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Helicoprion

Helicoprion is an extinct genus of large shark-like cartilaginous fish that lived from the Early to the Middle Permian, about 290-270 million years ago. Helicoprion is a member of the Eugeneodontiformes, an extinct order of cartilaginous fish within the clade Holocephali, a group today represented only by chimaeras. It is also the type genus of the Helicoprionidae, a family of eugeneodonts characterised by distinctive tooth structures called tooth whorls. Helicoprion was first named in 1899 by Alexander Karpinsky on the basis of fossils discovered in Russia and Australia, the generic name meaning "spiral saw". Although numerous species were subsequently assigned to the genus, only H. bessonowi, H. davisii, and H. ergassaminon are recognized following a 2013 revision. The three species are distinguished by the shape and spacing of their tooth crowns.

Research history
figures of various eugeneodont fossils, the holotype of H. davisii (WAMAG 9080) being shown at the top (fig. 1) The first known specimen of Helicoprion consists of a partial tooth whorl preserving 15 teeth, 14 of which are visible. It was discovered in Western Australia in a tributary of the Gascoyne River by a gold prospector named Mr. Davis, his first name being unknown. Now housed in the Western Australian Museum under the catalogue WAMAG 9080, the fossil was not found in situ, thus its precise stratigraphic origin remains uncertain. tooth whorl of H. bessonowi (TsNIGR 1/1865) More complete tooth whorls were discovered in the late 19th century by Alexander G. Bessonov of the Russian Academy of Sciences, in a quarry near the town of Krasnoufimsk in Sverdlovsk Oblast. He sent them to geologist Alexander Karpinsky, who first described them in a monograph published in Russian in 1899, followed later that year by a German translation. As the fossils he examined were sufficiently distinct from those of Edestus, Karpinsky assigned them to a new genus, which he named Helicoprion, with H. bessonowi as the type species. The generic name Helicoprion derives from the Ancient Greek (hélikos, "spiral") and (príōn, "saw"), in reference to the characteristic morphology of the tooth whorls, while the specific epithet bessonowi honors their discoverer. In his 1899 observations, Karpinsky also noted that the specimen described by Woodward in 1886 showed sufficient similarities with the Krasnoufimsk material to justify its provisional reassignment to Helicoprion, renaming it H. davisii. A few years later, in 1909, his colleague Oliver Perry Hay transferred the species once again, this time to Toxoprion, another newly established, related genus. In later publications, Karpinsky reaffirmed the position he had argued in 1899, an interpretation followed by several authors in the subsequent decades. His interpretation was finally confirmed in 1940, when German–American paleontologist Curt Teichert described much more complete fossils of H. davisii, which like the holotype, were also discovered in Western Australia. Since the genus Helicoprion was established by Karpinsky in 1899, numerous additional species have been described from fossils found across the world, although most originate from North America. In a morphometric revision published in 2013, American paleontologists Leif Tapanila and Jesse Pruitt reassessed all of these historical species assignments. Their analysis showed that, among the roughly 10 named species, only three possess features distinctive enough to be considered valid: H. bessonowi, H. davisii, and H. ergassaminon, with the remaining taxa regarded as either synonymous or doubtful. H. ergassaminon was first described in 1966 by Danish ichthyologist Svend Erik Bendix-Almgreen in a monograph devoted to several Helicoprion fossils housed in the paleontological collections of various universities in Idaho, USA. As with the generic name, the specific epithet derives from the Ancient Greek (ergasamenon), meaning "the one who has done work", in reference to the distinctive wear marks observed on the holotype. This latter, nicknamed "Idaho 5" and discovered in a now-abandoned mine near Fort Hall, was reported by Bendix-Almgreen to be stored at the University of Idaho in Moscow, Idaho. However, later searches failed to locate the specimen at that institution, and the fossil was subsequently considered lost. For nearly a century, this species was distinguished on the basis of tooth angle and height. However, in their 2013 revision, Tapanila and Pruitt demonstrated that these traits vary naturally within a single Helicoprion species, so regarded