Goniopholis

Goniopholis is an extinct genus of goniopholidid crocodyliform that lived in Europe and Africa during the Late Jurassic and Early Cretaceous.[1][2] 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.

Goniopholis
Temporal range: Late Jurassic - Early Cretaceous, 155–139.8 Ma
Swanage Crocodile Goniopholis kiplingi
Skull of the "Swanage Crocodile", G. kiplingi. Berriasian age (earliest Cretaceous)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Family: Goniopholididae
Genus: Goniopholis
Owen, 1841
Type species
Goniopholis crassidens
Owen, 1841
Species
  • G. baryglyphaeus Schwarz, 2002
  • G. crassidens Owen, 1841
  • G. kiplingi Andrade et al., 2011
  • G. simus Owen, 1878

Discovery and species

Goniopholis crassidens
G. crassidens holotype BMNH 3798
Goniopholis BW
G. simus restoration

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".[3]

Eggs attributed to Goniopholis were found in the Late Jurassic of Portugal.[4]

Formerly assigned species

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.[5]

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.[6][7][8]

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][9] Dollo's goniopholidid has also been assigned to Anteophthalmosuchus.[10]

Classification

Goniopholis simus skull
G. simus skull from Middle Purbeck Group
Goniopholis - Castilla-La Mancha Paleontological Museum (Cuenca, Spain)
Model of Goniopholis at the Castilla-La Mancha Paleontological Museum

Below is a cladogram including several Goniopholis species:[1]

Neosuchia
Atoposauridae

Theriosuchus pusillus

Theriosuchus guimarotae

Rugosuchus

Bernissartia

Eusuchia

Stolokrosuchus

Tethysuchia

Thalattosuchia

Goniopholididae

Calsoyasuchus valliceps

"Goniopholis" phuwiangensis

Eutretauranosuchus delfi

"Sunosuchus" junggarensis

Sunosuchus miaoi

Sunosuchus thailandicus

Siamosuchus phuphokensis

Amphicotylus lucasii

Denazinosuchus kirtlandicus

Nannosuchus gracilidens

Hulkepholis (Hulke's goniopholidid)

Anteophthalmosuchus (Hooley’s goniopholidid)

Anteophthalmosuchus (Dollo’s goniopholidid)

Goniopholis

Goniopholis baryglyphaeus

Goniopholis kiplingi

Goniopholis simus

References

  1. ^ a b c d e f g De Andrade, M. B.; Edmonds, R.; Benton, M. J.; Schouten, R. (2011). "A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus". Zoological Journal of the Linnean Society. 163: S66. doi:10.1111/j.1096-3642.2011.00709.x.
  2. ^ a b c Buscalioni, A.D.; Alcalá, L.; Espílez, E.; Mampel, L. (2013). "European Goniopholididae from the Early Albian Escucha Formation in Ariño (Teruel, Aragón, España)". Spanish Journal of Palaeontology. 28 (1): 103–122.
  3. ^ "BBC News - Rudyard Kipling inspires naming of prehistoric crocodile". BBC Online. 2011-03-20. Retrieved 20 March 2012.
  4. ^ Russo, J., Mateus O., Marzola M., & Balbino A. (2017). Two new ootaxa from the late Jurassic: The oldest record of crocodylomorph eggs, from the Lourinhã Formation, Portugal. PLOS ONE. 12, 1-23.
  5. ^ Piacentini Pinheiro, A.E.; da Costa Pereira, P.V.L.G.; de Souza, R.G.; Brum, A.S.; Lopes, R.T.; Machado, A.S.; Bergqvist, L.P.; Simbras, F.M. (2018). "Reassessment of the enigmatic crocodyliform "Goniopholis" paulistanus Roxo, 1936: Historical approach, systematic, and description by new materials". PLoS ONE. 13 (8): e0199984. doi:10.1371/journal.pone.0199984.
  6. ^ Allen, E. (2010). "Phylogenetic analysis of goniopholidid crocodyliforms of the Morrison Formation". Journal of Vertebrate Paleontology. 30 (Supp. 1): 52A. doi:10.1080/02724634.2010.10411819.
  7. ^ Pol, D.; Leardi, J.M.; Lecuona, A.; Krause, M. (2012). "Postcranial anatomy of Sebecus icaeorhinus (Crocodyliformes, Sebecidae) from the Eocene of Patagonia". Journal of Vertebrate Paleontology. 32 (2): 328. doi:10.1080/02724634.2012.646833.
  8. ^ 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 (3783): 1. doi:10.1206/3783.2.
  9. ^ Steven W. Salisbury; Darren Naish (2011). "Crocodilians". In Batten, D. J. English Wealden Fossils. The Palaeontological Association (London). pp. 305–369.
  10. ^ Martin, J.E.; Delfino, M.; Smith, T. (2016). "Osteology and affinities of Dollo's goniopholidid (Mesoeucrocodylia) from the Early Cretaceous of Bernissart, Belgium". Journal of Vertebrate Paleontology. doi:10.1080/02724634.2016.1222534.

