represent a significant evolutionary avenue within the Mollusca, but one that is poorly examined. Nonetheless, comprehensive phylogenetic analysis and documentation of diversity within this small, monophyletic, class-level taxon, well-preserved in the fossil record, is imminently tractable, and presents a singular opportunity in the study of molluscan evolution.