Edestus | |
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Jaw whorl of Edestus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Subclass: | Holocephali |
Order: | † Eugeneodontida |
Family: | † Edestidae |
Genus: | † Edestus Leidy, 1856 |
Species | |
Synonyms | |
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Edestus is an extinct genus of eugeneodontid holocephalian fish known from the Late Carboniferous (Pennsylvanian) of the United Kingdom, Russia, and the United States. Most remains consist of isolated curved blades or "whorls" that are studded with teeth, that in life were situated within the midline of the upper and lower jaws. Edestus is a Greek name derived from the word edeste (to devour), in reference to the aberrant quality and size of the species' teeth. [1] The largest species, E. heinrichi, has been conservatively estimated to reach greater than 6.7 m (22 ft) in length, around the size of the largest known great white shark, possibly making it the largest marine predator to have ever existed up to that point. [2]
Like its other relatives, such as Helicoprion , and unlike modern sharks, the species of Edestus grew teeth in curved blades or "whorls". In Edestus' case, only a single row of teeth occurred in the midline of each jaw, leading Edestus to sometimes be described as the "scissor tooth shark". The degree of curvature in the teeth brackets, along with size, are distinct in each species.
Edestus was first described by Joseph Leidy in 1856. The type species is Edestus vorax, the holotype specimen (ANSP 9899) is very fragmentary and of uncertain stratigraphic and geographic provenance, reported to be from Muskogee County, Oklahoma, though this has been questioned. [3] In 1855 Edward Hitchcock presented a specimen of Edestus to the American Association for the Advancement of Science meeting at Providence, Rhode Island the specimen (a tooth whorl) had been originally found in Parke County, Indiana. The specimen was loaned to famed British anatomist Richard Owen in 1861, who referred the specimen to Edestus and suggested that it was a fin spine. [4] Edestus minor was described in 1866 by John Strong Newberry based on AMNH FF477, a single crown that lacks most of the root from Posey County, Indiana. [4] Edestus heinrichi was described in 1870 by Newberry and Amos Henry Worthen from a now-lost specimen found in Illinois. [5] Edestus triserratus was described in 1904 by Edwin Tulley Newton, from GSM 31410, which was found in the Coal Measures in Staffordshire, England which is a partial tooth lacking the apex of the crown. [6] Oliver Perry Hay in 1912 was the first to describe an associated pair of upper and lower tooth whorls (USNM V7255), found in a coal mine near Lehigh, Iowa, which he assigned to the new species Edestus mirus, now thought to be a synonym of Edestus minor. [5] Specimens of Edestus are also known from the Moscow Basin on the Russian Platform. Edestus karpinskii was described by A.B. Missuna in 1908 from remains found near Kolomna around 100 km southeast of Moscow. [7] Edestus minusculus was described by Hay in 1912 from a specimen originally described by Alexander Karpinsky as Edestus cf. minor from a specimen found at the Myachkova quarry, near Moscow. [3]
The whorls of Edestus are composed of numerous serrated teeth that have long v-shaped roots that are stacked on top of each other akin to roof tiling. Up to a dozen teeth are present in each whorl at any one time. Teeth grew in the back end (posterior) of the whorl, and gradually migrated forward through the whorl, before being ejected at the front end (anterior). Up to 40 teeth could have grown in each whorl during the lifetime of the animal. The upper and lower whorls are distinct in their morphology, with the lower whorl having a greater degree of curvature. [5]
As with most cartilaginous fish, preserved skeletal material is rare, due to the cartilage of the skeleton having a poor potential for fossilisation. Several specimens with the preserved cranial remains of Edestus have been reported, the most important being FMNH PF2204, a crushed juvenile specimen, likely representative of E. heinrichi, which preserves both upper and lower blades in association with a well preserved chondrocranium and jaws. [8] The Meckel's cartilage of the lower jaw is approximately 1.5 times the length of the lower whorl, the end of the lower whorl extends forward beyond the edge of the Meckel's cartiilage. The Meckel's cartilage was articulated with a slender quadrate via a socket in the Meckel's cartilage which articlulated with a process of the quadrate. The quadrate at its other end articulated with the otic process of the chondrocranium, the structure that houses the brain and sensory organs. The upper tooth blade was rigidly held between two plates of palatine cartilage, which at their front ends are crescent shaped, matching the curvature of the tooth whorl, with the upper whorl extending forward beyond the cartilage. The chondrocranium was capped by a shield-shaped dorsal plate. [8]
