Dwykaselachus Temporal range: (Middle Permian) | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Order: | † Symmoriiformes |
Genus: | † Dwykaselachus Oelofsen, 1986 |
Type species | |
†Dwykaselachus oosthuizeni Oelofsen, 1986 |
Dwykaselachus (pronounced dwike-a-selak-us) 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. [1]
Prior to its discovery, symmoriiforms were thought to be related to sharks, in the group Elasmobranchii. However, CT scans of its relatively intact skull showed traits such as brain shape and inner ear structure that are shared with cartilaginous fish from the group Holocephali, which includes chimaeras. [2] This implies that the first major radiation of cartilaginous fish after the Devonian extinction was in fact holocephalians, rather than sharks as commonly believed. [3]
Dwykaselachus was first discovered in the 1980, in a nodule of sediments from the Dwyka Group of the Karoo Supergroup by amateur paleontologist Roy Oosthuizen, and originally described by Burger Wilhelm Oelofsen in 1986. [1]
In 2013, co-author Dr. Robert Gess, a researcher in the Geology Department and Albany Museum at Rhodes University in South Africa, CT-scanned [4] the skull of Dwykaselachus, and showed a symmoriiform morphology that resembles a 3D-preserved model. [5] At first, the skull was thought to belong a symmoriid shark, but after the CT-scanning, the image appeared to show anatomical structures that mark the specimen as an early relative of chimaeras. [6]
A research team led by Michael Coates from the University of Chicago Medical center has found that ghosts sharks, also named chimaeras, are related to the 280 million-year-old fish Dwykaselachus oosthuizeni. Chimaera-like features including tell-tale shapes of cranial nerves, nostrils and inner ears suggests that D. oosthuizeni was included in the group Symmoriiformes. [5] Although resembling sharks in appearance, Dwykaselachus was not actually a shark, but rather had diverged from a common ancestor with true sharks in the Devonian. [7]
The computed tomography (CT) analysis of Dwykaselachus shows a symmoriiform morphology with three-dimensional articulation. [8] [4] It exhibits some chondrichthyans features such as the large hepophyseal chamber and dorsally projecting endolymphatic duct. [2] The most visible shared specialization with chimaeroids is the offset between the dorsally prominent mesencephalon chamber and the ventral level of the telencephalon space. Moreover, Dwykaselachus share the characteristic chimaeroid elevation of the midbrain, relative to forebrain. [2]
The discovered skull has unusually ethmoid cartilages which include large hemispherical nasal capsules. [3] The nasal capsules are bridged by an internasal groove. Each capsule roof is shorter than the floor, suggesting that, unlike many sharks, the narial openings were directed slightly dorsally. [3] The capsule wall openings include a canal for the olfactory nerve (nerve I), a foramen for the profundus nerve (nerve V), and an opening in the floor, which resembles the subnasal fenestra of Doliodus. [3] The braincase roof is mostly complete, leave little space for the fontanelle. Therefore, a precerbral fontanelle, a signature of non-chimaeroid chondrichthyans, is either reduced or absent.
The phylogenetic analyses establish the importance of the shared similarities between Dwykaselachus and chimaeroids, suggesting symmoriiforms, including Dwykaselachus, comprise a sister clade to iniopterygians and holocephalans. [9] The detailed fossil chondrichthyans data used in Coates et al.’s paper provided strong evidence supporting that hypothesis. [10] Thus, expansion of holocephalian stem membership moves the chondrichthyans crown group divergence to a deeper phylogenetic node. [3]
Another study done by Coates et al. showed that phylogenetic analysis confirms Gladbachus as a stem chondrichthyan. Strong evidence was provided to support chondrichthyans as a crown clade. [11] It suggests that the initial evolutionary radiation of crown chondrichthyans is primarily post-Devonian, forming a significant component of the vertebrate recovery after the end-Devonian extinction. [6]
The current analyses focus on the conflicting patterns of character state distributions, implying repeated and convergent evolution of chondrichthyan-like specializations among the earliest total group members. [12] Early chondrichthyan species suggest that the morphological disparity in the early members of the chondrichthyan total group was probably substantially greater than that which is observed. [11]
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 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.
Holocephali, sometimes given the term Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primitive forms, and the only surviving group in the subclass is the order Chimaeriformes.
Placoderms are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Devonian periods. While their endoskeletons are mainly cartilaginous, their head and thorax were covered by articulated armoured plates, and the rest of the body was scaled or naked depending on the species.
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.
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.
The ptyctodontids ("folded-teeth") are placoderms of the order Ptyctodontida, containing the family Ptyctodontidae. With their big heads, big eyes, reduced armor and long bodies, the ptyctodontids bore a superficial resemblance to modern day chimaeras (Holocephali). Their armor was reduced to a pattern of small plates around the head and neck. Like the extinct and related acanthothoracids, and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on shellfish.
Symmoriiformes is an extinct order of stem-group holocephalians. Originally named Symmoriida by Zangerl (1981), it has subsequently been known by several other names. Lund (1986) synonymized the group with Cladodontida, while Maisey (2008) corrected the name to Symmoriiformes in order to prevent it from being mistaken for a family. The symmoriiform fossils record begins during the late Devonian. Most of them died out at the start of the Permian, but Dwykaselachus is known from the Artinskian-Kungurian of South Africa. Teeth described from the Valanginian of France and Austria indicate that members of the family Falcatidae might have survived until the Early Cretaceous; however, these teeth were also argued to be more likely neoselachian teeth.
Symmoriidae is an extinct family of cartilaginous fish belonging to the order Symmoriiformes. Merbers of the family are known from the Devonian and Carboniferous periods.
Falcatus is an extinct genus of falcatid chondrichthyan which lived during the early Carboniferous Period in Bear Gulch bay in what is now Montana.
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.
Symmorium is a dubious genus of extinct stethacanthid cartilaginous fish from the Devonian and Carboniferous of the United States (Illinois) and Russia. The type species, Symmorium reniforme, was named by Edward Drinker Cope in 1893 and several other species were originally classed under this genus, but they have since been classified into other genera such as Petalodus. Symmorium bears close similarity in size and appearance to Stethacanthus but the former is missing the "spine and brush" on its back. Some paleontologists think that the two forms are simply the males and females of related species, while other scientists think they were distinct genera.
Falcatidae is a family of Paleozoic cartilaginous fish belonging to the order Symmoriiformes. Members of this family include Falcatus, a small fish from the Bear Gulch Limestone of Montana. The family first appeared around the start of the Carboniferous, and there is some evidence that they survived well into the early Cretaceous, though its putative Cretaceous members were also argued to be more likely neoselachians.
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.
Ptomacanthus is an extinct genus of spiny shark, an early relative of living cartilaginous fishes.
Plesioselachus is an extinct genus of Late Devonian (Famennian) cartilaginous fish with uncertain classification, which contains only one species, P. macracanthus from the Waterloo Farm lagerstätte in South Africa. Known from a single incomplete articulated skeleton and some isolated remains, it is characterized by having a long dorsal spine with length about one third of body length.
Ozarcus is an extinct genus of symmoriiform cartilaginous fish from the Carboniferous period of Arkansas. The type species, Ozarcus mapesae, was named in 2014 based on cartilaginous skulls from the Serpukhovian-age Fayetteville Formation. The genus is named after the Ozark Mountains while the species was named after its discoverer, G. K. Mapes.