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 species rich, 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). No orthoconic and only a handful of 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.
Cameroceras is an extinct genus of endocerid cephalopod which lived in equatorial oceans during the entire Ordovician period. Like other endocerids, it was an orthocone, meaning that its shell was fairly straight and pointed. It was particularly abundant and widespread in the Late Ordovician, inhabiting the shallow tropical seas in and around Laurentia, Baltica and Siberia.
Discosorida are an order of cephalopods that lived from the beginning of the Middle Ordovician, through the Silurian, and into the Devonian. Discosorids are unique in the structure and formation of the siphuncle, the tube that runs through and connects the camerae (chambers) in cephalopods, which unlike those in other orders is zoned longitudinally along the segments rather than laterally. Siphuncle structure indicated that the Discosorida evolved directly from the Plectronoceratida rather than through the more developed Ellesmerocerida, as did the other orders. Finally and most diagnostic, discosorids developed a reinforcing, grommet-like structure in the septal opening of the siphuncle known as the bullette, formed by a thickening of the connecting ring as it draped around the folded back septal neck.
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.
Clitendoceras is a genus of cephalopods in the order Endocerida from the Lower Ordovician with an elongate shell with a slight downward, endogastric, curvature and a siphuncle that lies along the ventral margin. Common for endocerids, the chambers are short and the septa close spaced with sutures sloping forward across the back of the shell. Septal necks are short in the young, lengthening in the adult. Endocones are simple, but with the ventral side of the last formed projecting forward. The endosiphotube running down the middle is arched on top and somewhat flat on the lower side.
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).
Baltoceratidae is an extinct family of orthoconic cephalopods belonging to the subclass Nautiloidea endemic to what would be Asia, Australia, Europe, North America, and South America during the Ordovician living from about 480–460 mya, existing for approximately 20 million years.
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.
Plectronoceratoidea is a superorder or subclass containing primitive nautiloids from the Late Cambrian and Early Ordovician. This group is best considered a paraphyletic grade of early cephalopods, as it contains the ancestors of subsequent post-Cambrian cephalopod orders.
Intejocerida is the name given to a group of generally straight shelled nautiloid cephalopods originally found in Lower and Middle Ordovician sediments in the Angara River basin in Russia; defined in the Treatise as an order, and combined there with the Endocerida in the Endoceratoidea.
Hectoceras is a genus in the nautiloid cephalopod order Discosorida from the Upper Ordovician of Australia (Tasmania), known from a few isolated siphuncle specimens.
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.
Cyrtocerinida is an order of Ordovician nautiloid cephalopods. The order includes the families Cyrtocerinidae and Eothinoceratidae, as well as the genera Bathmoceras and Rummoceras.
Yanhecerida is a small order of Late Cambrian nautiloid cephalopods. They were similar to the more diverse Plectronocerida and Ellesmerocerida, with short shells, closely spaced septa, and diaphragms (partitions) within the siphuncle. Their most distinctive trait is the conical shape of the diaphragms, similar to the endocones which characterize the later nautiloid order Endocerida. While some authors have argued that Yanhecerida should be lumped into Ellesmerocerida, a phylogenetic analysis has maintained that Yanhecerida is a valid clade of early cephalopods closely related to a paraphyletic Ellesmerocerida.
Multiceratoidea is a major subclass or superorder of Paleozoic nautiloid cephalopods. Members of this group can be characterized by nautilosiphonate connecting rings, with an organic inner layer and outer layer of calcitic spherules and blades, similar to the modern nautilus. The earliest-diverging multiceratoids have oncomyarian muscle scars, though several orders trend towards a ventromyarian condition. Multiceratoid shells are generally short and curled, with a relatively small aperture (opening). Cameral deposits are never found among the multiceratoids, though several orders are known to bear endosiphuncular deposits within their siphuncles.
Riocerida, originally named as Rioceratida, is an extinct order of Ordovician nautiloid cephalopods. They were the earliest-diverging members of the major group Orthoceratoidea, with which they share dorsomyarian muscle scars. However, they also possess several plesiomorphic ("primitive") traits which are absent in other orthoceratoids: Cameral and endosiphuncular deposits are absent, and the siphuncle has a marginal position along the lower edge of the shell. The conch shape is slender, usually orthoconic (straight) or weakly cyrtoconic. The monophyly of Riocerida as a unique group is plausible but difficult to confirm, and they may instead be a paraphyletic grade or polyphyletic assemblage ancestral to all later orthoceratoids.