Sources

  • Buffetaut, E; Ingavat, R (1983). "Goniopholis phuwiangensis nov. sp., a new mesosuchian crocodile from the Mesozoic of North-eastern Thailand". Geobios. 16 (1): 79–91. doi:10.1016/S0016-6995(83)80048-5.
  • Holland, W. J. (1905). "A new crocodile from the Jurassic of Wyoming". Annals of the Carnegie Museum. 3 (3): 431–434. ISSN 0097-4463.
  • Mook, C. C. (1964). "New species of Goniopholis from the Morrison of Oklahoma". Oklahoma Geology Notes. 24: 283–287. ISSN 0030-1736.
  • Owen, R. 1878. Monograph on The Fossil Reptilia of the Wealden and Purbeck Formations, Supplement no. VII. Crocodilia (Goniopholis, Pterosuchus, and Suchosaurus). Palaeontological Society Monograph, p. 1-15.
  • Owen, R. (1879). "On the Association of dwarf crocodiles (Nanosuchus and Theriosuchus pusilus, e. g.) with the diminutive mammals of the Purbeck Shales". Quarterly Journal of the Geological Society of London. 35: 148–155. doi:10.1144/GSL.JGS.1879.035.01-04.02.
  • Salisbury, S. W.; Willis, P. M. A.; Peitz, S. & Sander, P. M. (December 1999). "The crocodilian Goniopholis simus from the Lower Cretaceous of north-western Germany". Special Papers in Palaeontology. 60: 121–148. ISBN 978-0-901702-67-8.
  • Schwarz, Daniela (2002). "A new species of Goniopholis from the Upper Jurassic of Portugal". Palaeontology. 45 (1): 185–208. doi:10.1111/1475-4983.00233.
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.

Blackhorse Quarry

Blackhorse Quarry is a Site of Special Scientific Interest in East Sussex, England and is of geological interest.The site has a great value of vertebrate palaeontological interest, it is the type locality for the Wadhurst Clay Telham Bond Bed. The

bone bed is a conglomerate of polished pebbles (indicating a former high energy surface), fish scales, coprolites, bones, teeth

and scutes. The reptiles recorded include turtles, crocodiles (Goniopholis), pterosaurs

(Ornithocheirus), and dinosaurs (Megalosaurus, Cetiosaurus, Iguanodon

Hylaeosaurus). This site has yielded large numbers of specimens in the past, and it

has excellent potential for further finds.

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.

Como Bluff

Como Bluff is a long ridge extending east-west, located between the towns of Rock River and Medicine Bow, Wyoming. The ridge is an anticline, formed as a result of compressional geological folding. Three geological formations, the Sundance, the Morrison, and the Cloverly Formations, containing fossil remains from the Late Jurassic of the Mesozoic Era are exposed. Nineteenth century paleontologists discovered many well-preserved specimens of dinosaurs, as well as mammals, turtles, crocodilians, and fish from the Morrison Formation. Because of this, Como Bluff is considered to be one of the major sites for the early discovery of dinosaur remains. Among the species discovered is the only known specimen of Coelurus. Significant discoveries were made in 22 different areas scattered along the entire length of the ridge. It is included on the National Register of Historic Places as well as the National Natural Landmark list.