H. ferrieri as a junior synonym of H. davisii. In their 2013 revision, Tapanila and Pruitt demonstrated that the characters attributed to H. sierrensis fall within the natural range of variation seen in H. davisii, while those of H. nevadensis correspond to the growth stage of H. bessonowi. The two species named by Wheeler were thus synonymized with H. davisii and H. bessonowi. Although the holotype of this species has since been reported lost, Tapanila and Pruitt noted that its anatomy closely resembles that of the largest known Helicoprion specimen. Nevertheless, neither specimen can be assigned to any of the three recognised species due to their incomplete preservation. Consequently, in the absence of more complete material attributable to the taxon, H. mexicanus is regarded as a nomen dubium. Contrary to ICZN requirements, though, the author provided no diagnosis establishing the distinctiveness of the taxon. As the specimen is also considered lost, Tapanila and Pruitt classified this species as a nomen nudum. Tapanila and Pruitt interpreted these observations as an artefact of the holotype’s partial preservation, since only the central portions of its teeth are preserved. The authors of the 2013 revision considered H. svalis to be strongly similar to H. bessonowi, but could not formally synonymise the two due to the incompleteness of the fossil material, so regarded it as a doubtful taxon. In their 2013 revision, though, Tapanila and Pruitt pointed out that the specimen is partially obscured by its surrounding matrix, which likely led to an underestimation of tooth height. Taking this into account, along with intraspecific variation, they regarded H. jingmenense as a junior synonym of H. davisii. , which was originally assigned to Helicoprion'' In 1916, Karpinsky erected the species H. clerci on the basis of fragmentary remains of a large tooth whorl discovered near Krasnoufimsk, the same locality that yielded the original fossils of H. bessonowi. In his description, however, he noted that the taxon might be sufficiently distinct to warrant its own genus. He formalised this proposal in 1924 by transferring the species to the genus Parahelicoprion, a name he had already used informally in several earlier publications. In 1986, a second species, P. mariosuarezi, was described by Bolivian geologist Dagmar Merino-Rodo and French palaeontologist Philippe Janvier on the basis of fossils recovered in Bolivia. In the same publication, however, the authors were the first to question the taxonomic status of the genus, noting the likely absence of clearly defined autapomorphies. This cast doubt on the monophyly of Parahelicoprion and complicated the assignment of additional species. In 2018 and 2023, Russian palaeontologists Sergey V. Naugolnykh and Dmitry V. Naumkin proposed that Parahelicoprion may, in fact, represent a junior synonym of Helicoprion, suggesting that the genus may have been named on the basis of fossils belonging to exceptionally large and old individuals of the latter. Other authors have continued to consider the genus to be distinct from Helicoprion. == Description ==
Description
Overall morphology of H. davisii. Like other chondrichthyan fish, Helicoprion had a skeleton made of cartilage. Around the jaws, this cartilage was mineralized, such as the Pennsylvanian to Triassic-age caseodontid eugeneodonts Caseodus, Fadenia, and Romerodus. No evidence has been found of the specialized gill basket and fleshy operculum present in living chimaeroids. Based on the proportional size of caseodontoid tooth whorls, Lebedev suggested in 2009 that Helicoprion individuals with tooth whorls in diameter could reach in total length, comparable to the size of modern basking sharks. Tooth size increases away from the center of the spiral, with the largest teeth possibly exceeding in height. The lower parts of the teeth form projections below the crown of the previous tooth. The lowest portion of the root below the enameloid tooth projections is referred to as the "shaft", and lies on jaw cartilage that covers the previous revolutions of the whorl. In a complete tooth whorl, the outermost part of the spiral terminates with an extended section of shaft that lacks the middle and upper portions of the tooth crown. Another well-preserved specimen of H. davisii, USNM 22577+494391 (nicknamed the "Sweetwood specimen"), has demonstrated that the inner surface of the palatoquadrate was covered with numerous small (~2 mm wide) teeth. The palatoquadrate teeth were low and rounded, forming a "pavement" that scraped against the tooth whorl. When seen from behind, the palatoquadrate