This article needs additional citations for verification .(September 2014) |
Holocephali Temporal range: | |
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Chimaera monstrosa , a rat fish | |
† Helicoprion davisii , a eugeneodontid | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Subclass: | Holocephali Bonaparte, 1832 |
Orders | |
Holocephali sensu lato (Euchondrocephali)
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Holocephali ("complete heads"), sometimes given the name Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. [1] The earliest fossils are of teeth and come from the Devonian period. The only living holocephalans are the chimaeras (Chimaeriformes), though during the late Paleozoic (Carboniferous and Permian) Holocephali was much more diverse, including an array of forms including those considerably different from modern Chimaeriformes, including shark-like predatory forms and slow, durophagous fish.
Chimaeras, also known as rat fish, or ghost sharks, include three living families and a little over 50 species of surviving holocephalans. These fishes move by using sweeping movements of their large pectoral fins. They are deep sea fish with slender tails, living close to the seabed to feed on benthic invertebrates. They lack a stomach, their food moving directly into the intestine.
Members of this taxon preserve today some features of elasmobranch life in Paleozoic times, though in other respects they are aberrant. They live close to the bottom and feed on molluscs and other invertebrates. The tail is long and thin and they move by sweeping movements of the large pectoral fins. The erectile spine in front of the dorsal fin is sometimes venomous. There is no stomach (that is, the gut is simplified and the 'stomach' is merged with the intestine), and the mouth is a small aperture surrounded by lips, giving the head a parrot-like appearance. The only surviving members of the group are the rabbit fish ( Chimaera ), and the elephant fishes ( Callorhinchus ). [2] [3]
The fossil record of the Holocephali starts during the Devonian period. [4] The record is extensive, but most fossils are of teeth, and the body forms of numerous species are not known, or at best poorly understood. Some experts[ who? ] further group the orders Petalodontiformes, Iniopterygiformes, and Eugeneodontida into the taxon "Paraselachimorpha", and treat it as a sister group to Chimaeriformes. However, as almost all members of Paraselachimorpha are poorly understood, most experts suspect this taxon to be either paraphyletic or a wastebasket taxon.
Lund & Grogan (1997) coined the subclass Euchondrocephali to refer to the total group of holocephalians, i.e. all fish more closely related to living holocephalians than to living elasmobranchs such as sharks and rays. Under this classification scheme, "Holocephali" would have a much more restricted definition. [5] Other authors have used Holocephali in a broad sense covering all fish more closely related to chimaeras than to sharks and rays. [6]
Based on genetic research, it is estimated the Holocephali split from the Elasmobranchii (the branch of chondrichthyans containing true sharks and rays) about 421 million years ago. [7]
While historically considered closely related to elasmobranchs, recent studies have found members of the shark-like "symmoriiformes", which may be paraphyletic, to be early diverging relatives of Holocephali, (with some of these studies choosing to exclude them from Holocephali proper). [8]
Taxonomy according to Joseph Nelson, 2006 [9] |
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† Extinct* position uncertain |
Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opercula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.
Chimaeras are cartilaginous fish in the order Chimaeriformes, known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively.
Elasmobranchii is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks, rays, skates, and sawfish. Members of this subclass are characterised by having five to seven pairs of gill slits opening individually to the exterior, rigid dorsal fins and small placoid scales on the skin. The teeth are in several series; the upper jaw is not fused to the cranium, and the lower jaw is articulated with the upper. The details of this jaw anatomy vary between species, and help distinguish the different elasmobranch clades. The pelvic fins in males are modified to create claspers for the transfer of sperm. There is no swim bladder; instead, these fish maintain buoyancy with large livers rich in oil.
Cladoselache is an extinct genus of shark-like chondrichthyan from the Late Devonian (Famennian) of North America. It was similar in body shape to modern lamnid sharks, but was not closely related to lamnids or to any other modern (selachian) shark. As an early chondrichthyan, it had yet to evolve traits of modern sharks such as accelerated tooth replacement, a loose jaw suspension, enameloid teeth, and possibly claspers.
