Goniopholididae

Goniopholididae is an extinct family of moderate-sized semi-aquatic crocodyliforms superficially similar to living crocodiles (but see below). They lived between the Early Jurassic and the Late Cretaceous.[1][2]

Goniopholididae
Temporal range: 196–66 Ma
Sinemurian - Maastrichtian
Goniopholis simus 3
Anteophthalmosuchus fossil, Musee d'Histoire Naturelle, Brussels
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Neosuchia
Family: Goniopholididae
Cope, 1875
Subgroups

See text.

Phylogeny

The following cladogram simplified after an analysis presented by Marco Brandalise de Andrade and colleagues in 2011.[3]

Neosuchia
Atoposauridae

Theriosuchus pusillus

Theriosuchus guimarotae

Rugosuchus

Bernissartia

Eusuchia

Stolokrosuchus

Tethysuchia

Thalattosuchia

Goniopholididae

Calsoyasuchus valliceps

"Goniopholis" phuwiangensis

Eutretauranosuchus delfi

"Sunosuchus" junggarensis

Sunosuchus miaoi

Chalawan thailandicus

Siamosuchus phuphokensis

Amphicotylus lucasii

Denazinosuchus kirtlandicus

Nannosuchus gracilidens

Hulkepholis willetti

Anteophthalmosuchus

Dollo’s goniopholidid

Goniopholis

Goniopholis baryglyphaeus

Goniopholis kiplingi

Goniopholis simus

Geographical distribution

Goniopholidids are known across Laurasia, ranging from North America, Europe and China from the Middle Jurassic,[1][4] and reaching Thailand by the Early Cretaceous.[5]

Biology

Compared to modern crocodilians, goniopholids are very unusual in several respects. They possessed two rows of rectangular, interlocking osteoderms like those of terrestrial crocodilymorphs like atoposaurids, that are relatively simple, do not extend far in their necks, as opposed to the ornate armours of modern crocodilians; likewise, unlike modern crocodilians but like many extinct forms like phytosaurs, they have ventral osteoderms as well. Their forelimbs are also proportionally very long, particularly in the humeri and wrist bones, being as long or longer than the hindlimbs, the opposite of the condition seen in modern crocodilians. Some like Anteophthalmosuchus also have forwardly oriented eyes, as opposed to the dorsally oriented eyes seen in modern forms. These suggest multiple biomechanical differences from modern species.[6][7]

References

  1. ^ a b Steel R. 1973. Crocodylia. Handbuch der Paläoherpetologie, Teil 16. Stuttgart: Gustav Fischer Verlag, 116 pp.
  2. ^ Tykoski RS, Rowe TB, Ketcham RA, Colbert MW. 2002. Calsoyasuchus valliceps, a new crocodyliform from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology 22 (3): 593-611.
  3. ^ Marco Brandalise de Andrade; Richard Edmonds; Michael J. Benton; Remmert Schouten (2011). "A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus". Zoological Journal of the Linnean Society. 163 (s1): S66–S108. doi:10.1111/j.1096-3642.2011.00709.x.
  4. ^ Maisch MW, Matzke AT, Stohr H. 2003. Sunosuchus (Archosauria, Crocodyliformes) from the Toutunhe Formation (Middle Jurassic) of the Southern Junggar Basin (Xinjiang, NW-China). Geobios 36 (4): 391-400.
  5. ^ Lauprasert, K.; Cuny, G.; Buffetaut, E.; Suteethorn, V.; Thirakhupt, K. (2007). "Siamosuchus phuphokensis, a new goniopholidid from the Early Cretaceous (ante-Aptian) of northeastern Thailand". Bulletin de la Société Géologique de France. 178 (3): 201–216. doi:10.2113/gssgfbull.178.3.201.
  6. ^ Salisbury, S. W. & Frey, E. 2000. A biomechanical transformation model for the evolution of semi-spheroidal articulations between adjoining vertebral bodies in crocodilians. In Grigg, G. C., Seebacher, F. & Franklin, C. E. (eds) Crocodilian Biology and Evolution. Surry Beatty & Sons (Chipping Norton, Aus.), pp. 85-134.
  7. ^ Salisbury, S. W. & Naish, D. (2011). Crocodilians. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 305-369.
Amphicotylus

Amphicotylus is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado and Oklahoma.

