Temporal range: Mississippian–Recent
|A shell of the scaphopod Antalis vulgaris from France|
The tusk shells or scaphopods // (Greek, "boat-footed") are a class of shelled marine mollusc with worldwide distribution, and are the only exclusively class of infaunal marine molluscs. Shells of species within this class range from about 0.5 to 15 cm in length. Members of the order Dentaliida tend to be significantly larger than those of the order Gadilida.
These molluscs live in soft substrates offshore (usually not intertidally). Because of this subtidal habitatand the small size of most species, many beachcombers are unfamiliar with them; their shells are usually not as common or as easily visible in the beach drift as the shells of sea snails and clams.
The mantle of a scaphopod is entirely within the shell. The foot extends from the larger end of the shell, and is used to burrow through the substrate. The scaphopod positions itself head down in the substrate, with the apical end of the shell (at the rear of the animal's body) projecting upward. This end seldom appears above the level of the substrate, however, as doing so exposes the animal to numerous predators. Most adult scaphopods live their lives entirely buried within the substrate.
Water enters the mantle cavity through the apical aperture, and is wafted along the body surface by cilia. There are no gills; the entire surface of the mantle cavity absorbs oxygen from the water. Unlike most other molluscs, there is no continuous flow of water with a separate exhalant stream. Instead, deoxygenated water is expelled rapidly back through the apical aperture through muscular action once every ten to twelve minutes.
A number of minute tentacles around the foot, called captacula, sift through the sediment and latch onto bits of food, which they then convey to the mouth. The mouth has a grinding radula that breaks the bit into smaller pieces for digestion. The radulae and cartilaginous oral bolsters of the Gadilidae are structured like zippers where the teeth actively crush the prey by opening and closing on it repeatedly, while the radulae and bolsters of the Dentaliidae work rachet-like to pull the prey into the esophagus, sometimes whole. The massive radula of the scaphopods is the largest such organ relative to body size of any mollusc (among whom, except for the bivalves, the presence of a which is a defining characteristic). The remainder of the digestive system consists of a digestive diverticulum, esophagus, stomach, and intestine. A digestive gland secretes enzymes into the stomach, but, unlike some other molluscs, does not digest the food directly itself. The anus opens on the ventral/ underside of the animal, roughly in the middle of the mantle cavity.
The scaphopod vascular system is rudimentary, lacking both heart and blood vessels; the blood is held in sinuses throughout the body cavity, and is pumped through the body by the rhythmic action of the foot. Metabolic waste is excreted through a pair of nephridia close to the anus.
The nervous system is generally similar to that of gastropods. One pair each of cerebral and pleural ganglia lie close to the oesophagus, and effectively form the animal's brain. A separate set of pedal ganglia lie in the foot, and a pair of visceral ganglia are set further back in the body. Scaphopods have no eyes, or other distinct sensory organs.
Reproduction and development
The group is composed of a two subtaxa, the Dentaliida (which may be paraphyletic) and the monophyletic Gadilida. The differences between the two orders is subtle and hinges on size and on details of the radula, shell, and foot. Specifically, the Dentaliids are the physically larger of the two families, and possess a shell that tapers uniformly from anterior (widest) to posterior (narrowest); they also have a foot which consists of one central and two lateral lobes and which bends into the shell when retracted. The Gadilids, on the other hand, are much smaller, have a shell whose widest portion is slightly posterior to its aperture, and have a foot which is disk-like and fringed with tentacles which inverts into itself when retracted (in this state resembling a pucker rather than a disk).
There is a good fossil record of scaphopods from the Mississippian onwards, making them the youngest molluscan class; but the origin of the group remains contentious.
The Ordovician Rhytiodentalium kentuckyensis has been interpreted as an early antecedent of the scaphopods, implying an evolutionary succession from ribeirioid rostroconch molluscs such as Pinnocaris. However, a competing hypothesis suggests a Devonian/Carboniferous origin from a non-mineralized ancestor, or from a more derived, Devonian, conocardioid rostroconch.
The shells of Dentalium hexagonum, a scaphopod mollusc, were strung on thread and used by the natives of the Pacific Northwest as shell money. Dentalium shells were also used to make belts and headdresses by the Natufian culture of the Middle East, and are a possible indicator of early social stratification.
Shells of the species Dentalium pretiosum were used as money.
- Steiner, G. . (1992). "Phylogeny and Classification of Scaphopoda". Journal of Molluscan Studies 58 (4): 385. doi:10.1093/mollus/58.4.385.
- Reynolds, Patrick D. (2002). "The Scaphopoda". Advances in marine biology. Advances in Marine Biology 42: 137–236. doi:10.1016/S0065-2881(02)42014-7. ISBN 9780120261420. PMID 12094723.
- Giribet, G.; Okusu, A, A.; Lindgren, A.R., A. R.; Huff, S.W., S. W.; Schrödl, M, M.; Nishiguchi, M.K., M. K. (May 2006). "Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons" (FREE FULL TEXT). Proceedings of the National Academy of Sciences of the United States of America 103 (20): 7723–7728. Bibcode:2006PNAS..103.7723G. doi:10.1073/pnas.0602578103. PMC 1472512. PMID 16675549.
- Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 432–434. ISBN 0-03-056747-5.
- Guralnick, R.; Smith, K. (1999). "Historical and biomechanical analysis of integration and dissociation in molluscan feeding, with special emphasis on the true limpets (Patellogastropoda: Gastropoda)". Journal of Morphology 241 (2): 175–195. doi:10.1002/(SICI)1097-4687(199908)241:2<175::AID-JMOR7>3.0.CO;2-0. PMID 10420163.
- Ellis L. Yochelson; Royal H. Mapes; Doris Heidelberger. "An enigmatic molluscan fossil from the Devonian of Germany: scaphopod or cephalopod?". Paläontologische Zeitschrift 81 (2): 118–122. doi:10.1007/BF02988386.
- Peel, J.S. (2004). "Pinnocaris and the origin of scaphopods.". Acta Palaeontologica Polonica 49 (4): 543–550.
- Crabtree, Pam J.; Campana, Douglas V. (eds.). Exploring Prehistory: How Archaeology Reveals Our Past (2nd ed.). p. 233. ISBN 0-07-297814-7.
For a comprehensive overview, see Reynolds, P. D. (2002). "The scaphopoda". Molluscan Radiation - Lesser-known Branches. Advances in Marine Biology 42. pp. 137–236. doi:10.1016/S0065-2881(02)42014-7. ISBN 9780120261420. PMID 12094723. .