Phragmocone

Last updated May 02, 2019

Cutaway of a nautilus shell showing the chambers NautilusCutawayLogarithmicSpiral.jpg — Cutaway of a nautilus shell showing the chambers

The phragmocone is the chambered portion of the shell of a cephalopod. It is divided by septa into camerae.

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishermen sometimes call them inkfish, referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Camera (cephalopod) feature of cephalopods

Camerae are the spaces or chambers enclosed between two adjacent septa in the phragmocone of a nautiloid or ammonoid cephalopod. 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.

In most nautiloids and ammonoids, the phragmocone is a long, straight, curved, or coiled structure, in which the camarae are linked by a siphuncle which determines buoyancy by means of gas exchange.

Nautiloids are a large and diverse group of marine cephalopods (Mollusca) belonging to the subclass Nautiloidea that began in the Late Cambrian and are represented today by the living Nautilus and Allonautilus. Nautiloids flourished during the early Paleozoic era, where they constituted the main predatory animals, and developed an extraordinary diversity of shell shapes and forms. Some 2,500 species of fossil nautiloids are known, but only a handful of species survive to the present day.

Ammonoids are an extinct group of marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids than they are to shelled nautiloids such as the living Nautilus species. The earliest ammonites appear during the Devonian, and the last species died out in the Cretaceous–Paleogene extinction event.

Siphuncle Strand of tissue passing longitudinally through the shell of a cephalopod mollusk

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 walls dividing the chambers.

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.

The Silurian is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago (Mya), to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the shortest period of the Paleozoic Era. As with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by several million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when up to 60% of marine genera were wiped out.

The Ascocerida are comparatively small, bizarre Orthoceratoids known only from Ordovician and Silurian sediments in Europe and North America, uniquely characterized by a deciduous conch consisting of a longiconic juvenile portion and an inflated breviconic adult portion that separate sometime in maturity.

The PaleozoicEra is the earliest of three geologic eras of the Phanerozoic Eon. It is the longest of the Phanerozoic eras, lasting from 541 to 251.902 million years ago, and is subdivided into six geologic periods : the Cambrian, Ordovician, Silurian, Devonian, Carboniferous, and Permian. The Paleozoic comes after the Neoproterozoic Era of the Proterozoic Eon and is followed by the Mesozoic Era.

Fossil record

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]

Trilobites are a group of extinct marine arachnomorph arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period, and they flourished throughout the lower Paleozoic era before beginning a drawn-out decline to extinction when, during the Devonian, all trilobite orders except the Proetids died out. Trilobites disappeared in the mass extinction at the end of the Permian about 252 million years ago. The trilobites were among the most successful of all early animals, existing in oceans for over 300 million years.

Related Research Articles

Endocerida is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in its siphuncle.

Septa are thin walls or partitions between the internal chambers (camerae) of the shell of a cephalopod, namely nautiloids or ammonoids.

Orthocerida is an order of extinct Orthoceratoid cephalopods also known as the Michelinocerda 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. They were most common however from the Ordovician to the Devonian.

The body whorl is part of the morphology of the shell in those gastropod mollusks that possess a coiled shell. The term is also sometimes used in a similar way to describe the shell of a cephalopod mollusk.

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.

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.

Knightoconus antarcticus is a species of monoplacophoran from the Cambrian Minaret Formation of Antarctica. It is thought to represent an ancestor to the cephalopods. It had a chambered conical shell, but lacked a siphuncle. The absence of a siphuncle has been taken as evidence against cephalopod ancestry, as the factors that would influence the siphuncle to penetrate preexisting septal chambers remains unknown. The prevailing argument suggests that a strand of tissue remained attached to the previous septum as the mollusc moved forwards and deposited its next septum, stopping that new septum from closing completely and becoming mineralised itself. Ten or more septa are found in mature individuals, occupying around a third of the shell - septa form very early and have been found in specimens as small as 2 mm in length. Unlike monoplacophoran fossils, there is no evidence of muscle scarring in Knightoconus fossils. Scars from the closely related Hypseloconus have been used to determine its orientation. Knightoconus started life with an exogastric shell, that becomes endogastric as the organism grew.

Brevicoceras is an extinct nautiloid genus from the order Oncocerida with wide distribution in the Middle Devonian in Eastern North America, Russia and Morocco. Nautiloids form a broad group of shelled cephalopods that were once diverse and numerous but are now represented by only a handful of species in two genera.

Microbaltoceras is an extinct genus of the cephalopod family Baltoceratidae that lived in what would be North America during the Early Ordovician. It was named by Rousseau Flower (1964) and assigned by him to the Baltoceratidae. The type species is Microbaltoceras minore which was found in the Threadgill Member of the Tanyard Formation in Gillespie County, Texas, USA.

Sactoceras is an extinct nautiloid cephalopod that lived during the Ordovician and Silurian in what would become North America, Europe, and Asia.

The cephalopods have a long geological history, with the first nautiloids found in late Cambrian strata, and purported stem-group representatives present in the earliest Cambrian lagerstätten.

Shimanskya is a late Carboniferous fossil tentatively interpreted as an early spirulid.

Hematitida is a group of coleoid cephalopods known from the early Carboniferous Period. They are the oldest definite coleoids, although there are controversial claims for even older coleoids from the Devonian. Fossil hematitidans have so far been found only in Arkansas and Utah of the United States. The only family described so far is Hematitidae.

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–from arms-most to 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, 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.

References

↑ Henderson, ROBERT A.; McNamara, Kenneth J. (1985). "Taphonomy and ichnology of cephalopod shells in a Maastrichtian chalk from Western Australia". Lethaia. 18 (4): 305. doi:10.1111/j.1502-3931.1985.tb00710.x.
↑ Arnold Davis, R. H. B. Fraaye, Char, Richard (2001). "Trilobites within nautiloid cephalopods". Lethaia. 34 (1): 37. doi:10.1080/002411601300068251.

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[Henderson1985-1] Henderson, ROBERT A.; McNamara, Kenneth J. (1985). "Taphonomy and ichnology of cephalopod shells in a Maastrichtian chalk from Western Australia". Lethaia. 18 (4): 305. doi:10.1111/j.1502-3931.1985.tb00710.x.

[Davis2007-2] Arnold Davis, R. H. B. Fraaye, Char, Richard (2001). "Trilobites within nautiloid cephalopods". Lethaia. 34 (1): 37. doi:10.1080/002411601300068251.