Dracopristis Temporal range: Carboniferous (Kasimovian), | |
---|---|
![]() | |
An illustration of D. hoffmanorum | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Subclass: | Elasmobranchii |
Order: | † Ctenacanthiformes |
Family: | † Heslerodidae |
Genus: | † Dracopristis Hodnett et al. 2021 |
Species: | †D. hoffmanorum |
Binomial name | |
†Dracopristis hoffmanorum Hodnett et al. 2021 | |
Dracopristis (meaning 'dragon shark') is an extinct genus of ctenacanth (a group of shark-like cartilaginous fish) 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, USA. 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".
Dracopristis possessed large dermal denticles along its head, along with rows of short, multi-cusped teeth in its jaws and very large spines on its dorsal fins, the latter of which inspired the name of the genus. The dorsal spines of the type specimen are about 0.57 m (1 ft 10 in) in length, while the entire body was approximately 2 m (6 ft 7 in) in length. Its large spines were likely used for defense against larger fish, while Dracopristis itself was likely a benthic predator that inhabited shallow, brackish-water environments.
The holotype of D. hoffmanorum was first discovered in May of 2013, [1] [2] when John-Paul Hodnett unearthed the specimen from the late Carboniferous-aged strata of Albuquerque, New Mexico. [2] [3] [4] The fossils were discovered within the Tinajas member of the Atrasado Formation, which, based on index fossils (such as conodonts), has been dated to the Kasimovian stage of the late Pennsylvanian, which is equivalent to the American Missourian. [1] [5] The specific locality which produced the specimen, the Kinney Brick Quarry, is considered a Konservat-Lagerstätten due to the exceptional quality of its fossils. [5]
Dracopristis was formally named, described, and classified by Hodnett and coauthors in 2021. [4] [5] The holotype (catalogued as NMMNH P-68537) is an articulated skeleton, embedded in a matrix of calcareous shale. [5] This specimen belonged to an adult female and represents the most complete ctenacanth fossil so far discovered. [5] [6] Around 87–90% of the cartilaginous skeleton, the animal's coating of tooth-like dermal denticles ,and impressions of its body outline are all preserved. [4] [5] The specimen was studied via CT scanning in 2014, which clarified details of its skeletal anatomy that were otherwise obscured by the matrix. [6] A second assigned specimen (NMMNH P-19181) consists of a neurocranium from a juvenile individual, and was originally misidentified as belonging to Orthacanthus huberi. [5] [7]
The genus name, Dracopristis, is derived from the Latin words draco, meaning 'dragon', and pristis, meaning 'shark'. [5] The dorsal spines, facial denticles, and rows of teeth have been suggested to give the animal a "dragon-like" appearance, [2] [3] [5] and -pristis is a common suffix given to shark-like fishes. [2] [5] [8] [note 1] The specific name, hoffmanorum, is in honor of Ralph and Jeanette Hoffman who own the Kinney Brick Quarry and helped conduct research on the taxon. [5] Prior to its scientific description, what is now the holotype of Dracopristis hoffmanorum was informally referred to as the "Godzilla shark" or, [4] [12] alternatively, the "Manzano ctenacanth" after the Manzano Mountains. [1] [13] The "Godzilla" nickname was inspired by the genus' defining exaggerated back spines and teeth, which are said to resemble the famous movie monster of the same name. [3] [4]
The type specimen of Dracopristis hoffmanorum is around 2.06 m (6 ft 9 in) in length, [5] although some earlier sources erroneously state lengths of up to 2.5 m (8 ft 2 in). [1] [6] Although the specimen is crushed, the skull, gill arches and dorsal fins remain in articulation, and the pelvic girdle and anal fins are particularly well preserved. [1] D. hoffmanorum had an elongated and dorsoventrally (from top to bottom) flattened body, which is also indicated by the shape of preserved soft tissue impressions and the arrangement of articulated patches of dermal denticles. [5]
The skull of Dracopristis was broad – nearly as wide as it was long – with eye sockets positioned far forward and a short rostrum. Fenestrae which housed cranial nerves are preserved, but the interior of the skull is too severely crushed for its internal nervous anatomy to be studied, even with the use of CT scans. Compared to other ctenacanths, the jaws were proportionally large and robust. As in living sharks, the rear portions of the palatoquadrates (upper jaws) articulated with the otic processes (equivalent to ear canals). However, unlike living sharks, the forward processes of the palatoquadrates were rigidly connected to the cranium, and additional "anterodorsal processes" anchored the rear portions of the palatoquadrates to the skull, rendering the upper jaws akinetic and inflexible (termed amphistyly). The Meckel's cartilages (lower jaws) articulated with the palatoquadrates. The hyomandibular arch was well-developed, and five gill arches were present which descended in size towards the posterior (back) of the body. [5]
The scapulocoracoids were fused at their coracoidal portions, while the pelvic girdles were unfused. Mineralized neural and haemal arches are preserved along the length of the body, although the vertebral centra are not known. The holotype of D. hoffmanorum preserves two dorsal fins, an anal fin, pectoral fins, pelvic fins, and most of the heterocercal caudal fin. Both dorsal fins were proportionally large, [5] as were the broad, triangular pectoral fins. [2] The fins were supported by rods of cartilage known as radials and ceratotrichia, with a triangular basal cartilage plate also supporting each of the dorsal fins. While the known individual lacks pelvic claspers, this is attributed to the specimen being female, and males of the species are presumed to have possessed these. The anal fin was supported by a plate of cartilage, and was rounded in shape. [5]