forms a paired jaw joint with the Meckel's cartilage. No evidence is seen for articulation between the jaws and the hyomandibula, which helps to suspend the jaws of modern sharks. The Meckel's cartilage of Helicoprion has an additional projection right before the joint with the palatoquadrate. This extra process, unique to Helicoprion, likely served to limit jaw closure to prevent the whorl from puncturing the skull. Another unique characteristic of Helicoprion is that the preserved labial cartilage forms a synchondrosis (fused joint) with the upper surface of Meckel's cartilage. This joint is facilitated by a long facet on the upper edge of Meckel's cartilage. The labial cartilage provides lateral support for the tooth whorl, widening near the root of each revolution. By wedging into the palatoquadrate while the mouth is closed, the upper edge of the labial cartilage helps to spread out the forces used to limit the extent of the jaw closure. The rear portion of the labial cartilage has a cup-like form, protecting the developing root of the last and youngest revolution of the tooth-whorl. Scales Tooth-like chondrichthyan scales, specifically known as odontodes, have been found associated with H. bessonowi remains in Kazakhstan. They are broadly similar to scales of other eugeneodonts such as Sarcoprion and Ornithoprion. The scales have a cap-shaped base with a concave lower surface. The crowns are conical and covered with serrated, longitudinal ridges. The scales may be monodontode (with one crown per base) or polyodontode (with multiple crowns extending from a single base). Compared to other eugeneodonts, the scales of Helicoprion are more strongly pointed. == Classification ==
Classification
in Pocatello Skull data from IMNH 37899 reveal several characteristics, such as an autodiastylic (two-jointed) jaw suspension with a nonsuspensory hyomandibula, which confirm the placement of Helicoprion within the chondrichthyan subclass Holocephali (or the more broadly defined Euchondrocephali). The relationship between Helicoprion and living chimaeras is very distant, but had been previously suspected based on details of its tooth anatomy. As most eugeneodonts are based on fragmentary tooth remains, relationships within the group remain unclear. A cladogram illustrating the group's possible relations, drawn from Rainer Zangerl's 1981 volume of the Handbook of Paleoichthyology, is provided below. == Paleobiology ==
Paleobiology
Whorl function Due to the narrow nature of ''Helicoprion's jaws, suction feeding is unlikely to have been effective, and Helicoprion is thought to have been a bite feeder. Biomechanical modelling in a 2015 paper by American biologist Jason B. Ramsay and colleagues suggests that the teeth in the whorl had distinct functions depending on where they were in the spiral. The foremost teeth served to snag and pull prey further into the mouth, while the middle teeth speared, and the hind teeth punctured and brought prey further into the throat. The prey would be squeezed between the whorl and the palatoquadrates during feeding. The labial cartilage served to provide support for the whorl. The unusual, saw-like tooth whorl and the lack of wear on the teeth of Helicoprion'' implies a diet of soft-bodied prey, as hard-shelled prey would simply slip out of the mouth. Helicoprion may have started with a wide gape during prey capture, followed by smaller jaw opening and closing cycles to further transport prey into the mouth as is done by modern bite-feeding sharks. While modern sharks shake their heads from side to side to facilitate sawing and cutting their prey, the teeth of Helicoprion would likely further cut the prey during the jaw opening, due to the arc-like path of the front teeth. Helicoprion likely used a series of rapid, forceful jaw closures to initially capture and push prey deeper into the mouth, followed by repeated opening and closing of the jaw to saw through prey. Shortly after his initial monograph, Karpinsky published a new argument suggesting that the whorl represented a curved, plate-covered tail resembling that of seahorses. This proposal was immediately criticized by several researchers. The Belgian paleontologist Ernest Van Den Broeck emphasized the fragility of the structure and argued that it would be better protected if it were a paired oral apparatus located within the animal's cheeks. The British paleoichthyologist Arthur Smith Woodward expanded on this idea and proposed that each whorl represented a dental battery belonging to a large shark. The Belgian paleontologist Gustave Simoens illustrated Karpinsky’s various hypotheses