No postcranial remains are known of Edestus. However, poscranial remains are known for other eugeneodont genera. Eugeneodonts with preserved postcrania include the Pennsylvanian to Triassic-age caseodontoids Caseodus, Fadenia, and Romerodus. [9] [10] [11] These taxa have a fusiform (streamlined, torpedo-shaped) body plan, with triangular pectoral fins. There is a single large and triangular dorsal fin without a fin spine, and a tall, forked caudal fin which externally appears to be homocercal (with two equally-sized lobes). This general body plan is shared by active, open-water predatory fish such as tuna, swordfish, and lamnid sharks. Eugeneodonts also lack pelvic and anal fins, and judging by Romerodus, they would have had broad keels along the side of the body up to the caudal fin. Fadenia had five well-exposed gill slits, possibly with a vestigial sixth gill. There is no evidence of the specialized gill basket and fleshy operculum present in living chimaeroids. [9] [10]
The cranium of FMNH PF2204 is around 25 centimetres (9.8 in) in length, and the associated upper and lower whorls have lengths of 10.4 and 8 centimetres (4.1 and 3.1 in), respectively. The largest known individuals of E. heinrichi are an upper whorl 32 centimetres (13 in) in length and a lower whorl 43 centimetres (17 in) in length. The estimated minimum length of the skulls of these individuals based on allometry are around 77 and 134 centimetres (30 and 53 in), respectively. Based on a 5:1 body length to head ratio, this suggests that individuals of E. heinrichi could reach lengths of about or more than 6.7 m (22 ft). [5] This estimate has been corroborated by Engelman (2023) based on allometric scaling, who also suggested that Edestus was likely the largest marine predator to have ever existed up until that point, given that earlier large predators like Dunkleosteus were probably smaller than previously thought. [2]
Due to the unusual nature of the tooth whorls and the historic lack of cranial material, many hypotheses for how the whorls functioned have been proposed. Early hypotheses suggested that they were indeed teeth or were defensive spines located on the fins. Wayne M. Itano proposed that the whorls were arranged vertically to slash prey. [12] In the description of the cranial material of Edestus, Tapanila and colleagues (2018) found that the tooth whorls functioned as effective grasping and slicing tools for soft bodied prey. The jaws of Edestus were operated by a two gear system, involving double jointing of the quadrate bone allowing for forward and backward movement of the lower jaw, similar to the streptostyly seen in living squamates, with an estimated force output of 1907 newtons. [8] The authors proposed that during prey approach, adductor muscles pulled the Meckel's cartilage of the lower jaw upwards and forwards to close the jaw, causing each tooth to slice roughly three times its length, and further push the prey into the teeth of the upper whorl. During the subsequent opening of the jaw mouth the Meckel's cartilage moved backwards and downwards by the adductor muscles, causing further slicing. [8]
13 species of Edestus have been named, but in a 2019 morphometric analysis only 4 were considered valid, which span a six million year interval in the Late Carboniferous (313-307 million years ago). Species of Edestus are divided into two groups, those which have asymmetrical crowns that slant forwards, and those with symmetrical ones. The earliest known species are from the late Bashkirian of the United Kingdom, with the first appearances in Russia and the United States during the Moscovian, corresponding to a rise in sea level. [5] These localities were situated in paleoequatorial tropical latitudes. Most remains of Edestus are found in marine shales that overlie coal swamp deposits as a result of marine transgression events. [5]
Edestusnewtoni, described by Arthur Smith Woodward in 1916 from the "Millstone Grit" of Yorkshire, United Kingdom, has a much greater curvature of the whorl than other species of Edestus, and has sometimes been placed in its own genus Lestrodus . A placement in a separate genus is supported by its lack of a convex bulge opposite the tooth crowns, which is present on all other species of Edestus. [5]
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.
