Goniopholis Temporal range: Late Jurassic - Early Cretaceous, | |
---|---|
Holotype skull of the "Swanage Crocodile", G. kiplingi on display at the Dorset Museum. Berriasian age (earliest Cretaceous). | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Clade: | Crocodyliformes |
Family: | † Goniopholididae |
Genus: | † Goniopholis Owen, 1841 |
Type species | |
†Goniopholis crassidens Owen, 1841 | |
Species | |
Goniopholis (meaning "angled scale") is an extinct genus of goniopholidid crocodyliform that lived in Europe and North America during the Late Jurassic and Early Cretaceous. [1] [2] Like other goniopholidids, it resembled living crocodilians, and probably had a similar ecology as semi-aquatic ambush predators.
The type species of the genus is G. crassidens which is known from the Berriasian of England, and the referable species G. simus from the Berriasian of NW Germany, might be conspecific. Other species that are referable to Goniopholis include G. kiplingi from the Berriasian of England, and G. baryglyphaeus from the Late Jurassic (Kimmeridgian) of Portugal making it the oldest known Goniopholis species. [1] [2] The species G. kiplingi honors the author Rudyard Kipling, "in recognition for his enthusiasm for natural sciences". [1] G. kiplingi had skull reaching 475.6 mm (18.72 in), it is one of the largest goniopholidid along with Amphicotylus milesi which had skull reaching 43 cm (17 in). [1] [3] Based on skull length, total body length of G. kiplingi is estimated at 3.47 m (11.4 ft). [1]
Eggs attributed to Goniopholis were found in the Late Jurassic of Portugal. [4]
A partial skeleton of an indeterminate species of Goniopholis has been recovered from the Berriasian aged Angeac-Charente bonebed of France. [5]
Goniopholis have been inferred to have been ectothermic on the basis of bone histology and stable isotope analysis. [6]
Two species were referred to Goniopholis from Brazil. Goniopholis hartti from the Lower Cretaceous of Brazil is in fact a member of the genus Sarcosuchus . [1] G. paulistanus, based on two tooth crowns and a disassociated fragment of the right tibia from the Upper Cretaceous Bauru Group, has been reassigned to Itasuchidae and given its own genus Roxochampsa . [7]
From North America, G. lucasii and G. kirtlandicus are currently placed in their own genera Amphicotylus and Denazinosuchus , respectively, [1] while G. felix, G. gilmorei, and G. stovalli, all from the Morrison Formation, are referable to Amphicotylus and closely related to Eutretauranosuchus which are known from the same formation. [8] [9] [10]
G. phuwiangensis is known from NE Thailand, but this species is fragmentary and was recently reassigned to Sunosuchus . Nannosuchus from the Early Cretaceous (Berriasian stage) of England and Spain currently considered to be valid, was referred to as G. gracilidens by some authors. [1]
Willett's / Hulke's, Hooley's and Dollo's goniopholidids represent several complete specimens previously classified as either G. simus or G. crassidens, [1] and one of them was recently re-described as the new species, G. willetti. More recently these specimens were removed from Goniopholis, and two of them, Hooley's and Hulke's goniopholidids, have been already reassigned to their own genera Anteophthalmosuchus and Hulkepholis , respectively. [2] [11] Dollo's goniopholidid has also been assigned to Anteophthalmosuchus. [12]
Koumpiodontosuchus aprosdokiti from England was initially identified as a juvenile Goniopholis.
Like other goniophoilids, Goniopholis bears a superficial resemblance to modern crocodilians. However, unlike modern crocodilians and like other goniopholidids, the dermal armour covering the back was composed of two rows of large rectangular scutes running parallel down each side of the midline, with a "peg and groove" mechanism articulating the sets of plates together, with the outer edge of the plates deflected downwards. [1] [13]
Goniopholidids likely had a similar ecology to modern crocodilians as semi-aquatic ambush predators. [14]
Below is a cladogram including several Goniopholis species: [1]
Neosuchia |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Elosuchus is an extinct genus of neosuchian crocodyliform that lived during the Middle Cretaceous of what is now Africa.
Calsoyasuchus is a genus of crocodylomorph 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 specific name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones. It has often been interpreted as the earliest diverging member of Goniopholididae, but other studies have recovered it in various other positions.
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 neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia between the Middle Jurassic and the Late Cretaceous.
Baurusuchidae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America and possibly Pakistan. Baurusuchidae has been, in accordance with the PhyloCode, officially defined as the least inclusive clade containing Cynodontosuchus rothi, Pissarrachampsa sera, and Baurusuchus pachecoi. Baurusuchids have been placed in the suborder Baurusuchia, and two subfamilies have been proposed: Baurusuchinae and Pissarrachampsinae.
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. 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 in North America. It is 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. It was a semiaquatic crocodylomorph.
Amphicotylus is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was described in 1878.
Paramacellodus is an extinct genus of scincomorph lizards from the Early Cretaceous of England and France, and the Late Jurassic of Portugal and the western United States. The type species, Paramacellodus oweni, was named in 1967 from the earliest Cretaceous (Berriasian) Purbeck Group in Dorset, England. Additional material referable to a species of Paramacellodus, possibly P. oweni, has been described from the Morrison Formation, specifically in Como Bluff, Wyoming, and Dinosaur National Monument, Utah. An indeterminate species is known from the Berriasian aged Angeac-Charente bonebed in France. Paramacellodus belongs to an extinct family of scincomorphs called Paramacellodidae, which spanned most of Laurasia during the Late Jurassic and Early Cretaceous and represented one of the earliest evolutionary radiations of lizards.
Narynsuchus is an extinct genus of giant Goniopholidid Crocodylian. Because it has not yet been formally described, it is currently considered a nomen nudum. The name was first used in 1990 for fossil material found from a formation called the Balabansai Svita, which forms a lens in site FTA-30 of the Sarykamyshsai 1 locality in the Fergana Valley of Kyrgyzstan. The formation dates back to the Callovian stage of the Middle Jurassic. The type species of "Narynsuchus" is "N. ferganensis".Being semi-aquatic it is very similar to modern crocodiles. It ranged from 7–8 metres in length, and would have had a very similar lifestyle to the American alligator or Nile crocodile.
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.
Ziphosuchia is a clade of mesoeucrocodylian crocodyliforms that includes notosuchians and sebecosuchians.
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.
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.
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.
Roxochampsa is an extinct genus of crocodylomorph from the Late Cretaceous of Brazil belonging to the sebecosuchian clade Itasuchidae. The type species is R. paulistanus.