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The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and Spirula. In the case of the cuttlefish, the siphuncle is indistinct and connects all the small chambers of that animal's highly modified shell; in the other cephalopods it is thread-like and passes through small openings in the septa (walls) dividing the camerae (chambers). Some older studies have used the term siphon for the siphuncle, though this naming convention is uncommon in modern studies to prevent confusion with a mollusc organ of the same name.
Nautiloids are a group of marine cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living Nautilus and Allonautilus. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species. They flourished during the early Paleozoic era, when they constituted the main predatory animals. Early in their evolution, nautiloids developed an extraordinary diversity of shell shapes, including coiled morphologies and giant straight-shelled forms (orthocones). Only a handful of rare coiled species, the nautiluses, survive to the present day.
Endocerida is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. Endocerida was a diverse group of cephalopods that lived from the Early Ordovician possibly to the Late Silurian. Their shells were variable in form. Some were straight (orthoconic), others curved (cyrtoconic); some were long (longiconic), others short (breviconic). Some long-shelled forms like Endoceras attained shell lengths close to 6 metres (20 ft). The related Cameroceras is anecdotally reported to have reached lengths approaching 9 metres (30 ft), but these claims are problematic. The overwhelming majority of endocerids and nautiloids in general are much smaller, usually less than a meter long when fully grown.
Endoceras is an extinct genus of large, straight shelled cephalopods from the Middle and Upper Ordovician that gives its name to the Nautiloid order Endocerida. The cross section in the mature portion is slightly wider than high, but is narrower laterally in the young. Sutures are straight and transverse. Endoceras has a large siphuncle, located close to the ventral margin, composed of concave segments, especially in the young but which may be tubular in the adult stage. Endocones are simple, subcircular in cross section, and penetrated by a narrow tube which may contain diaphragms reminiscent of the Ellesmerocerid ancestor.
The Ellesmerocerida is an order of primitive cephalopods belonging to the subclass Nautiloidea with a widespread distribution that lived during the Late Cambrian and Ordovician.
The Proterocameroceratidae were the first of the Endocerida. They began early in the Ordovician with Proendoceras or similar genus which had developed endocones, replacing the diaphragms of the ellesmerocerid ancestor.
Bathmoceras is a primitive cephalopod genus from the Middle and Upper Ordovician. It is a member of the order Cyrtocerinida and is the only genus in the family Bathmoceratidae.
The Tarphycerida were the first of the coiled cephalopods, found in marine sediments from the Lower Ordovician to the Middle Devonian. Some, such as Aphetoceras and Estonioceras, are loosely coiled and gyroconic; others, such as Campbelloceras, Tarphyceras, and Trocholites, are tightly coiled, but evolute with all whorls showing. The body chamber of tarphycerids is typically long and tubular, as much as half the length of the containing whorl in most, greater than in the Silurian Ophidioceratidae. The Tarphycerida evolved from the elongated, compressed, exogastric Bassleroceratidae, probably Bassleroceras, around the end of the Gasconadian through forms like Aphetoceras. Close coiling developed rather quickly, and both gyroconic and evolute forms are found in the early middle Canadian.
Anthoceras is a genus of straight, annulated, proterocamerioceratids from the Lower Ordovician, found in North America, NW Australia, and Siberia. The cross section is circular, the siphuncle moderately large, and marginal. Segments are constricted ; septal necks hemichoantici to subholochoantic ; connecting rings thick. Endocones are long and slightly asymmetric.
The Ellesmeroceratidae constitute a family within the cephalopod order Ellesmerocerida. They lived from the Upper Cambrian to the Lower Ordovician. They are characterized by straight and endogastric shells, often laterally compressed, so the dorso-ventral dimension is slightly greater than the lateral, with close spaced sutures having shallow lateral lobes and a generally large tubular ventro-marginal siphuncle with concave segments and irregularly spaced diaphragms. Connecting rings are thick and layered, externally straight but thickening inwardly with the maximum near the middle of the segment so as to leave concave depressions on internal siphuncle molds. Septal necks are typically orthochoanitic but vary in length from almost absent (achoanitic) to reaching halfway to the previous septum (hemichoanitic) and may even slope inwardly (loxochoanitic).
The Trigonoceratoidea are a superfamily within the Nautilida that ranged from the Devonian to the Triassic, thought to have contained the source for the Nautilaceae in which Nautilus is found.
Orthoceratoidea is a major subclass of nautiloid cephalopods. Members of this subclass usually have orthoconic (straight) to slightly cyrtoconic (curved) shells, and central to subcentral siphuncles which may bear internal deposits. Orthoceratoids are also characterized by dorsomyarian muscle scars, extensive cameral deposits, and calciosiphonate connecting rings with a porous and calcitic inner layer.
Endoceratidae is a family of large to very large straight shelled nautiloid cephalopods belonging to the order Endocerida that lived during the Middle and Late Ordovician. They include the largest known Paleozoic invertebrates, represented by Endoceras and Cameroceras.
Chazyoceras is a moderately large endocerid included in the Endoceratidae with a Nanno type apex and a ventral siphuncle with a holochoanitic wall, characteristic of the family. The siphuncle swelling at the apex is subtriangular in longitudinal profile. The endocones are of medium length.
Bajkaloceras is a straight-shelled orthoceroid, and possibly a member of the Intejocerida, from the Angara River basin in central Russia, named by Balashov in 1962. Its age, as given in the Treatise on Invertebrate Paleontology is Arenigian.
Proterovaginoceras is a medium to large sized endocerid from the Early and Middle Ordovician included in the family Endoceratidae.
Cyrtogomphoceras is a genus of nautiloid cephalopods, recognized by its large breviconic shell with a notable endogastric curvature. The shell is fusiform in profile, reaching maximum width at or near the base of body chamber, which narrows toward the aperture. The siphuncle is large and slightly removed from the ventral side, that with the concave longitudinal profile. Siphuncle segments are short, as are chambers; septal necks recurved, connecting rings thick, bullettes at the apical end of the rings swollen. Cameral deposits are lacking.
Parryoceras is a cyrtogomphoceratid similar to Strandoceras but with broadly expanded siphuncle segments and a nearly straight ventral line. The mature shell is thickened on the inside just behind the aperture resulting in a constriction in the internal mold at the adoral end.
The Allotrioceratidae is a family of Middle Ordovician fossils, established by Rousseau Flower, 1955, originally including Allotrioceras and Mirabilocras, assigned inferentially to the Endocerida and known only from structures interpreted as siphuncles. Later Williamsoceras, Cacheoceras, and Perkinsoceras were added.
Bisonocerida is an order of Ordovician to Silurian nautiloid cephalopods. Members of this order were originally placed in the order Endocerida, but later investigation argued that this broad usage of Endocerida was a polyphyletic assemblage encompassing two different groups of independent origin. Bisonocerida was differentiated from Endocerida in 2012 in order to resolve this issue.
References
- Flower, R.H. 1976. Some Whiterock and Chazy Endoceroids. Part II, Mem.28; New Mexico Bureau of Mines and Mineral Resources, 1976.
- Techert, C. 1964. Endoceratoidea. Treatise on Invertebrate Paleontology, Park K. Teichert & Moore (eds). Geological Society of America and Univ Kansas Press.