Helicoprion is an extinct genus of shark-like eugeneodont fish. Almost all fossil specimens are of spirally arranged clusters of the individuals' teeth, called "tooth whorls", which in life were embedded in the lower jaw. As with most extinct cartilaginous fish, the skeleton is mostly unknown. Fossils of Helicoprion are known from a 20 million year timespan during the Permian period from the Artinskian stage of the Cisuralian to the Roadian stage of the Guadalupian. The closest living relatives of Helicoprion are the chimaeras, though their relationship is very distant. The unusual tooth arrangement is thought to have been an adaption for feeding on soft bodied prey, and may have functioned as a deshelling mechanism for hard bodied cephalopods such as nautiloids and ammonoids. In 2013, systematic revision of Helicoprion via morphometric analysis of the tooth whorls found only H. davisii, H. bessonowi and H. ergassaminon to be valid, with some of the larger tooth whorls being outliers.
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 taillight shark is a little-known species of shark in the family Dalatiidae and the only member of its genus. It is known from only four specimens collected from deep oceanic waters in the southern Atlantic Ocean and Pacific Ocean. A small shark with a laterally compressed body and a bulbous snout, this species has unusual adaptations that indicate a specialized lifestyle: its pectoral fins are paddle-like and may be used for propulsion, unlike other sharks and it has a pouch-like gland on its abdomen that emits clouds of luminescent blue fluid. This shark is likely aplacental viviparous and a formidable predator for its size.
The quagga catshark is a species of catshark, belonging to the family Scyliorhinidae. A small, slim-bodied shark reaching 37 cm (15 in) in length, it has a distinctive color pattern of narrow, dark brown vertical bars, which resemble those of the quagga. Its head is short and flattened, with a pointed snout tip that is not upturned.
The slender sawtail catshark is a little-known species of catshark, part of the family Scyliorhinidae, endemic to northern Australia. It is found over the continental slope in 290–470 m (950–1,540 ft) on water. Growing to 34 cm (13 in) long, this shark has a slim gray body with four dark saddle markings below the dorsal fins and on the caudal fin, as well as a prominent crest of enlarged dermal denticles along the dorsal edge of the caudal fin. The slender sawtail catshark is not valued by fisheries but is taken as bycatch. The International Union for Conservation of Nature (IUCN) presently lacks enough information to assess its conservation status.
The dwarf sawtail catshark is a little-known species of catshark, belonging to the family Scyliorhinidae, found exclusively in the deep waters off Luzon in the Philippines. Unlike other members of its genus, this slender, diminutive shark has a short, rounded snout and very short furrows at the corners of its jaws. It has indistinct darker saddles beneath each dorsal fin and two dark bands on the caudal fin, as well as a prominent crest of enlarged dermal denticles along the upper caudal fin margin.
The dwarf lanternshark is a species of dogfish shark in the family Etmopteridae and is the smallest shark in the world, reaching a maximum known length of 20 cm (8 in). It is known to be present only on the upper continental slopes off Colombia and Venezuela, at a depth of 283–439 m (928–1,440 ft). This species can be identified by its small size at maturity, long flattened head, and pattern of black ventral markings and a mid-dorsal line. Like other members of its genus, it is capable of producing light from a distinctive array of photophores. Reproduction is aplacental viviparous, with females gestating two or three young at a time. The dwarf lanternshark is not significant to commercial fisheries, but could be threatened by mortality from bycatch; the degree of impact from human activities on its population is unknown.
Onychodus is a genus of prehistoric lobe-finned fish which lived during the Devonian Period. It is one of the best known of the group of onychodontiform fishes. Scattered fossil teeth of Onychodus were first described from Ohio in 1857 by John Strong Newberry. Other species were found in Australia, England, Norway and Germany showing that it had a widespread range.
