The phragmocone is the chambered portion of the shell of a cephalopod. It is divided by septa into camerae.
In most nautiloids and ammonoids, the phragmocone is a long, straight, curved, or coiled structure, in which the camerae are linked by a siphuncle which determines buoyancy by means of gas exchange.
Despite this benefit, such a large shell adds to the mass of the animal, and hence is disadvantageous in catching fast-moving prey. Some nautiloids, such as the Silurian Ascocerida, dropped the phragmocone upon maturity, presumably to increase speed and maneuverability. They thus became the early Paleozoic equivalent of coleoids. The early coleoids and belemnoids adopted a different approach: the phragmocone was retained but became internal and reduced. In general the shell in cephalopods tends to be vestigial or absent.
Being the only biomineralised part of most cephalopods, the phragmocone is typically the only part to enter the fossil record. It is sometimes infilled with sediment, with sediment presumably getting in through the siphuncle. [1] There are occasions where trilobites have been preserved within phragmocones, presumably where they crawled in for refuge. [2]
Ammonoids are extinct spiral shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living coleoids than they are to shelled nautiloids. The earliest ammonoids appeared during the Devonian, with the last species vanishing during or soon after the Cretaceous–Paleogene extinction event. They are often called ammonites, which is most frequently used for members of the order Ammonitida, the only living group of ammonoids from the Jurassic up until their extinction.
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.
Septa are thin walls or partitions between the internal chambers (camerae) of the shell of a cephalopod, namely nautiloids or ammonoids.
Camerae are the spaces or chambers enclosed between two adjacent septa in the phragmocone of a nautiloid or ammonoid cephalopod molluscus. These can be seen in cross-sections of a nautilus shell and in the polished cross-sections of ammonites. In life these chambers are filled with gas, mediated by the siphuncle, and used to control buoyancy.
Orthocerida, also known as the Michelinocerida, is an order of extinct orthoceratoid cephalopods that lived from the Early Ordovician possibly to the Late Triassic. A fossil found in the Caucasus suggests they may even have survived until the Early Cretaceous, and the Eocene fossil Antarcticeras is sometimes considered a descendant of the orthocerids although this is disputed. They were most common however from the Ordovician to the Devonian.
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.
Aulacocerida is an order of primitive coleoid cephalopods, possibly derived from michelinoceraitids (Orthocerida) early in the Devonian, which in turn gave rise to the Belemnites.
Plectronocerida is a primitive order from which subsequent cephalopod orders are ultimately derived.
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.
Boletzkyida is a primitive order of teuthid coleoid cephalopod: the boletzkyids are thought to be the earliest forms of coleoid cephalopods, and appear to form a link between nautiloid orthocerids and more advanced coleoids. Boletzkyida was named and described by Bandel, Reitner, and Sturmer in 1983 (B.R.&S) from specimens found in the Lower Devonian black slate in Germany.
Wardoceras is an extinct nautiloid genus from the late Early Ordovician of Western Utah, assigned to the orthocerid family, Michelinoceratidae
Sactoceras is an extinct nautiloid cephalopod that lived during the Ordovician and Silurian in what would become North America, Europe, and Asia.
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.
The cephalopods have a long geological history, with the first nautiloids found in late Cambrian strata.
Shimanskya is a late Carboniferous fossil tentatively interpreted as an early spirulid.
Phragmoteuthida is an order of extinct coleoid cephalopods characterized by a fan-like teuthoid pro-ostracum attached to a belemnoid-like phragmocone.
Belemnitida is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. The parts are, from the arms-most to the tip: the tongue-shaped pro-ostracum, the conical phragmocone, and the pointy guard. The calcitic guard is the most common belemnite remain. Belemnites, in life, are thought to have had 10 hooked arms and a pair of fins on the guard. The chitinous hooks were usually no bigger than 5 mm (0.20 in), though a belemnite could have had between 100 and 800 hooks in total, using them to stab and hold onto prey.
Antarcticeras is an extinct genus of enigmatic cephalopod from the Eocene of Antarctica. It contains a single species, A.nordenskjoeldi. It is either considered the last of the "orthocone"-type cephalopods, the only member of its subclass Paracoleoidea & a descendant of the orthoceratids, and a remarkable example of convergent evolution with coleoid cephalopods, or an oegospid squid and a transitional form in the development of the modern squid gladius, of which it is the only preserved example.
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