Anteophthalmosuchus

Anteophthalmosuchus (meaning "forward-pointing eye crocodile") is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England, eastern Spain, and western Belgium.

Calsoyasuchus

Calsoyasuchus (meaning "[Dr. Kyril] Calsoyas' crocodile") 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.

Chalawan (genus)

Chalawan (from Thai: ชาละวัน [t͡ɕʰāːlāwān]) is an extinct genus of pholidosaurid mesoeucrocodylian known from the Early Cretaceous Phu Kradung Formation of Nong Bua Lamphu Province, northeastern Thailand. It contains a single species, Chalawan thailandicus.

Coelognathosuchia

Coelognathosuchia is an extinct clade of neosuchian crocodyliforms that includes Goniopholididae and Pholidosauridae, two families of superficially crocodile-like aquatic crocodyliforms from the Mesozoic. 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.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 diagonostic 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).A close relationship between Goniopholididae and Pholidosauridae conflicts with the hypothesis that pholidosaurids are instead more closely related to the family Dyrosauridae. This alternate phylogeny was found in many analyses, including Jouve et al. (2006), Pol and Gasparini (2009), and de Andrade et al. (2011), who named the clade Tethysuchia. 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.

Coelosuchus

Coelosuchus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils have been found from the Graneros Shale of the Benton Group in Wyoming, and are of Cenomanian age. It was slightly over 1 meter in length.

Dakotasuchus

Dakotasuchus (meaning "Dakota [Sandstone] crocodile") is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Cenomanian-age Upper Cretaceous Dakota Sandstone of Kansas. The type specimen was found in an iron-cemented sandstone concretion near Salina. This concretion was broken into two large pieces; more of the specimen was probably present originally, but by the time it was found only the torso and short portions of the neck and tail remained. Twenty pairs of bony scutes ran down the midline of the back. The vertebrae lacked the procoelous articulation (concave anterior and convex posterior faces) of more derived crocodyliforms. Dakotasuchus had short broad shoulder blades, suggesting it had stout powerful forelimbs and perhaps terrestrial habits. M. G. Mehl, who described the genus, estimated the length of the type individual when complete to have been 3–4 metres (9.8–13.1 ft). The type species is D. kingi, named for Professor King, a former dean of Kansas Wesleyan University. Mehl did not classify his new genus to a more inclusive group than Mesosuchia (a paraphyletic group replaced by Mesoeucrocodylia). Robert Carroll assigned Dakotasuchus to Goniopholididae in 1988.

Garden Park, Colorado

Garden Park is a paleontological site in Fremont County, Colorado, known for its Jurassic dinosaurs and the role the specimens played in the infamous Bone Wars of the late 19th century. Located 10 km (6.2 mi) north of Cañon City, the name originates from the area providing vegetables to the miners at nearby Cripple Creek in the 19th century. Garden Park proper is a triangular valley surrounded by cliffs on the southeast and southwest and by mountains to the north; however, the name is also refers to the dinosaur sites on top and along the cliffs. The dinosaur sites now form the Garden Park Paleontological Resource Area, which is overseen by the Bureau of Land Management.

Goniopholis

Goniopholis is an extinct genus of goniopholidid crocodyliform that lived in Europe and Africa during the Late Jurassic and Early Cretaceous. Being semi-aquatic it is very similar to modern crocodiles. It ranged from 2–4 metres in length, and would have had a very similar lifestyle to the American alligator or Nile crocodile.

Hulkepholis

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 (including "Dollo's goniopholidid").

Mesosuchia

"Mesosuchia" is an obsolete name for a group of terrestrial, semi-aquatic, or fully aquatic crocodylomorph reptiles. The marine crocodile Metriorhynchus had paddle-like forelimbs, Dakosaurus andiniensis had a skull that was adapted to eat large sea reptiles, and Shamosuchus was adapted for eating molluscs and gastropods. Shamosuchus also looked like modern crocodiles and was very closely related to their direct ancestor.

The "Mesosuchia" were formerly placed at Suborder rank as within Crocodylia. The "first" crocodiles were placed within their own suborder, Protosuchia; whilst extant species where placed within Suborder Eusuchia (meaning 'true crocodiles'). Mesosuchia were the crocodylians "in between". As it is a paraphyletic group however, it is not considered valid anymore. It is replaced by its phylogenetic equivalent Mesoeucrocodylia, which contains the taxa herein, the Crocodylia, and some allied forms more recently discovered.

