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Ellesmeroceratoidea
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 several new ...
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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 ident ...
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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 (= Ellesm ...
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Plectronocerida
Plectronocerida is a primitive order from which subsequent cephalopod orders are ultimately derived.Curt Teichert, 1988. Main Features of Cephalopod Evolution. The Mollusca Vol. 12 Paleontology and Neontology of Cephalopds; Academic Pres Inc. Occurrence Plectronoceratids are known from the Upper Cambrian (upper Franconian – middle Trempealeauan) of China and North America (Minnesota, Wisconsin). Two families are recognized (Flower, 1964),Flower, Rousseau H. 1964. The Nautiloid Order Ellesmeroceratida (Cephalopoda); Memoir 12, New Mexico Bureau of Mines and Mineral Resources, Socorro, New Mexico the generally straight to endogastric Plectronoceratidae and the slightly exogastric Balkoceratidae. Diagnostic characters Members of the Plectronocerida are characterized as follows. Shells are generally small, some even tiny, laterally compressed, curved (cyrtochonic) or straight (orthoconic). Most cyrtoconic forms are endogastric, with the ventral side longitudinally concave, or t ...
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Protostome Superorders
Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's members, although the reverse is typically true of its sister clade, Deuterostomia. Well known examples of protostomes are arthropods, molluscs, annelids, flatworms and nematodes. They are also called schizocoelomates since schizocoely typically occurs in them. Together with the Deuterostomia and Xenacoelomorpha, these form the clade Bilateria, animals with bilateral symmetry, anteroposterior axis and three germ layers. Protostomy In animals at least as complex as earthworms, the first phase in gut development involves the embryo forming a dent on one side (the blastopore) which deepens to become its digestive tube (the archenteron). In the sister-clade, the deuterostomes (), the original dent becomes the anus while the gut eventually tun ...
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Multiceratoidea
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 (with numerous small muscle attachments ringing the body chamber), though several orders trend towards a ventromyarian condition (with muscle scar area concentrated at the bottom of the body chamber). 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. When originally defined in 2013, Multiceratoidea included four nautiloid orders: Ellesmerocerida, Oncocerida, Discosorida, and Ascocerida. The order Tarphycerida was considered a potential member of ...
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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 ...
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Clarkoceras
''Clarkoceras'' is a genus of breviconic ellesmerocerid cephalopods, one of only two genera known to have crossed from the Late Cambrian, Trempealeauan, into the Early Ordovician, Gasconadian. (Flower 1964, Teichert 1988); the other being '' Ectenolites''. Description Clarkoceras has a rapidly expanding, laterally compressed, relatively short, endogastrically cyrtoconic shell; The upper or dorsal side is more strongly convex longitudinally than the lower or ventral side is concave. Sutures are essentially straight and close spaced indicating very short camerae (chambers). The siphuncle is relatively large, 0.3 the dorsoventral dimension and is ventral, although not necessarily marginal. (Flower 1964, Furnish and Glenister 1964). Septal necks are straight, reaching about half way back to the previous septum and the connecting rings are thick and layered Taxonomy ''Clarkoceras'' was first thought by Clarke to be a pilocerid, who in 1897 give it the name ''Piloceras newton-winche ...
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Ectenolites
''Ectenolites'' is a genus of small, slender, cylindrical Ellesmeroceratids that resemble '' Ellesmeroceras'' but are smaller and proportionally narrower. Septa, as typical for ellesmerocerids, are close spaced with shallow lobes on either flank. The body chamber is proportionally long, the shell itself slightly compressed. The dorsal side at the beginning of the shell, opposite the aperture and body chamber, is strongly convex so to produce a sense of endogastric curvature with the apex and siphuncle 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 ... aligned.Flower R.H.1964, The Nautiloid Order Ellesmeroceratida (Cephalopoda). Memoir 12, New Mexico Bureau of Mines and Mineral Resources, Socorro, NM The siphuncle in ''Ectenolites'' lies along the ventral side; is tubular, composed ...
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Evolutionary Grade
A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of species united by morphological or physiological traits, that has given rise to another group that has major differences from the ancestral condition, and is thus not considered part of the ancestral group, while still having enough similarities that we can group them under the same clade. The ancestral group will not be phylogenetically complete (i.e. will not form a clade), so will represent a paraphyletic taxon. In order to fully understand evolutionary grades, one must first get a better understanding of Phylogenetics, defined as "''In biology, is the study of the evolutionary history and relationships among individuals or groups of organisms (e.g. species, or populations). These relationships are discovered through phylogenetic i ...
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Yanhecerida
Yanhecerida is a small order of Late Cambrian (mid- Stage 10) 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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ... has maintained that Yanhecerida is a valid clade of early cephalopods closely related to a paraphyletic Ellesmerocerida (and by extension all post-Cambian cephalopods). References {{reflist Prehistoric nautiloids Preh ...
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Paraphyly
In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In contrast, a monophyletic group (a clade) includes a common ancestor and ''all'' of its descendants. The terms are commonly used in phylogenetics (a subfield of biology) and in the tree model of historical linguistics. Paraphyletic groups are identified by a combination of synapomorphies and symplesiomorphies. If many subgroups are missing from the named group, it is said to be polyparaphyletic. The term was coined by Willi Hennig to apply to well-known taxa like Reptilia ( reptiles) which, as commonly named and traditionally defined, is paraphyletic with respect to mammals and birds. Reptilia contains the last common ancestor of reptiles and all descendants of that ancestor, including all extant reptiles as well as the extinct synapsids, ...
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Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The Ordovician, named after the Celtic Britons, Welsh tribe of the Ordovices, was defined by Charles Lapworth in 1879 to resolve a dispute between followers of Adam Sedgwick and Roderick Murchison, who were placing the same Rock (geology), rock beds in North Wales in the Cambrian and Silurian systems, respectively. Lapworth recognized that the fossil fauna in the disputed Stratum, strata were different from those of either the Cambrian or the Silurian systems, and placed them in a system of their own. The Ordovician received international approval in 1960 (forty years after Lapworth's death), when it was adopted as an official period of the Paleozoic Era by the International Union of Geological Sciences, Intern ...
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