Stethacanthus is an extinct genus of shark-like cartilaginous fish which lived from the Late Devonian to Late Carboniferous epoch, dying out around 298.9 million years ago. Fossils have been found in Australia, Asia, Europe and North America.
The Bear Gulch Limestone is a limestone-rich geological lens in central Montana, renowned for the quality of its marine fossils from the late Mississippian subperiod, about 324 million years ago. It is exposed over a number of outcrops northeast of the Big Snowy Mountains, and is often considered a component of the more widespread Heath Formation. The Bear Gulch Limestone reconstructs a diverse, though isolated, marine ecosystem which developed near the end of the Serpukhovian age in the Carboniferous period. It is a lagerstätte, a particular type of rock unit with exceptional fossil preservation of both articulated skeletons and soft tissues. Bear Gulch fossils include a variety of fish, invertebrates, and algae occupying a number of different habitats within a preserved shallow bay.
Symmoriiformes is an extinct order of cartilaginous fish. Originally named Symmoriida by Zangerl (1981), the name has since been corrected to Symmoriiformes to avoid confusion with a family. The symmoriiform fossils record begins during the late Devonian, and most had become extinct by the start of the Permian, with the genus Dwykaselachus from the Artinskian-Kungurian of South Africa being the latest known uncontroversial occurrence. Teeth described from the Valanginian of France and Austria indicate that members of the family Falcatidae might have survived until the Early Cretaceous; however, it has since been proposed that these teeth more likely belonged to neoselachian sharks.
Stethacanthidae is an extinct family of prehistoric sharks. It is estimated to have existed approximately between 380 and 300 million years ago. Members of this family are noted for their peculiar dorsal fin.
Callorhinchus, the plough-nosed chimaeras or elephantfish, are the only living genus in the family Callorhinchidae. A few extinct genera only known from fossil remains are recognized. Callorhinchus spp. are similar in form and habits to other chimaeras, but are distinguished by the presence of an elongated, flexible, fleshy snout, with a vague resemblance to a ploughshare. They are only found in the oceans of the Southern Hemisphere along the ocean bottom on muddy and sandy substrates. They filter feed, with small shellfish making up the bulk of their diet. The plough-nosed chimaera lays eggs on the ocean floor that hatch at around 8 months. They are currently not a target of conservation efforts; however, they may be susceptible to overfishing and trawling.
The silver chimaera, or ginzame, is a species of holocephalan in the family Chimaeridae. They are found in the deep sea along the coast of East Asia, from Japan to Indonesia. They are chondrichthyans, closely related to sharks and rays, which means that they have a fully cartilaginous skeleton with no true bones.
Orodus is an extinct genus of cartilaginous fish that lived from the late Pennsylvanian to the early Permian in what is now North America. O. greggi reached around 2–4 metres (6.6–13.1 ft) long, while O. micropterygius just reached 1 metre (3.3 ft). It was a member of the holocephali subclass, meaning that it was more closely related to chimaeras and their relatives, and not elasmobranchs like sharks and rays.
Iniopterygiformes is an extinct order of cartilaginous fish known only from the Carboniferous period of the United States. Iniopterygians are characterized by large, superficially wing-like pectoral fins positioned upwards behind the head, from which the name of the group is derived. Iniopterygians are also noted to possess proportionally large skulls and eyes, armor plates composed of dentin, and "tooth-whorls" of fused teeth. Their elongated pectoral fins bore large, denticle-covered spines, and they are thought to have used them to swim using a "flying" motion. The iniopterygians were comparatively small chondrichthyans, with the largest species reaching only 50 cm (20 in) in length.
Squaloraja is an extinct genus of ray-like marine chimaeriform fish from the Early Jurassic of Europe.
