Nautiloids
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Nautiloids
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. In a broad sense, "nautiloid" refers to a major cephalopod subclass or collection of subclasses (Nautiloidea ''sensu lato''). Nautiloids are typically considered one of three main groups of cephalopods, along with the extinct ammonoids (ammonites) and living coleoids (such as squid, octopus, and kin). While ammonoids and coleoids are monophyletic clades with exclusive ancestor-descendant rel ...
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Ellesmerocerida
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. Morphology The Ellesmerocerida are characterized by shells that are typically small, some even tiny, with close-spaced septa and relatively large ventral siphuncles. In some genera (e.g. ''Paleoceras''), the septa are uniformly spaced. Shells of ellesmerocerids are typically smooth and compressed and vary in form. They may be breviconic (short) or longiconic (elongate), straight (orthoconic) or curved (cyrtoconic). Cyrtoconic forms are usually endogastric, with longitudinally convex ventral margins. The apeces of straight forms typically have an endogastric curvature. Some may have grown to as much as 15 cm. Siphuncle segments are tubular or concave. Septal necks are short. Connecting rings which may appear layered are thick and typically wedge shaped with their maximum width at or near where t ...
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Plectronoceratoidea
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. Plectronoceratoidea contains several exclusively Cambrian cephalopod orders: Plectronocerida, Protactinocerida, and Yanhecerida. Under some classification schemes, the plectronoceratoid grade may also contain the paraphyletic Cambro-Ordovician Ellesmerocerida. A few older studies consider Plectronocerida to be descended from Ellesmerocerida (rather than ancestral), but this is no longer believed to be the case. The Plectronocerida is the earliest group of the four, and may have given rise to the other three. Of these four orders, only the Ellesmerocerida crossed (barely) into the Ordovician, with two genera, '' Ectenolites'' and '' Clarkoceras''. Ordovician ellesmerocerids in turn likely gave rise to severa ...
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Cephalopod
A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles ( muscular hydrostats) modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology. Cephalopods became dominant during the Ordovician period, represented by primitive nautiloids. The class now contains two, only distantly related, extant subclasses: Coleoidea, which includes octopuses, squid, and cuttlefish; and Nautiloidea, represented by '' Nautilus'' and '' Allonautilus''. In the Coleoidea, the molluscan shell has been internalized or is absent, whereas in the Nautiloidea, the external shell remains. About 800 living species of cephalopods have been i ...
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Orthoceratoidea
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 (a small number of large scars concentrated at the top of the body chamber), extensive cameral deposits, and calciosiphonate connecting rings with a porous and calcitic inner layer. Currently, Orthoceratoidea comprises the orders Riocerida, Dissidocerida, Actinocerida, Pseudorthocerida, Lituitida and Orthocerida. Orthocerida is a noteworthy paraphyletic order which is ancestral to the major cephalopod groups such as the extinct ammonoids and living coleoids (cephalopods without external shells, including squids, octopus, cuttlefish, etc.). Taxonomy As a superorder, Orthoceratiodea was one of six superorders within the Nautiloidea, the others being the Plectronoceratoidea (= E ...
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Nautilida
The Nautilida constitute a large and diverse order of generally coiled nautiloid cephalopods that began in the mid Paleozoic and continues to the present with a single family, the Nautilidae which includes two genera, ''Nautilus'' and '' Allonautilus'', with six species. All told, between 22 and 34 families and 165 to 184 genera have been recognised, making this the largest order of the subclass Nautiloidea. Classification and phylogeny Current classification The current classification of the Nautilida, in prevalent use, is that of Bernhard Kummel (Kummel 1964) in the Treatise which divides the Nautilida into five superfamilies, the Aipocerataceae, Clydonautilaceae, Tainocerataceae, and Trigonocerataceae, mostly of the Paleozoic, and the later Nautilaceae. These include 22 families and some 165 or so genera (Teichert and Moore 1964) Other concepts Shimansky 1962 (in Kummel 1964) divided the Nautilida into five suborders, the mostly Paleozoic Centroceratina, Liroceratina, Ru ...
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Endoceratoidea
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 . The related '' Cameroceras'' is anecdotally reported to have reached lengths approaching , 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. Morphology Endocerids had a relatively small body chamber as well as a proportionally large siphuncle, which in some genera reached nearly half the shell diameter. This suggests that much of the visceral mass may have been housed within the siphuncle itse ...
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Nautilus
The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus '' Nautilus''. Though it more specifically refers to species '' Nautilus pompilius'', the name chambered nautilus is also used for any of the Nautilidae. All are protected under CITES Appendix II. Depending on species, adult shell diameter is between 4 and 10 inches. Nautilidae, both extant and extinct, are characterized by involute or more or less convolute shells that are generally smooth, with compressed or depressed whorl sections, straight to sinuous sutures, and a tubular, generally central siphuncle.Kümmel, B. 1964. Nautiloidae-Nautilida, in the Treatise on Invertebrate Paleontology, Geological Society of America and Univ of Kansas Press, Teichert and Moore eds. Having survived rel ...
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Endocerida
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 . The related '' Cameroceras'' is anecdotally reported to have reached lengths approaching , 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. Morphology Endocerids had a relatively small body chamber as well as a proportionally large siphuncle, which in some genera reached nearly half the shell diameter. This suggests that much of the visceral mass may have been housed within the siphuncle i ...
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Tarphycerida
The Tarphycerida were the first of the coiled cephalopods, found in marine sediments from the Lower Ordovician (middle and upper Canad) 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,Furnish and Glenister 1964; Nautiloidea - Tarphycerida; In the ''Treatise on Invertebrate Paleontology'' Vol K; Teichert and Moore, (eds) GSA and U of Kansas Press 1964 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 i ...
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Monophyly
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic groups are typically characterised by shared derived characteristics (synapomorphies), which distinguish organisms in the clade from other organisms. An equivalent term is holophyly. The word "mono-phyly" means "one-tribe" in Greek. Monophyly is contrasted with paraphyly and polyphyly as shown in the second diagram. A ''paraphyletic group'' consists of all of the descendants of a common ancestor minus one or more monophyletic groups. A '' polyphyletic group'' is characterized by convergent features or habits of scientific interest (for example, night-active primates, fruit trees, aquatic insects). The features by which a polyphyletic group is differentiated from others are not inherited from a common ancestor. These definitions have take ...
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Octopus
An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the center point of the eight limbs. The soft body can radically alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates. Octopuses inhabit various regions of the ocean, including coral reefs, pelagic waters, and the seabed; some live in the intertidal zone and others at abyssal depths. Most species grow quickly, mature earl ...
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Orthocone
An orthocone is an unusually long straight shell of a nautiloid cephalopod.; During the 18th and 19th centuries, all shells of this type were named '' Orthoceras'', creating a wastebasket taxon, but it is now known that many groups of nautiloids developed or retained this type of shell. An orthocone can be thought of as like a ''nautilus'', but with the shell straight and uncoiled. It was previously believed that these represented the most primitive form of nautiloid, but it is now known that the earliest nautiloids had shells that were slightly curved. An orthoconic form evolved several times among cephalopods, and among nautiloid cephalopods is prevalent among the ellesmerocerids, endocerids, actinocerids, orthoceratoids, and bactritids. Orthocones existed from the Late Cambrian to the Late Triassic, but they were most common in the early Paleozoic. Revivals of the orthocone design later occurred in other cephalopod groups, notably baculitid ammonites in the Cretaceous P ...
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