Sarcoprion is an extinct genus of eugeneodont holocephalan from the Permian of Greenland. Similar to other helicoprionids such as Agassizodus and Helicoprion, it possessed tooth whorls on the symphysis of the jaw as well as flattened, pavement-type teeth. It is distinguished from other members of its family by the presence of sharp, symphyseal teeth on both the upper and lower jaws. The tooth whorl on the lower jaw bore sharp, compact tooth crowns, while a row of backward facing, triangular teeth was present on the roof of the mouth. The preserved material does not show evidence of a distinct upper jaw, implying it may have been fused to the cranium, reduced, or lost entirely. The type and only species in the genus is S. edax.
Parahelicoprion is an extinct genus of shark-like cartilaginous fish known from the early Permian of the Ural Mountains of Russia and the Copacabana Formation of Bolivia. Members of the genus possessed a row of large, serrated tooth crowns on the lower jaw, called a tooth whorl, and are considered members of the order Eugeneodontida. The genus name refers to relation with Helicoprion, another eugeneodont from the Ural Mountains which also bore a curved tooth whorl along the midline of the lower jaw. Two species of Parahelicoprion are assigned; the Russian P. clerci and the Bolivian P. mariosuarezi.
The Eugeneodontida, sometimes also called Eugeneodontiformes, is an extinct and poorly known order of cartilaginous fishes. They possessed "tooth-whorls" on the symphysis of either the lower or both jaws and pectoral fins supported by long radials. They probably lacked pelvic fins and anal fins. The palatoquadrate was either fused to the skull or reduced. Now determined to be within the Holocephali, their closest living relatives are chimaeras. The eugeneodonts are named after paleontologist Eugene S. Richardson, Jr. The group first appeared in the fossil record during the late Mississippian (Serpukhovian). The youngest eugeneodonts are known from the Early Triassic. The geologically youngest fossils of the group are known from the Sulphur Mountain Formation, Vardebukta Formation and Wordie Creek Formation (Greenland).
The Edestidae are a poorly known, extinct family of shark-like eugeneodontid holocephalid cartilaginous fish.
The phallic catshark is a little-known species of catshark, belonging to the family Scyliorhinidae. It is found on or near the ocean floor, in the deep waters off New Caledonia and Vanuatu. A slender species attaining a length of 46 cm (18 in), it is characterized by a long caudal fin bearing a crest of enlarged dermal denticles along the dorsal margin, and very long claspers in adult males. This shark is gray-colored, with four dark saddles along the back and tail.
Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.
Toxoprion is an extinct genus of eugeneodont holocephalans whose fossils are found in marine strata from the Early Carboniferous until the Late Permian near Eureka, Nevada.
Ornithoprion is a monotypic genus of extinct eugeneodont chondrichthyan in the family Caseodontidae. The type and 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 anatomy of the eugeneodonts, a group which were previously known primarily from isolated tooth whorls. The genus derives its name from its pointed, bill-like armored skull and large eyes, which vaguely resemble the features of a bird, while the species name honors Oscar Hertwig. It is known from specimens preserving the skull and anterior portion of the body.
Agassizodus is an extinct genus of eugeneodont holocephalian from the Carboniferous. It belongs to the family Helicoprionidae, which is sometimes called Agassizodontidae. Like other members of its family, it possessed a symphyseal tooth whorl, which was likely present at the tip of the lower jaw and associated with lateral crushing toothplates. The type species, A. variabilis, was originally named Lophodus variabilis until the name "Lophodus" was determined to be preoccupied.
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 type and 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, which 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. Unlike its larger relatives, Romerodus grew to approximately 50 cm in total length.
Paredestus is a monotypic genus of extinct eugeneodont holocephalan from the Early Triassic of Canada. The type and only species, P. bricircum represents the last known member of the superfamily Edestoidea and among the last known eugeneodonts. It was named in 2008 based on tooth and jaw material, with the holotype representing the only known specimen.