Compared to its relatives, Dracopristis possessed proportionally very large dorsal fin spines. [1] [14] In the type specimen, the anterior (front) spine is roughly 57 cm (22 in) long (about 27% of the length of the body), while the shorter posterior (back) spine is roughly 40 cm (16 in) long. Both spines are ornamented with rows of small, rounded denticles along their lateral sides and two rows of larger, recurved denticles along their anterior sides. The first spine is strongly swept backwards, while the second is much straighter. Both dorsal spines articulate with the basal cartilages of the dorsal fins. [5]
The skin was coated in fine dermal denticles, the size and shape of which varied depending on their position on the body. The largest of these were leaf-shaped and distributed along the back of the head, whereas the smaller, shorter denticles were present along the fins and rostrum. The bases of the dermal denticles were wide, and while they were tightly packed, they did not overlap with one another. Mucous membrane denticles up to 5 mm (0.20 in) in diameter were present within the mouth and gill basket. [5]
The largest teeth of D. hoffmanorum could reach up to 2 cm (0.79 in) in width, and were arranged into 12 laterally (towards the outside) positioned rows. [1] [4] Their appearance is similar to those of other ctenacanthiforms such as Glikmanius and Heslerodus in sharing a cladodont (multi-cusped) crown morphology and a reniform (d-shaped) root. [1] Unlike other known ctenacanths, however, the cusps on the teeth of Dracopristis are relatively short, broad, and triangular in shape. [1] [5] The teeth possess five cusps each, with the central cusp being more than twice the height of the outer four. The tooth cusps are ornamented with multiple rows of denticles down their length, the arrangement of which is unique to this species. The dentition is morphologically homodont (teeth do not vary in shape), although the rows of teeth do decrease in size further into the mouth. The teeth on the frontmost row are over twice the crown height of those of the backmost row. In the 2021 description, it is suggested that tooth replacement was very slow. [5]
Phylogenetic position of D. hoffmanorum as reconstructed by Hodnett and colleagues in 2021. [5] |
Dracopristis was part of the chondrichthyan order Ctenacathiformes, [5] which were likely part of the subclass Elasmobranchii and were thus relatives of living sharks and rays. [5] [15] Despite often being informally called "sharks", [5] [12] [16] ctenacanths were outside of the clade Selachimorpha and are not considered "true" sharks. [17] Compared with selachimorph sharks, the ctenacanths had greatly enlarged ornamented fin spines, proportionally large mouths, and akinetic jaw suspension. [4] [5] Genera such as Bandringa and Sphenacanthus (traditionally considered ctenacanthiforms) [18] [19] have been recovered elsewhere within the Euselachii, indicating the group as traditionally defined may not be monophyletic. [5] [15] Rather than being elasmobranchs, it has alternatively been suggested that ctenacanths instead diverged much earlier and were on the stem of Chondrichthyes. [20]
In the 2021 description of Dracopristis hoffmanorum, cladistic analyses were performed which found that the genus was most closely related to the Devonian genus Ctenacanthus. [5] It was also suggested that ctenacanthiforms are closer to crown group euselachians than to other cladodont chondricthyans such as Symmoriiformes. [5] In a 2024 publication describing the ctenacanths Troglocladodus trimblei and Glikmanius careforum, Hodnett and coauthors placed Dracopristis in the newly-erected family Heslerodidae alongside the genera Glikmanius, Heslerodus, Avonacanthus , and Kaibabvenator . [16]
During the Pennsylvanian, New Mexico was covered by a vast seaway. [4] Dracopristis would have lived in the shallow coastal waters of this seaway, and probably would have been an ambush predator. [2] It may have hunted small animals such as fish and crustaceans, [21] and its teeth show adaptations for grasping and crushing its prey. [2] [4] As in other ctenacanths, its dorsal fin spines may have served as protection from larger predators. [4] [14] The shape of Dracopristis' pectoral fins suggests it lived a nektobenthic (bottom dwelling) lifestyle, due to similarities to those of living benthic sharks. [2] D. hoffmanorum may have been a specialist of brackish-water habitats, and its ecology has been compared with that of the bull shark and common sawfish. [2] [5]
During the Carboniferous, the Atrasado Formation consisted of estuarine and lagoonal habitats, [1] evidenced by fish specimens which show adaptations to both freshwater and marine environments. [21] The quality of many of the Kinney Brick Quarry fossils may have been the result of an anoxic zone, which possibly prevented scavengers disturbing them. [21] Dracopristis shared the estuary with the larger, related ctenacanthiform Glikmanius occidentalis, [5] which may have preyed upon it. [2] [21] The quarry has also yielded numerous specimens of the filter feeding acanthodian Acanthodes , symmoriiforms similar to Cobelodus , [1] hybodonts, holocephalans, ray-finned palaeonisciformes, lungfish, coelacanths, and megalichthyoforms. [1] [21] Rarer genera, such as the large eugeneodont (whorl-tooth shark) Campyloprion and the aforementioned Glikmanius, might have only occasionally migrated into the estuary from deeper, marine habitats. [5] [21] In total, more than 31 other distinct fish genera have been identified from the site, [21] [22] and it is considered one of the most diverse and well preserved fossil fish faunas in the Southwestern United States. [1]
Chimaeras are cartilaginous fish in the order Chimaeriformes, known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last two names are also applied to Opisthoproctidae and Siganidae, respectively.
