Hulkepholis is an extinct genus of goniopholidid neosuchian 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:

NeosuchiaAtoposauridae 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
AtoposauridaeTheriosuchus pusillus Theriosuchus guimarotae
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
Rugosuchus Bernissartia Eusuchia
Rugosuchus
Bernissartia Eusuchia
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
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
Tethysuchia Thalattosuchia
Tethysuchia
Thalattosuchia
GoniopholididaeCalsoyasuchus 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
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
"Goniopholis" phuwiangensis Eutretauranosuchus delfi "Sunosuchus" junggarensis Sunosuchus miaoi Sunosuchus thailandicus
"Goniopholis" phuwiangensis
Eutretauranosuchus delfi "Sunosuchus" junggarensis
Eutretauranosuchus delfi
"Sunosuchus" junggarensis
Sunosuchus miaoi Sunosuchus thailandicus
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
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
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
Denazinosuchus kirtlandicus
Nannosuchus gracilidens
Hulkepholis (Hulke's goniopholidid) Anteophthalmosuchus (Hooley's goniopholidid) Anteophthalmosuchus (Dollo's goniopholidid) Goniopholis Goniopholis baryglyphaeus Goniopholis kiplingi Goniopholis simus
Hulkepholis (Hulke's goniopholidid) Anteophthalmosuchus (Hooley's goniopholidid) Anteophthalmosuchus (Dollo's goniopholidid)
Hulkepholis (Hulke's goniopholidid)
Anteophthalmosuchus (Hooley's goniopholidid) Anteophthalmosuchus (Dollo's goniopholidid)
Anteophthalmosuchus (Hooley's goniopholidid)
Anteophthalmosuchus (Dollo's goniopholidid)
GoniopholisGoniopholis baryglyphaeus Goniopholis kiplingi Goniopholis simus
Goniopholis baryglyphaeus
Goniopholis kiplingi Goniopholis simus
Goniopholis kiplingi
Goniopholis simus