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 PreꞒ Ꞓ O S D C P T J K Pg N
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

G. crassidens holotype BMNH 3798

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

G. simus skull from Middle Purbeck Group

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. 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–S108. doi:10.1111/j.1096-3642.2011.00709.x.
2. 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. PMC 6070184. PMID 30067779.
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. hdl:2246/6449.
9. Steven W. Salisbury; Darren Naish (2011). "Crocodilians". In Batten, D. J. (ed.). 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. 36 (6): e1222534. 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 (1–4): 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.