and, drawing on histological data, asserted with certainty that the whorls were tooth-bearing structures located within the mouth. In 1911, Karpinsky depicted the whorls as components of the dorsal fins. In a 1996 textbook, Janvier presented a similar reconstruction, albeit with sharp teeth at the front of the upper jaw and rows of low crushing teeth in the back of the jaw. In 2008, American paleoartist Mary Parrish created a new reconstruction for the renovated Ocean Hall at the Smithsonian Museum of Natural History. Designed under the guidance of American researchers Robert W. Purdy, Victor Springer, and Matthew Carrano, Parrish's reconstruction places the whorl deeper within in the throat. This hypothesis was justified by the argument that the teeth supposedly had no wear marks, and the assumption that the whorl would have created a drag-inducing bulge on the chin of the animal if located in a symphyseal position. They envisioned the tooth whorl as a structure derived from throat denticles and designed to assist swallowing. This would hypothetically negate the disadvantages the tooth whorl would produce if positioned further forward in the jaw. This reconstruction was criticized for the overly intricate and potentially ineffective design of such a structure, if solely used to assist swallowing. In 2009, Lebedev provided further support for a reconstruction similar to those proposed by Bendix-Almgreen (1966). An H. bessonowi tooth whorl found in Kazakhstan preserved radial scratch marks; the whorl was also found near several wide, tuberculated teeth similar to those of the eugeneodont Campodus. Lebedev's reconstruction presented a cartilage-protected tooth whorl in a symphysial position at the front of the long lower jaw. When the mouth was closed, the tooth whorl would fit into a deep longitudinal pocket on the upper jaw. Both the pocket in the upper jaw and the edges of the lower jaw would have been lined with dense rows of Campodus-like teeth. This was similar to the situation reported in related helicoprionids such as Sarcoprion and Agassizodus. As for Helicoprion's ecology, it was compared to modern cetaceans such as Physeter (the sperm whale), Kogia (dwarf and pygmy sperm whales), Grampus (Risso's dolphin), and Ziphius (Cuvier's beaked whale). These fish-and squid-eating mammals (ichthyoteuthophage) have reduced dentition, often restricted to the tip of the lower jaw. Lebedev's reconstruction approximates modern views on Helicoprion's anatomy, though the hypothetical long jaw has been superseded by CT data since 2013. == Paleoecology ==
Paleoecology
Although the precise stratigraphic level of some specimens remains unknown, the genus Helicoprion is recorded over a span of roughly 20 million years during the Permian, from the Artinskian stage of the Cisuralian to the Roadian of the Guadalupian, i.e. approximately between 290 and 270 million years ago. More than half of the fossils assigned to the genus belong to H. davisii, primarily recovered from the Phosphoria Formation in Idaho. Approximately 25% represent H. bessonowi, known from the Divya Formation in the Ural Mountains of Russia. Other specimens come from more geographically dispersed deposits worldwide, including additional U.S. states as well as Australia, Canada, China, Kazakhstan, Japan, Laos, Mexico, and Norway. The wide geographic distribution of Helicoprion suggests it was a pelagic animal which may only have migrated into shallower waters periodically. During the Artinskian the Phosphoria Formation represented a deep inland sea with an anoxic, muddy bottom. Based on the large number of Helicoprion specimens discovered in the formation, the Phosphoria Sea may have been either a nursery habitat for breeding Helicoprion, or potentially a hunting ground. Helicoprion is believed to have been the apex predator of its environment. The genus' diet likely consisted primarily of cephalopods such as belemnoids, nautiloids and ammonoids, and it also may have hunted cartilaginous and unarmored bony fish as well. Naugolnykh has suggested that, in addition to nektonic cephalopods, Helicoprion fed on benthic invertebrates such as annelid worms caght on the seabed. == Extinction ==
Extinction
The disappearance of Helicoprion is not associated with a larger extinction event, and has been suggested to have been a background extinction. Because of its highly specialized feeding mechanism and large size, Helicoprion would have been particularly vulnerable to environmental or ecological changes, although the exact reason the genus died out is unknown. ==Notes==
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