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Amphicotylus Temporal range: Late Jurassic, Tithonian PreꞒ Ꞓ O S D C P T J K Pg N ↓
Skull (OMNH 2392) of Amphicotylus stovalli from Oklahoma
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauria
Clade:	Pseudosuchia
Clade:	Crocodylomorpha
Clade:	Crocodyliformes
Family:	†Goniopholididae
Genus:	†Amphicotylus Cope, 1878
Type species
†Amphicotylus lucasii Cope, 1878
Species
†A. felix (Marsh, 1877) †A. gilmorei? (Holland, 1905) †A. lucasii Cope, 1878 †A. stovalli (Mook, 1964) †A. milesi Yoshida et al., 2021

Amphicotylus is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was described in 1878.

Discovery and species

Amphicotylus was first described by Edward Drinker Cope in 1878 based on dorsal and lumbar vertebrae, ribs and osteoderms. Based on these remains, found in the same locality as Camarasaurus supremus, Cope determined the animal to have been smaller than the extant American Alligator and named the species A. lucasii after Superintendent Lucas who initially made the discovery. Cope also collected skull material from the same locality and level, however did not refer it to Amphicotylus. The cranial material was examined by Charles C. Mook in 1942 who referred it to the type species not only on the basis of its locality, but also its matching size, general morphological characters and the absence of any other crocodilian remains from the area. Mook furthermore uses this skull to establish a provisional neotype to account for the fragmentary nature of the original material described by Cope.

In the 1930s the Works Progress Administration discovered further remains in Oklahoma which were initially investigated by J. W. Stovall, noting similarities to Amphicotylus gilmorei. The remains were officially described by Mook in 1964 as a species of Goniopholis, Goniopholis stovalli. In 2012 Eric Randall Allen argued that the goniopholidids found in the Morrison Formation differ significantly from their European counterparts in the anatomy of the palate, with the choanae fully splitting the palatines unlike in the taxa from the UK and mainland Europe. Consequently, Allen reasons that Goniopholis should exclusively refer to European forms while Amphicotylus should serve as a constituent for Morrison goniopholidids previously referred to Goniopholis, such as "Goniopholis" felix. Furthermore, it is noted that there are no significant differences between A. lucasii and the only known material of A. gilmorei, with the only notable morphological character being artificially created by breakage, rendering it a junior synonym of the genus' type species.

Excavations in the East Camarasaurus Quarry, Wyoming, in 1993 yielded the largest and most complete goniopholidid skeleton found to date. Since 1996 the fossil has been exhibited and stored in the Gunma Museum of Natural History in Japan before it was described by Yoshida et al. in 2021 as Amphicotylus milesi, in honor of Clifford Miles for his contribution to the excavation and museum curation.

Description

Amphicotylus had a roughly triangular, platyrostral skull similar in shape to that of modern crocodilians. They differ from European goniopholidids in several morphological characters, most notably the anatomy of the palate. In members of Amphicotylus, the palatines are completely separated from one another by the choana while in Goniopholis the palatines make broad contact prior to the choana. This trait is shared by Eutretauranosuchus. Amphicotylus milesi is the largest species with a skull length of 43 cm.

The anatomy of the palate and hyoid in A. milesi show similar adaptations to extant crocodilians, with the anterior portion of the elongated choana possibly being covered in soft tissue similar to the state observed in embryos of American alligators. Based on the morphology of the hyoid apparatus and its muscle attachments, A. milesi would have been able to raise the ventral fold of the gullar valve, separating narial and oral passages. This adaptation for semi-aquatic life would have allowed Amphicotylus and other Neosuchians to completely cut out the oral cavity from respiration, giving it the ability to open its mouth underwater while continuing to breathe as long as the nares are above the surface. This pushes the development of a crocodilian-like respiratory system to the beginnings of Neosuchian evolution in the Jurassic period.

Phylogeny

Studies have repeatedly found Amphicotylus to be nested deep within Goniopholididae, typically located just outside the clade that contains most European members of the group. Denazinosuchus, a late Cretaceous member of Goniopholididae, was recovered as a sister taxon to Amphicotylus by Puértolas-Pascual et al. 2015, but as part of the "European clade" by Yoshida et al. 2021. Yoshida et al. also include Sunosuchus thailandicus without following the revisions by Martin et al. 2013 that found it to be a distinct genus of pholidosaurid, Chalawan thailandicus. The phylogenetic tree recovered by Yoshida et al. is depicted below.