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 a genus of extinct 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.
The Bahamas sawshark, Pristiophorus schroederi, is a sawshark of the family Pristiophoridae, found in the western Central Atlantic Ocean from the Bahamas and Cuba at depths of between 400 and 1,000 m. These sharks are at least 80 cm long.
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 viper dogfish or viper shark is a rare species of dogfish shark in the family Etmopteridae, and the only extant member of its genus. It has been found in the Pacific Ocean off southern Japan, the Bonin Islands, Pacific Ocean off northern Taitung County and the Northwestern Hawaiian Islands. This species inhabits upper continental slopes and seamounts. It may migrate vertically, shifting between bottom waters 270–360 m (890–1,180 ft) deep during the day and upper waters less than 150 m (490 ft) deep at night. A slender, black shark reaching 54 cm (21 in) in length, the viper dogfish can be recognized by its narrow, triangular jaws and well-spaced, fang-like teeth. It also has two spined dorsal fins, dermal denticles with faceted crowns, and numerous light-emitting photophores concentrated on its ventral surface.
The sailfin roughshark is a species of dogfish shark in the family Oxynotidae, found in the eastern North Atlantic from Scotland to Senegal between latitudes 61°N and 11°N, at depths of between 265 and 720 m. Its length is up to 1.2 m.
The blurred lanternshark is a little-known species of dogfish shark in the family Etmopteridae, found around the world in benthic and pelagic habitats from a depth of 110 m (360 ft) to over 1 km (0.62 mi) down. This shark forms the E. pusillus species group with the smooth lanternshark, which are distinguished from other members of its family by having irregularly arranged, flat-topped dermal denticles that give them a "smooth" appearance. Both species are slender-bodied with long heads, two dorsal fins bearing spines, no anal fins, and light-emitting photophores. The blurred lanternshark is larger, reaching 67 cm (26 in) or more in length. This species feeds on small squid, fishes, and fish eggs, and is ovoviviparous. It has been assessed as of Least Concern by the International Union for Conservation of Nature, because of its wide distribution and lack of threat from fishing pressure.
The quagga catshark is a species of shark belonging to the family Pentanchidae, the deepwater catsharks. 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 green lanternshark is a species of dogfish shark in the family Etmopteridae, found in the western central Atlantic Ocean. This species usually occurs on the upper continental slope below a depth of 350 m (1,150 ft). Reaching 26 cm (10 in) in length, the green lanternshark has a slender body with a long, thin tail and low, conical dermal denticles on its flanks. It is dark brown or gray with ventral black coloration, which contain light-emitting photophores that may serve a cryptic and/or social function. Green lanternsharks are thought to be gregarious and may attack their prey, squid and octopus often larger than themselves, in packs. Reproduction is aplacental viviparous, with females giving birth to litters of one to three young. This relatively common shark is an occasional, valueless bycatch of commercial fisheries; currently it does not appear to be significantly threatened by human activities.
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.
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, in reference to the aberrant quality and size of the species' teeth. 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.
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.
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.
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.
Tribodus is an extinct genus of hybodont. It lived during the mid Cretaceous (Albian-Cenomanian) with fossils being known from northern South America, North Africa, and southern Europe.
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.
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.
The Eugeneodontidae is an extinct family of cartilaginous fish known from the Carboniferous, Permian, and potentially Devonian periods of the United States, Iran, and possibly Belgium, Poland and Russia. The eugeneodontids are known primarily from fossilized teeth, as well as the remains of the jaws, gills and in some species nearly complete skeletons with coatings of dermal denticles. The genera Bobbodus, Gilliodus and the type genus Eugeneodus are included within the family. Eugeneodontidae is considered the sister group to the family Caseodontidae, and the two are included together within the suborder Caseodontoidea and the order Eugeneodontida. The type genus, and by extension the family and the order Eugeneodontida are named in honor of Paleontologist Eugene S. Richardson Jr.