Orthacanthus | |
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Fossil interpretation of juvenile specimen of O. platypternus (top) and skeletal reconstruction of juvenile specimen of O. bohemicus (bottom) | |
Front view of skull, American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Subclass: | Elasmobranchii |
Order: | † Xenacanthida |
Family: | † Orthacanthidae Heyler and Poplin, 1989 |
Genus: | † Orthacanthus Agassiz, 1843 |
Species | |
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Orthacanthus is an extinct genus of fresh-water xenacanthiform cartilaginous fish, named by Louis Agassiz in 1843, [3] [4] ranging from the Upper Carboniferous [3] into the Lower Permian. [1] Orthacanthus had a nektobenthic life habitat, with a carnivorous diet. [5] Multiple authors have also discovered evidence of cannibalism in the diet of Orthacanthus and of "filial cannibalism" where adult Orthacanthus preyed upon juvenile Orthacanthus. [6] Synonyms of the genus Orthacanthus are Dittodus Owen, 1867, Didymodus Cope, 1883, Diplodus Agassiz, 1843, [5] Chilodus Giebel, 1848 (preoccupied by Chilodus Müller & Troschel, 1844). [3]
During the Late Carboniferous-Early Permian, Orthacanthus was an apex predator of freshwater swamps and bayous in Europe and North America. [6] Mature Orthacanthus reached nearly 3 meters (10 feet) in length. [6] Orthacanthus teeth have a minimum of three cusps, two principal cusps, and an intermediate cusp, where the principal cusps are variously serrated, with complex base morphology. [7] [4] Additionally, Orthacanthus can be diagnosed by major transverse axes of proximal ends at a 45-degree angle to and often almost parallel to the labial margin of the base between the cusps. [4] Deformed teeth are characteristic of the xenacanthiform sharks and of Orthacanthus. [8]
The two genera Orthacanthus and Pleuracanthus were erected by Louis Agassiz based on isolated "ichthyodorulites" from the British Carboniferous System, and at the time were mistakenly thought of as the first indicators of skates. [9] They were initially found in the United Kingdom in Dudley, Leeds, North Wales, Carluke, and Edinburgh. [9] Three additional species from the Carboniferous formation of Ohio were described by John Strong Newberry, but two of them were junior homonyms of another species, Orthacanthus gracilis (Giebel, 1848). Accordingly, these two species received replacement names, O. adamas Babcock 2024 and O. lintonensis Babcock, 2024. [3] Teeth associated with Diplodus, a genus of sharks, was found in the Carboniferous slates of England in Stafford, Carluke, and Burdiehouse, and in Nova Scotia. [9] A well preserved impression from Ruppelsdorf, Bohemia, was described by Goldfuss, and in a separate paper, the same specimen was described under the name Xenacanthus dechenii. One year later, in 1849, Dr. Jordan mistakenly identified this specimen as the remains of a fossil shark, Triodus sessilis. This mistake was corrected and the specimen was identified as Xenacanthus by Mr. Schnur. [9]
The larger teeth of Orthacanthus compressus and Orthacanthus texensis are differentiated by a more pronounced basal tubercle in O. compressus. [7] The basal tubercle of a typical tooth file is on the apical button of the underlying tooth. [8] The larger adult teeth of O. compressus have a wider rather than longer base, similar to O. texensis, and tend to have serrations on both carinae of each cusp, while the medial carinae of smaller adult teeth are not serrated. [7] The juvenile teeth of O. compressus are longer than wide, have a thinner base, and lack serrations, similar to O. platypternus teeth. [7]
Orthacanthus platypternus from the Craddock Bonebed shark layer in Texas, USA, shows evidence of resorption, and the equivalent of an "enamel pearl." [8] Some of the teeth specimens found at this location show evidence of resorption, which has not been previously observed in other faunal members at the same location. [8] Where the superjacent basal tubercle is expected to be resorbed if the teeth were to undergo resorption, the apical button is resorbed instead. [8]
The difference in characteristics between the large and small O. compressus adult teeth might indicate sexual dimorphism. [7]
The spines of O. platypternus showing 3 to 4 dentine layers are interpreted to be subadults or young adults, and are separated into two size classes where females have the largest spines in comparison to males, indicating sexual dimorphism. [10]