References

1. ^ Yoshida, J.; Hori, A.; Kobayashi, Y.; Ryan, M.J.; Takakuwa, Y.; Hasegawa, Y. (8 December 2021). "A new goniopholidid from the Upper Jurassic Morrison Formation, USA: novel insight into aquatic adaptation toward modern crocodylians". The Royal Society. 8 (12). doi:10.1098/rsos.210320. PMC 8652276. PMID 34909210.
2. Pritchard, A. C.; Turner, A. H.; Allen, E. R. & Norell, M. A. (2013). "Osteology of a North American Goniopholidid (Eutretauranosuchus delfsi) and Palate Evolution in Neosuchia". American Museum Novitates (3783): 1–56. doi:10.1206/3783.2. hdl:2246/6449. S2CID 73539708.
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. ^ Puértolas-Pascual, E.; Canudo, J.I. & Sender, L.M. (2015). "New material from a huge specimen of Anteophthalmosuchus cf. escuchae (Goniopholididae) from the Albian of Andorra (Teruel, Spain): Phylogenetic implications". Journal of Iberian Geology. 41 (1): 41–56. doi:10.5209/rev_JIGE.2015.v41.n1.48654.
5. ^ E. D. Cope. 1878. Descriptions of new extinct Vertebrata from the Upper Tertiary and Dakota Formations. 'Bulletin of the United States Geological and Geographical Survey of the Territories, 4(2):379-396
6. Mook, C.C. (2 April 1942). "Skull Characters of Amphicotylus lucasii Cope". American Museum Novitates (1165).
7. ^ Allen, E.R. (2012). "Analysis of North American goniopholidid crocodyliforms in a phylogenetic context". M.C. thesis, University of Iowa.
8. "水中適応では最古...ワニ祖先の新種　福島県立博物館のグループ" [The oldest in water adaptation: A group of new species of crocodile ancestors at the fukushima prefectural museum]. 9 December 2021. Retrieved 9 December 2021.

v t e Neosuchia
Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Pseudosuchia Clade: Metasuchia Pseudosuchia see Pseudosuchia Neosuchia see below↓
Neosuchia Neosuchia †Burkesuchus †Isisfordia †Karatausuchus †Khoratosuchus †Montsecosuchus †Stolokrosuchus? †Susisuchus †Tadzhikosuchus? †Wahasuchus? †Zhyrasuchus? †Thalattosuchia? †Tethysuchia Pholidosauridae Anglosuchus Chalawan Crocodilaemus Dakotasuchus Elosuchus Meridiosaurus Oceanosuchus Pholidosaurus Sarcosuchus Terminonaris Vectisuchus Woodbinesuchus Dyrosauridae Acherontisuchus Anthracosuchus Arambourgisuchus Atlantosuchus Brachiosuchus Cerrejonisuchus Chenanisuchus Congosaurus Dorbignysuchus Dyrosaurus Fortignathus Guarinisuchus Hyposaurus Luciasuchus Phosphatosaurus Rhabdognathus Rodeosuchus Sabinosuchus? Sokotosuchus Tilemsisuchus †Atoposauridae Alligatorellus Alligatorium Aprosuchus Atoposaurus Knoetschkesuchus Sabresuchus Theriosuchus Varanosuchus †Stomatosuchidae Laganosuchus Stomatosuchus †Paluxysuchidae Deltasuchus Paluxysuchus †Goniopholididae Amphicotylus Anteophthalmosuchus Calsoyasuchus Coelosuchus Denazinosuchus Diplosaurus Eutretauranosuchus Goniopholis Hulkepholis Nannosuchus Ophiussasuchus Oweniasuchus Pinacosuchus? Siamosuchus Sunosuchus Symptosuchus? †Bernissartiidae Bernissartia Koumpiodontosuchus †Paralligatoridae Batrachomimus Brillanceausuchus Kansajsuchus Paralligator Rugosuchus Scolomastax Shamosuchus Tarsomordeo Turanosuchus Wannchampsus Yanjisuchus Eusuchia see below↓
Eusuchia Eusuchia †Borealosuchus †Confractosuchus †Dolichochampsa †Gilchristosuchus †Heterosuchus †Isisfordia? †Portugalosuchus †Pristichampsus †Susisuchus? †Titanochampsa? †Atoposauridae? †Bernissartiidae? †Paralligatoridae? †Hylaeochampsidae Acynodon? Hylaeochampsa Iharkutosuchus Pachycheilosuchus? Pietraroiasuchus Turcosuchus Unasuchus †Allodaposuchidae Agaresuchus Allodaposuchus Arenysuchus? Ischyrochampsa Lohuecosuchus Massaliasuchus Musturzabalsuchus? †Aegyptosuchidae Aegyptosuchus Aegisuchus †"Thoracosaurs" Argochampsa Eogavialis Eosuchus Eothoracosaurus Thoracosaurus †Planocraniidae Boverisuchus Duerosuchus Planocrania Crocodilia see Crocodilia for extant taxa, and Extinct Crocodilia for extinct ones
See also: Coelognathosuchia Category

• Goniopholidids
• Late Jurassic crocodylomorphs of North America
• Late Jurassic reptiles of North America
• Morrison fauna
• Fossil taxa described in 1878
• Taxa named by Edward Drinker Cope
• Prehistoric pseudosuchian genera