Denazinosuchus

Denazinosuchus is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Upper Cretaceous Fruitland Formation and Kirtland Formation (late Campanian-early Maastrichtian) of the San Juan Basin, New Mexico. It is the most abundant and readily identifiable mesoeucrocodylian of the San Juan Basin, mostly due to its distinctive subrectangular, flattened, and sparsely pitted bony armor. It was first described in 1932 by Carl Wiman on the basis of a skull as a species of Goniopholis, G. kirtlandicus. Spencer G. Lucas and Robert M. Sullivan redescribed the species in 2003 and gave it its own genus, Denazinosuchus. To date, Denazinosuchus is only known from skull material, armor, and a thigh bone.

Diplosaurus

Diplosaurus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils have been found from the western United States and range from Late Jurassic to Early Cretaceous in age. The genus was first named and described in a paper written in 1877 by Othniel Charles Marsh. The generic name, derived from Greek διπλόος, diploos, "double", probably refers to the "biconcave vertebrae" Marsh mentions as a distinctive trait compared to modern forms. The type species is Diplosaurus felix. In 1890 Karl Alfred von Zittel recombined this with Goniopholis into a Goniopholis felix, but today this is generally rejected.

Along with D. felix (sometimes misspelled D. felise), a second species of Diplosaurus was constructed in 1877 by Marsh after he recombined Hyposaurus vebbii, named by Edward Drinker Cope in 1874, with Diplosaurus into a new combination, Diplosaurus vebbii. D. nanus was named by Marsh in 1895 but is now considered to be a nomen dubium.

Eutretauranosuchus

Eutretauranosuchus is a genus of goniopholidid mesoeucrocodylian. It is known from several specimens collected from the Late Jurassic-age Morrison Formation, including fossils from Garden Park and Dry Mesa Quarry in Colorado and Como Bluff in Wyoming. The type species is E. delfsi. Charles Mook described Eutretauranosuchus in 1967 from a skull and partial skeleton of a subadult individual uncovered at Garden Park in the 1950s by workers from the Cleveland Museum of Natural History. This individual was only about 1.77 metres (5.8 ft) long, and weighed perhaps 18 kilograms (40 lb), but older individuals could reach substantially larger sizes. The skull of Eutretauranosuchus differed from that of contemporaneous Goniopholis by the presence of an additional opening in the palate, among other details. Eutretauranosuchus would have been a semiaquatic predator, its prey including fish and small to medium-sized terrestrial vertebrates.

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.

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").

Itasuchidae

Itasuchidae is a Gondwanan family of mesoeucrocodylians that lived during the Cretaceous period. It was a clade of terrestrial crocodyliforms that evolved a rather dog-like form, and were terrestrial carnivores.

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.

Pholidosaurus

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 stage 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.

Roxochampsa

Roxochampsa is an extinct genus of crocodylomorph from the Late Cretaceous of Brazil belonging to the sebecosuchian clade Itasuchidae.

Sarcosuchus

Sarcosuchus (; meaning "flesh crocodile") is an extinct genus of crocodyliform and distant relative of living crocodylians that lived 112 million years ago. It dates from the early Cretaceous Period of what is now Africa and South America and is one of the largest crocodile-like reptiles that ever lived. It was almost twice as long as the modern saltwater crocodile and weighed up to 8 tonnes (7.9 long tons; 8.8 short tons).

The first remains were discovered during several expeditions led by the French paleontologist Albert-Félix de Lapparent, spanning from 1946 to 1959, in the Sahara. These remains were fragments of the skull, vertebrae, teeth, and scutes. In 1964, an almost complete skull was found in Niger by the French CEA, but it was not until 1997 and 2000 that most of its anatomy became known to science, when an expedition led by the American paleontologist Paul Sereno discovered six new specimens, including one with about half the skeleton intact and most of the spine.

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.

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