Ornithoprion is a genus of extinct cartilaginous fish in the family Caseodontidae. The only species, O. hertwigi, lived during the Moscovian stage of the Pennsylvanian, between 315.2 to 307 million years ago, and is known from black shale deposits in what is now the Midwestern United States. The discovery and description of Ornithoprion, performed primarily via radiography, helped clarify the skull anatomy of eugeneodonts; a group which includes O. hertwigi and which were previously known primarily from isolated teeth. The genus name, which is derived from the ancient Greek órnith- meaning 'bird' and príōn meaning 'saw', was inspired by the animal's vaguely bird-like skull and the saw-like appearance of the lower teeth. The species name honors Oscar Hertwig.
Dwykaselachus is an extinct genus of symmoriiform, a cartilaginous fish that lived in what is now South Africa during the Permian period around 280 million years ago. It was first discovered in the 1980s, in a nodule of sediments from the Karoo Supergroup. Dwykaselachus was named based on Dwyka Group, the group of sedimentary geological formation in the southeastern part of Africa. It represents the place where the type species Dwykaselachus oosthuizeni was found.
Ctenacanthiformes is an extinct order of cartilaginous fish. They possessed ornamented fin spines at the front of their dorsal fins and cladodont-type dentition, that is typically of a grasping morphology, though some taxa developed cutting and gouging tooth morphologies. Some ctenacanths are thought to have reached sizes comparable to the great white shark, with body lengths of up to 7 metres (23 ft) and weights of 1,500–2,500 kilograms (3,300–5,500 lb), while others reached lengths of only 30 centimetres (12 in). The earliest ctenacanths appeared during the Frasnian stage of the Late Devonian, with the group reaching their greatest diversity during the Early Carboniferous (Mississippian), and continued to exist into at least the Middle Permian (Guadalupian). Some authors have suggested members of the family Ctenacanthidae may have survived into the Cretaceous based on teeth found in deep water deposits of Valanginian age in France and Austria, however, other authors contend that the similarity of these teeth to Paleozoic ctenacanths is only superficial, and they likely belong to neoselachians instead.
Dracopristis is an extinct genus of ctenacanth that lived during the Carboniferous period in North America, around 307 million years ago. The species was discovered in the Kinney Brick Quarry in New Mexico, US. Like many fossils from the site, the fossils of Dracopristis are very well-preserved. A single species is known, Dracopristis hoffmanorum, which is named in honor of Ralph and Jeanette Hoffman, the owners of the quarry. Prior to being scientific named, D. hoffmanorum was informally referred to as the "Godzilla shark".
Romerodus is an extinct genus of cartilaginous fish in the family Caseodontidae. While it and the rest of its family were historically considered elasmobranchs related to sharks and rays, they are now regarded as holocephalans, a diverse subclass which is today only represented by chimaeras. Romerodus is known from the Carboniferous and possibly Permian periods of North America, and the only named species, R. orodontus, was discovered in organic shale deposits in the U.S. state of Nebraska. It is one of few members of its order, the Eugeneodontida, that is known from multiple complete, well preserved body fossils, and is thus an important taxon for understanding the anatomy and ecology of less well preserved eugeneodonts such as Helicoprion. The genus name honors paleontologist Alfred Romer.
Squatinactis is a genus of extinct elasmobranch chondrichthyan known from the Carboniferous aged Bear Gulch Limestone in Montana. This fish was discovered in 1974 by Richard Lund. The type specimen, named CMNH 46133, consists of a brain case, poorly preserved jaws and gills, a pectoral fin, and a partial vertebral axis. This creatures most startling feature were its broad pectoral fins which resembled those of stingrays and angel sharks (Squatina). The holotype specimen has about 15 teeth in its jaw. This creature is named after the angel shark. Remains found in the South Urals of Russia and the Eyam Limestone of Derbyshire, England, have been tentatively identified as those belonging to S. caudispinatus.
Fishes are a paraphyletic group and for this reason, the class Pisces seen in older reference works is no longer used in formal taxonomy. Traditional classification divides fish into three extant classes, and with extinct forms sometimes classified within those groups, sometimes as their own classes: