Coelognathosuchia | |
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Skull of Goniopholis simus | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Neosuchia |
Clade: | † Coelognathosuchia Martin et al., 2014 |
Families | |
Coelognathosuchia is an extinct clade of neosuchian crocodyliforms that includes all taxa more closely related to the family Pholidosauridae than to Bernissartia fagesii or Eusuchia. Martin et al. (2014) named the clade after finding goniopholidids and pholidosaurids to group together in their phylogenetic analysis of crocodyliform evolutionary relationships. In their analysis, Pholidosauridae was monophyletic and Goniopholididae was paraphyletic, being an assemblage of successively more basal taxa within Coelognathosuchia. Coelognathosuchia itself was positioned near the base of the larger clade Neosuchia as the sister group to a clade containing the Early Cretaceous neosuchian Bernissartia and Eusuchia, the group that includes all modern crocodilians and their closest extinct relatives. [1]
Martin et al. named Coelognathosuchia from the Greek κοῖλος (koĩlos, "concave"), γνάθος (gnáthos, "jaw") and σοῦχος (soũchos, "crocodile"), after a small depression on the surface of the skull between the maxilla and jugal bones in both goniopholidids and pholidosaurs. Other diagnostic features of Coelognathosuchia include orbits (eye sockets) that are narrower than the frontal bone that separates them and smaller than the supratemporal fenestrae (two holes at the back of the skull roof) behind them, a notch between the premaxilla and maxilla at the tip of the snout, and the reduction or absence of the antorbital fenestrae (a pair of holes in the snout in front of each orbit). [1]
A close relationship between Goniopholididae and Pholidosauridae conflicts with the hypothesis that pholidosaurids are instead more closely related to the family Dyrosauridae. [1] This alternate phylogeny was found in many analyses, including Jouve et al. (2006), [2] Pol and Gasparini (2009), [3] and de Andrade et al. (2011), who named the clade Tethysuchia. [4] In both studies, Goniopholididae was found to be a more distantly related clade within Neosuchia. In their description of Pholidosaurus specimens from southwestern France, Martin and colleagues reiterated their opinion that Dyrosauridae is not as closely related to Pholidosauridae as stated by previous authors by pointing to numerous shared characters between the Cherves-de-Cognac pholidosaurid remains and Goniopholididae. [5]
Thalattosuchia is the name given to a clade of marine crocodylomorphs from the Early Jurassic to the Early Cretaceous that had a cosmopolitan distribution. They are colloquially referred to as marine crocodiles or sea crocodiles, though they are not members of Crocodilia. The clade contains two major subgroupings, the Teleosauroidea and Metriorhynchoidea. Within Metriorhynchoidea, the Metriorhynchidae displayed extreme adaptions for life in the open ocean, including the transformation of limbs into flippers, the development of a tail fluke, and smooth, scaleless skin.
Dyrosauridae is a family of extinct neosuchian crocodyliforms that lived from the Late Cretaceous (Maastrichtian) to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South America. Over a dozen species are currently known, varying greatly in overall size and cranial shape. A majority were aquatic, some terrestrial and others fully marine, with species inhabiting both freshwater and marine environments. Ocean-dwelling dyrosaurids were among the few marine reptiles to survive the Cretaceous–Paleogene extinction event.
Elosuchus is an extinct genus of neosuchian crocodyliform that lived during the Middle Cretaceous of what is now Africa.
Calsoyasuchus is a genus of goniopholidid mesoeucrocodylian that lived in the Early Jurassic. Its fossilized remains were found in the Sinemurian-Pliensbachian-age Kayenta Formation on Navajo Nation land in Coconino County, Arizona, United States. Formally described as C. valliceps, it is known from a single incomplete skull which is unusually derived for such an early crocodile relative. This genus was described in 2002 by Ronald Tykoski and colleagues; the species name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones.
Rugosuchus is an extinct genus of neosuchian crocodyliform from the late Early Cretaceous of China. It is known from most of a skull, a partial postcranial skeleton, and a second partial skeleton including part of the hips. It was described by Xiao-Chun Wu and colleagues in 2001, with R. nonganensis as the type species. At the time of its description, it was the most complete crocodyliform from northeastern China, and only the second known.
Goniopholididae is an extinct family of moderate-sized semi-aquatic crocodyliforms superficially similar to living crocodiles. They lived between the Early Jurassic and the Late Cretaceous.
Pholidosauridae is an extinct family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe, Africa, North America and South America. The pholidosaurids first appeared in the fossil record during the Bathonian stage of the Middle Jurassic. Jouve & Jalil (2020) described postcranial material of a pholidosaurid from the Paleocene (Danian) of Ouled Abdoun Basin (Morocco), representing the most recent record of the family. The authors also reinterpreted putative Maastrichtian dyrosaurid Sabinosuchus as a pholidosaurid, and argued that at least two independent pholidosaurid lineages reached the Maastrichtian, among which one survived the Cretaceous–Paleogene extinction event. Before the publication of this study it was thought that the family became extinct during the Late Turonian stage of the Late Cretaceous.
Eutretauranosuchus is an extinct genus of goniopholidid crocodyliform. E. delfsi is the only known species within the genus.
Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to Crocodylus niloticus than to Notosuchus terrestris. Neosuchia is very diverse and may be polyphyletic, as the clade has undergone many revisions since it was first named in 1988. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages, often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.
Stolokrosuchus is an extinct genus of neosuchian crocodylomorph that lived during the Early Cretaceous. Its fossils, including a skull with a long thin snout and bony knobs on the prefrontal, have been found in Niger. Stolokrosuchus was described in 2000 by Hans Larsson and Boubacar Gado. The type species is S. lapparenti. They initially described it as related to Peirosauridae, if not a member of that family. One study has shown it to be related to Elosuchus. However, more recent works usually find Stolokrosuchus to be one of the basalmost neosuchian, only distantly related to the elosuchid or pholidosaurid, Elosuchus.
Amphicotylus is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was described in 1878.
Mahajangasuchidae is an extinct family of neosuchian crocodyliforms. It currently contains two genera, Mahajangasuchus and Kaprosuchus, both of which lived during the Late Cretaceous in Gondwana. It is defined as the most inclusive clade containing Mahajangasuchus insignis but not Notosuchus terrestris, Simosuchus clarki, Araripesuchus gomesii, Baurusuchus pachecoi, Peirosaurus torminni, Goniopholis crassidens, Pholidosaurus schaumbergensis, or Crocodylus niloticus. Phylogenetically, Mahajangasuchidae is placed just outside pholidosaurids and more derived neosuchians.
Metasuchia is a major clade within the superorder Crocodylomorpha. It is split into two main groups, Notosuchia and Neosuchia. Notosuchia is an extinct group that contains primarily small-bodied Cretaceous taxa with heterodont dentition. Neosuchia includes the extant crocodylians and basal taxa, such as peirosaurids and pholidosaurids. It is phylogenetically defined by Sereno et al. (2001) as a clade containing Notosuchus terrestris, Crocodylus niloticus, and all descendants of their common ancestor.
Pholidosaurus is an extinct genus of neosuchian crocodylomorph. It is the type genus of the family Pholidosauridae. Fossils have been found in northwestern Germany. The genus is known to have existed during the Berriasian-Albian stages of the Early Cretaceous. Fossil material found from the Annero and Jydegård Formations in Skåne, Sweden and on the island of Bornholm, Denmark, have been referred to as a mesoeucrocodylian, and possibly represent the genus Pholidosaurus.
Sunosuchus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils are known from China, Kyrgyzstan, and Thailand and are Jurassic in age, although some may be Early Cretaceous. Four species are currently assigned to the genus: the type species S. miaoi and the species S. junggarensis, S. shartegensis, and S. shunanensis. All species are from China. Goniopholis phuwiangensis, also from Thailand, was reassigned to Sunosuchus by Andrade et al. (2011). The material from Kyrgyzstan has not been assigned to any species.
Anteophthalmosuchus is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England, eastern Spain, and western Belgium.
Tethysuchia is an extinct clade of neosuchian mesoeucrocodylian crocodylomorphs from the late Middle Jurassic to the Early Eocene of Asia, Europe, North America and South America. It was named by the French paleontologist Eric Buffetaut in 1982 as a suborder. Tethysuchia was considered to be a synonym of Dyrosauridae or Pholidosauridae for many years. In most phylogenetic analyses the node Dyrosauridae+Pholidosauridae was strongly supported. De Andrade et al. (2011) suggested that Tethysuchia be resurrected for that node. They defined it as a node-based taxon "composed of Pholidosaurus purbeckensis and Dyrosaurus phosphaticus, their common ancestor and all its descendants". In their analysis they found that the support for Tethysuchia is actually stronger than the support for Thalattosuchia. The following cladogram shows the position of Tethysuchia among the Neosuchia sensu this study.
Paluxysuchus is an extinct genus of neosuchian crocodyliform known from the Early Cretaceous Twin Mountains Formation of north-central Texas. It contains a single species, Paluxysuchus newmani. Paluxysuchus is one of three crocodyliforms known from the Early Cretaceous of Texas, the others being Pachycheilosuchus and an unnamed species referred to as the "Glen Rose Form". Paluxysuchus has a long, flat skull that is probably transitional between the long and narrow skulls of many early neosuchians and the short and flat skulls of later neosuchians.
Hulkepholis is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England and eastern Spain. It contains two species, the type species, Hulkepholis willetti, and also H. plotos. Hulkepholis is most closely related to both species of Anteophthalmosuchus.
Chalawan is an extinct genus of pholidosaurid mesoeucrocodylian known from the Late Jurassic or Early Cretaceous Phu Kradung Formation of Nong Bua Lamphu Province, northeastern Thailand. It contains a single species, Chalawan thailandicus, with Chalawan shartegensis as a possible second species.