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Hulkepholis Temporal range: Early Cretaceous, 135–112 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauria
Clade:	Pseudosuchia
Clade:	Crocodylomorpha
Clade:	Crocodyliformes
Family:	†Goniopholididae
Genus:	†Hulkepholis Buscalioni et al., 2013
Type species
†Goniopholis willetti Salisbury & Naish, 2011
Species
†H. plotos Buscalioni et al., 2013 †H. rori Arribas et al., 2019 †H. willetti Salisbury & Naish, 2011

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").

Discovery

H. willetti is known from a single holotype specimen, BMNHB 001876 from the Wealden Group of the Isle of Wight that includes a well-preserved nearly complete skull. It was collected by Edgar W. Willett at Cuckfield, West Sussex, from the Valanginian-aged Grinstead Clay Member, of the Hastings Group, Wealden Supergroup. Willett showed the specimen to the Geological Society of London in or around 1877 or 1878, and it was then Hulke (1878) described it as a specimen of Goniopholis crassidens, an identification derived from a similar tooth form. The specimen was considered lost among the crocodyliform research community but was actually safely accessioned at the Booth Museum in Brighton. It was identified as "the long-lost crocodilian of Mr Willett" by Steve (1998), "Willett’s specimen" by Salisbury et al. (1999) and "Hulke’s specimen" by Andrande et al. (2011). It was assigned to a new species of Goniopholis, Goniopholis willetti, by Salisbury and Naish in 2011 honoring its collector. A broad phylogenetic analysis of crocodyliforms published the same year by Andrande et al. (2011), found the specimen to be the sister taxon of the clade formed by Anteophthalmosuchus hooleyi and the unnamed "Dollo’s goniopholidid". Thus it was reassigned to its own genus by Buscalioni et al. (2013), creating the combinatio nova, Hulkepholis willetti. The generic name honors John Whitaker Hulke who described the specimen as belonging to Goniopholis.

A second species, Hulkepholis plotos, was first described and named by A.D. Buscalioni, L. Alcalá, E. Espílez and L. Mampel in 2013. The specific name is derived from Greek mythology πλοτός, plotos, the drifter. It is known solely from the holotype AR-1/56, a partial skeleton which consists of AR-1-2045, a nearly complete but crushed skull; AR-1-2048, 4859, 4860, three vertebrae; AR-1-2046, a rib; AR-1-2048, a metapodial; and AR-1-2049, 4861, 4862, three osteoderms. It was collected from the early Albian-aged Escucha Formation, at Santa Maria Mine located in the municipality of Ariño, Teruel Province, of Aragon, along with the closely related Anteophthalmosuchus escuchae.

Phylogeny

Goniopholis willetti was included in a phylogenetic analysis of goniopholidids that was published soon before the specimen was redescribed. It was found to be most closely related to the clade formed by a specimen called "Dollo's goniopholidid", and "Hooley’s goniopholidid", now named Anteophthalmosuchus hooleyi. Below is a cladogram from that analysis:

References

1. ^ Steven W. Salisbury; Darren Naish (2011). "Crocodilians". In Batten, D. J. (ed.). English Wealden Fossils. The Palaeontological Association (London). pp. 305–369.
2. Naish, Darren (2 December 2011). "The Wealden Bible: English Wealden Fossils, 2011". Tetrapod Zoology. Scientific American Blogs. Retrieved 5 December 2011.
3. ^ 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. doi:10.7203/sjp.28.1.17835. S2CID 73570030.
4. 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. hdl:2318/1635521. S2CID 89199731.
5. Naish, Darren (24 September 2012). "In pursuit of Early Cretaceous crocodyliforms in southern England: ode to Goniopholididae". Tetrapod Zoology. Scientific American Blogs. Retrieved 24 September 2012.
6. ^ 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.

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
• Early Cretaceous crocodylomorphs of Europe
• Fossil taxa described in 2011
• Fossil taxa described in 2013
• Early Cretaceous reptiles of Europe
• Fossils of Great Britain
• Fossils of Spain
• Prehistoric pseudosuchian genera