The dorsal spines of Orthacanthus platypternus from the Craddock Bone Bed in Texas, USA, preserve a highly vascularized wall mainly composed of centrifugally growing dentine (the outer layer of the wall of the spine) in a succession of inwardly growing dentine layers that line the pulp cavity. [10] These dentine layers are likely deposited periodically in accordance with seasonal variations in water temperature and food availability. [10] More specifically, the periodic nature of the dentine layer deposits could be due to variation in calcium phosphate deposition following the changes in water temperature. [11] Spines of individuals with 1-2 dentine layers are likely juveniles and result in the smallest sizes, whereas individuals showing at least 3-4 dentine layers result in two separate size classes. [10] The cross section is oval near the opening of the pulp cavity and circular/subtriangular in the distal part of the non-denticulated region and circular in the denticulated region. [10] The pulp cavity of the spine is filled with calcite, quartz, and opaque minerals. [10]
The spine is superficially inserted in the skin, where it grows and moves from a deep position in the dermis where trabecular dentine forms, to a superficial location where centrifugally growing lamellar dentine forms. [11] The number of denticles per annual cycle vary with growth rate, and are independent dermal elements formed by the dermal papilla and secondarily attached by dentine to the spine proper. [11] The density of denticulation also varies with the growth rate of the occipital spine. [11] The ratio of length of denticulated region to total length of the spine changes throughout ontogeny. [11]
The teeth of Orthacanthus texensis and Orthacanthus platypternus from bonebeds from the Lower Permian of Texas, and the teeth of Orthacanthus compressus from the Upper Pennsylvanian of Nebraska and Dunkard Basin of the central Appalachians were used to determine the origin of O. texensis and O. platypternus. [7] It has been proposed that both O. texensis and O. platypternus could be derived from O. compressus, where juvenile features of O. compressus are retained in the adult teeth of O. platypternus via paedomorphosis, and the juvenile features of O. compressus teeth are observed in the adult teeth of O. texensis. [7]
The taxon Mimia sits outside of a clade that contains two monophyletic sister groups. The first monophyletic sister group defines a clade that includes the stem-group chondrichthyans which is visualized as a sister group of Doliodus and a large clade that comprises the cladodont sharks and Orthacanthus. The “cladodont sharks” plus Orthacanthus comprise two monophyletic sister groups: Orthacanthus and Cladodoides and Tamiobatis on one side, and the Symmoriiformes on the other side. The second monophyletic sister group characterizes crown chondrichthyans which contains two monophyletic sister groups ( Euchondrocephali on one side and Euselachii on the other side). [12]
The following cladogram follows a 2011 analysis by Alan Pradel and colleagues. [12]
A 2013 analysis of oxygen and strontium isotope composition of the teeth and spines of Late Carboniferous and Early Permian shark taxa was performed to infer the hydrochemistry of their ambient water, thus contributing to the controversy between an obligate freshwater or euryhaline diadromous lifestyle. [13] Facies interpretations in the Permian of North America suggested that salinity tolerances of xenacanthiforms were restricted to near marine environments whereas only Orthacanthus could tolerate brackish water environments. [13] A study covering the morphology and histology of dorsal spines of Orthacanthus platypternus also reported that "The comparative analyses of the ontogenetic stages of the recorded specimens of O. platypternus and their distribution along different facies and localities indicate that this species was euryhaline, diadromous with a catadromous life-cycle which was strongly regulated by the semi-arid, seasonally dry tropical climate affecting western Pangea during the Early Permian. [10] " The 2013 analysis provided evidence leaning towards an obligate freshwater lifestyle of the sharks from Variscan European basins, and nonmarine ratios suggested tooth formation was influenced by meteoric waves enriched by evaporation. [13] Euryhaline adaptation was not confirmed in the 2013 analysis. [13]
Orthacanthus and Triodus have a predator-prey relationship in which Orthacanthus preyed on Triodus. [14] Cranial remains of specimens of both Orthacanthus and Triodus from the Upper Carboniferous in Puertollano basin, Spain, give evidence of this predator-prey relationship. [14] Numerous and well preserved cephalic elements of Triodus were associated with the cranial remains of Orthacanthus, and is explained by the inclusion of occipital spines of Triodus in the buccal cavity of Orthacanthus. [14] Additional evidence is the co-occurrence of one Orthacanthus spine with many Triodus spines, which likely penetrated the soft tissue and cartilage of the mouth of Orthacanthus, similarly to modern sharks that feed on stingrays where the spines of stingrays have been found within and around the buccal cavities of Carcharhinus, Galeocerdo, Negaprion and Sphyrna.