The "Mesosuchia" was composed as:

Family Hsisosuchidae

Family Gobiosuchidae

Infraorder Notosuchia

Family Notosuchidae

Family Sebecidae

Family Baurusuchidae

Infraorder Neosuchia

Family Trematochampsidae

Family Peirosauridae

Genus Lomasuchus

Genus Montealtosuchus

Family Elosuchidae

Family Atoposauridae

Family Dyrosauridae

Family Pholidosauridae

Genus Sarcosuchus

Infraorder Thalattosuchia - Sea "Crocodiles"

Family Teleosauridae

Family Metriorhynchidae

Genus Dakosaurus

Family Goniopholididae

Family Paralligatoridae

Genus Shamosuchus

Nannosuchus

Nannosuchus (meaning "dwarf crocodile") is an extinct genus of goniopholidid mesoeucrocodylian from the Berriasian of England.

Neosuchia

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 (the Nile Crocodile) 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. Neosuchians first appear in the Early Jurassic with the earliest known goniopholid Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages.

Pholidosauridae

Pholidosauridae is an extinct family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe (Denmark, England, France, Germany, Spain and Sweden), Africa (Algeria, Niger, Mali, Morocco and Tunisia), North America (Canada and the United States) and South America (Brazil and Uruguay). The pholidosaurids first appeared in the fossil record during the Bathonian stage of the Middle Jurassic and became extinct during the Late Turonian stage of the Late Cretaceous.Sarcosuchus is one of the best known pholidosaurs. It is believed to have attained lengths of up to 12 metres (39 ft 4 in) and weighed up to 8 tonnes (7.9 long tons; 8.8 short tons). One genus, Suchosaurus, once thought to be a pholidosaur, has since been shown to be a spinosaurid theropod dinosaur.

Pliogonodon

Pliogonodon is an extinct genus of crocodylomorph. The type species, P. priscus, was named by Joseph Leidy in 1856. The holotype, known as USNM 7448, is a worn and broken tooth found from Phoebus Landing on the Cape Fear River in North Carolina. Although the age of the strata in which the tooth was found was not recorded, it is thought to have come from Miocene-age beds. The holotype and another tooth found at the same location are all that is known from the genus. The genus is considered a nomen dubium because of the lack of diagnostic features possessed by the teeth, and has been suggested to be synonymous with the alligatoroid Deinosuchus. Although Carroll (1988) assigned the genus to the basal neosuchian family Goniopholididae, current consensus is that Pligonodon is a Deinosuchus specimen.The two teeth are conical and curve slightly inward, estimated to be around 2 inches (5.1 cm) in length if they had been fully preserved. The enamel is wrinkled and the base of the crowns are hollow.

Siamosuchus

Siamosuchus is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the pre-Aptian-age Lower Cretaceous Sao Khua Formation of eastern Thailand. It is known from a partial skull, most of the right half of the postcranial skeleton, and some bony scutes. Siamosuchus was described by Lauprasert and colleagues in 2007. The type species is S. phuphokensis. Siamosuchus may be closely related to the European genus Goniopholis.

Sunosuchus

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.

Symptosuchus

Symptosuchus is an extinct genus of goniopholidid mesoeucrocodylian. It is known from the Late Cretaceous of Argentina. Argentine paleontologist Florentino Ameghino named the genus in 1899, along with the type species S. contortidens. It was formally described by Carlos Rusconi in 1934.

Woodbinesuchus

Woodbinesuchus (meaning "Woodbine Formation crocodile") is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Cenomanian-age Upper Cretaceous Woodbine Formation of Texas. Crocodyliform fossils are widespread in the formation. The type specimen of Woodbinesuchus, SMU 74626, came from near the base of the formation in Tarrant County. It was found by Johnny Allen Byers (Ted & Mary's favorite boy) and J. Maurice in May 1990. The specimen includes the lower jaw and a variety of postcranial elements, such as vertebrae, limb bones, shoulder and hip bones, and bony armor. The lower jaw was elongate and slender, with the two halves joined from the tip to the sixteenth teeth, and there was no mandibular fenestra. Yuong–Nam Lee named Woodbinesuchus in 1997, naming the type species W. byersmauricei to honor the discoverers. He tentatively assigned Woodbinesuchus to the Goniopholidae.

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