Examination of Orthacanthus coprolites from Canada by Aodhan O' Gogain et al. revealed that in times of hardship, Orthacanthus was likely cannibalistic, as teeth from juvenile Orthacanthus were found within the coprolites of adults. [15] [16] Orthacanthus had a diet that consisted of actinopterygians, acanthodians, dipnoans, xenacanthids and tetrapods, based on analysis of coprolites and gut contents. [6] There have also been suspicions of filial cannibalism due to the presence of juvenile Orthacanthus teeth inside an Orthacanthus coprolite. The feces of Orthacanthus has a spiral shape due to a corkscrew-shaped rectum. [6]
The paleobiogeographical distribution of O. platypternus suggests ontogenetic habitat partitioning. [10] Ontogenetic niche theory predicts that individuals may change their habitat or diet to maintain optimal growth rates or to improve trade-offs between mortality risk and growth. [17] While smaller individuals likely lived in shallower waters such as in small ponds and stream channels of the coastal plain, larger individuals likely lived in deeper water such as the fluvio-lacustrine (rivers and lakes) and marginal marine areas. [10]
The oldest known specimen of Orthacanthus, Diplodus problematicus, was found in New Brunswick, Canada, in the Lower Devonian (Emsian, c. 407 to 393 million years ago). [5] Other specimens have been found in locations including the US, the United Kingdom, Poland, and France. [5]
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 clefts 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.
Acanthodii or acanthodians is an extinct class of gnathostomes. They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, and chimaeras. Acanthodians possess a mosaic of features shared with both osteichthyans and chondrichthyans. In general body shape, they were similar to modern sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteians.
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.
Xenacanthus is an extinct genus of xenacanth cartilaginous fish. It lived in freshwater environments, and fossils of various species have been found worldwide.
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.
Xenacanthida is an order or superorder of extinct shark-like chondrichthyans known from the Carboniferous to Triassic. They were native to freshwater, marginal marine and shallow marine habitats. Some xenacanths may have grown to lengths of 5 m (16 ft). Most xenacanths died out at the end of the Permian in the End-Permian Mass Extinction, with only a few forms surviving into the Triassic.
Stethacanthidae is an extinct family of prehistoric holocephalians. 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.
Stereosternum tumidum is an extinct genus of mesosaur marine reptile from the Early Permian of Brazil and also the Great Karoo Basin of South Africa. The taxon mesosaur is a monophyletic group containing Brazilosaurus sanpauloensis and Mesosaurus tenuidens.
Triodus is an extinct genus of xenacanthiform cartilaginous fish that lived from the Carboniferous to the Permian. In 2017, a new species Triodus richterae was described from the Rio do Rasto Formation of Brazil.
Sphenacanthus is an extinct genus of a chondrichtyan xenacanthiform that belongs to the Sphenacanthidae family and lived from the Late Devonian, through Carboniferous until the Late Permian period in Scotland, Spain, Russia and Brazil. It lived 359 million years ago, and probably it was one of the first member of the elasmobranchians, the lineage that leads to the modern sharks. Sphenacanthus probably hunts small fishes and, unlike their modern-day relatives, its inhabited fresh water lagoons. Sphenacanthus had seven fins, two in the upper part and five in the underside, and it had a heterodont dentition and mandibles relatively long and deeper. Sphenacanthus serrulatus is still only known from incomplete neurocranial remains and associated dermal material. These suggest that it was a relatively large shark, probably well over one meter in length when fully grown. Its body form was probably similar to that of other phalacanthous sharks.
Hybodontiformes, commonly called hybodonts, are an extinct group of shark-like cartilaginous fish (chondrichthyans) which existed from the late Devonian to the Late Cretaceous. Hybodonts share a close common ancestry with modern sharks and rays (Neoselachii) as part of the clade Euselachii. They are distinguished from other chondrichthyans by their distinctive fin spines and cephalic spines present on the heads of males. An ecologically diverse group, they were abundant in marine and freshwater environments during the late Paleozoic and early Mesozoic, but were rare in open marine environments by the end of the Jurassic, having been largely replaced by modern sharks, though they were still common in freshwater and marginal marine habitats. They survived until the end of the Cretaceous, before going extinct.
Plicatodus is a prehistoric cartilaginous fish in the family Xenacanthidae that lived in Europe during the late Carboniferous and Early Permian Periods. It was described by Oliver Hampe in 1995, and the type species is Plicatodus jordani. The type locality for this genus is the Saar-Nahe basin.
The Archer City Formation is a geological formation in north-central Texas, preserving fossils from the Asselian and early Sakmarian stages of the Permian period. It is the earliest component of the Texas red beds, introducing a tropical ecosystem which will persist in the area through the rest of the Early Permian. The Archer City Formation is preceded by the cool Carboniferous swamp sediments of the Markley Formation, and succeeded by the equally fossiliferous red beds of the Nocona Formation. The Archer City Formation was not named as a unique geological unit until the late 1980s. Older studies generally labelled its outcrops as the Moran or Putnam formations, which are age-equivalent marine units to the southwest.
The Arroyo Formation, sometimes termed the Lower Clear Fork Formation, is a geologic formation in Texas. It preserves fossils dating back to the Kungurian stage of the Permian period. It is the lower-most portion of the Clear Fork Group, part of a series of fossiliferous Permian strata in the south-central United States known as the red beds.
Antarctilamna is an extinct genus of Devonian cartilaginous fish originally exemplified by Antarctilamna prisca from South Eastern Australia and Antarctica. The latest occurring described species is Antarctilamna ultima from the Waterloo Farm lagerstätte in South Africa. Antarctilamna has robust ctenacanthid-like spines which lack a deep insertion area, and are borne in front of the first dorsal fin; in addition to distinctive diplodont teeth with small intermediate cusps. Antarctilamna-like spines, known from the Bunga Beds locality in Australia have been ascribed to A. prisca.
Barbclabornia is an extinct genus of xenacanth from the Early Permian and possibly upper Pennsylvanian of North America. The genus contains a single described species: B. luedersensis. It has been found in several places within Asselian and Atinskian formations, including the Clear Fork, Albany, Wichita, and Dunkard Groups. There are possible examples from the Gzhelian-aged Admire, Monongahela, and Conemaugh groups.
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.
Cretacladoides is a genus of chondrichthyan, possibly a falcatid, found in France and Austria. Known solely from teeth, mainly found in the Klausrieglerbach locality of Austria, it consists of two species, C. ogiveformis and C. noricum. Assuming a falcatid identity, it is the most recent member of the family, which otherwise became extinct at the end of the Carboniferous.
Dracopristis is an extinct genus of ctenacanth that lived around 307 million years ago during the Pennsylvanian subperiod of the Carboniferous period. A single species is known, Dracopristis hoffmanorum, which is named in honor of Ralph and Jeanette Hoffman.
Lebachacanthus is a genus of extinct xenacanth cartilaginous fish known from the late Carboniferous-Early Permian of Europe. Well-preserved specimens, originally identified as Orthacanthus, are known from Meisenheim Formation in Germany. During the late Paleozoic, xenacanths were the apex predators of freshwater ecosystems, preying on small amphibians.
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