Hulkepholis

Last updated

Hulkepholis
Temporal range: Early Cretaceous, 135–112  Ma
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Pseudosuchia
Superorder: Crocodylomorpha
Family: Goniopholididae
Genus: Hulkepholis
Buscalioni et al., 2013
Type species
Goniopholis willetti
Salisbury & Naish, 2011
Species
  • H. plotosBuscalioni et al., 2013
  • H. roriArribas et al., 2019
  • H. willettiSalisbury & 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"). [1] [2] [3] [4]

Contents

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. [3] 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). [5] It was assigned to a new species of Goniopholis , Goniopholis willetti, by Salisbury and Naish in 2011 honoring its collector. [1] 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". [6] 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. [3]

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 . [3]

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: [6]

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

Related Research Articles

<i>Goniopholis</i> Extinct genus of reptiles

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

<i>Suchosaurus</i> Extinct genus of dinosaurs

Suchosaurus is a spinosaurid dinosaur from Cretaceous England and Portugal, originally believed to be a genus of crocodile. The type material, consisting of teeth, was used by British palaeontologist Richard Owen to name the species S. cultridens in 1841. Later in 1897, French palaeontologist Henri-Émile Sauvage named a second species, S. girardi, based on two fragments from the mandible and one tooth discovered in Portugal. Suchosaurus is possibly a senior synonym of the contemporary spinosaurid Baryonyx, but is usually considered a dubious name due to the paucity of its remains, and is considered an indeterminate baryonychine. In the Wadhurst Clay Formation of what is now southern England, Suchosaurus lived alongside other dinosaurs, as well as plesiosaurs, mammals, and crocodyliforms.

Calsoyasuchus is a genus of crocodylomorph 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 specific name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones. It has often been interpreted as the earliest diverging member of Goniopholididae, but other studies have recovered it in various other positions.

Rugosuchus is an extinct genus of neosuchian crocodyliform from the late Early Cretaceous of China. It is known from most of a skull, a partial postcranial skeleton, and a second partial skeleton including part of the hips. It was described by Xiao-Chun Wu and colleagues in 2001, with R. nonganensis as the type species. At the time of its description, it was the most complete crocodyliform from northeastern China, and only the second known.

<span class="mw-page-title-main">Goniopholididae</span> Extinct family of reptiles

Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia between the Middle Jurassic and the Late Cretaceous.

<span class="mw-page-title-main">Escucha Formation</span>

The Escucha Formation is a geological formation in La Rioja and Teruel provinces of northeastern Spain whose strata date back to the late Aptian to middle Albian stages of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.

<i>Dakotasuchus</i> Extinct genus of reptiles

Dakotasuchus is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Cenomanian-age Upper Cretaceous Dakota Sandstone of Kansas. The type specimen was found in an iron-cemented sandstone concretion near Salina. This concretion was broken into two large pieces; more of the specimen was probably present originally, but by the time it was found only the torso and short portions of the neck and tail remained. Twenty pairs of bony scutes ran down the midline of the back. The vertebrae lacked the procoelous articulation of more derived crocodyliforms. Dakotasuchus had short broad shoulder blades, suggesting it had stout powerful forelimbs and perhaps terrestrial habits. M. G. Mehl, who described the genus, estimated the length of the type individual when complete to have been 3–4 metres (9.8–13.1 ft). The type species is D. kingi, named for Professor King, a former dean of Kansas Wesleyan University. Mehl did not classify his new genus to a more inclusive group than Mesosuchia. Robert Carroll assigned Dakotasuchus to Goniopholididae in 1988. In 2017, fossils of Dakotasuchus kingi which consisted of a coracoid, scutes, a dorsal vertebrate and postcranial bones were found in Utah, specifically in the Cedar Mountain Formation's Mussentuchit Member.

Eutretauranosuchus is an extinct genus of goniopholidid crocodyliform. E. delfsi is the only known species within the genus.

<span class="mw-page-title-main">Neosuchia</span> Clade of reptiles

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 than to Notosuchus terrestris. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages in North America. It is often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.

<i>Isisfordia</i> Extinct genus of reptiles

Isisfordia is an extinct genus of crocodyliform closely related to crocodilians that lived in Australia during the Middle Cretaceous (Albian–Cenomanian).

<i>Amphicotylus</i> Extinct genus of reptiles

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

<i>Pholidosaurus</i> Extinct genus of reptiles

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-Albian stages 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.

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.

<i>Susisuchus</i> Extinct genus of reptiles

Susisuchus is an extinct genus of neosuchian mesoeucrocodylian crocodyliform from the Early Cretaceous of Brazil. Fossils have been found from the Nova Olinda Member of the Aptian-age Crato Formation in the Araripe and Lima Campos Basins of northeastern Brazil. Named in 2003, Susisuchus is the sole member of the family Susisuchidae, and is closely related to the clade Eusuchia, which includes living crocodilians. The type species is S. anatoceps, known from a single partial articulated skeleton that preserves some soft tissue. A second species, S. jaguaribensis, was named in 2009 from fragmentary remains.

Anteophthalmosuchus is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England, eastern Spain, and western Belgium.

<i>Proa valdearinnoensis</i> Extinct species of reptile

Proa is a genus of basal styracosternan iguanodont known from the Early Cretaceous Escucha Formation of Teruel Province, Spain.

Paluxysuchus is an extinct genus of neosuchian crocodyliform known from the Early Cretaceous Twin Mountains Formation of north-central Texas. It contains a single species, Paluxysuchus newmani. Paluxysuchus is one of three crocodyliforms known from the Early Cretaceous of Texas, the others being Pachycheilosuchus and an unnamed species referred to as the "Glen Rose Form". Paluxysuchus has a long, flat skull that is probably transitional between the long and narrow skulls of many early neosuchians and the short and flat skulls of later neosuchians.

<i>Europelta</i> Extinct genus of dinosaurs

Europelta is a monospecific genus of nodosaurid dinosaur from Spain that lived during the Early Cretaceous in what is now the lower Escucha Formation of the Teruel Province. The type and only species, Europelta carbonensis, is known from two associated partial skeletons, and represents the most complete ankylosaur known from Europe. Europelta was named in 2013 by James I. Kirkland and colleagues. Europelta has an estimated length of 5 metres and weight of 1.3 tonnes, making it the largest member of the clade Struthiosaurini.

<span class="mw-page-title-main">Coelognathosuchia</span> Extinct clade of reptiles

Coelognathosuchia is an extinct clade of neosuchian crocodyliforms that includes all taxa more closely related to the family Pholidosauridae than to Bernissartia fagesii or Eusuchia. Martin et al. (2014) named the clade after finding goniopholidids and pholidosaurids to group together in their phylogenetic analysis of crocodyliform evolutionary relationships. In their analysis, Pholidosauridae was monophyletic and Goniopholididae was paraphyletic, being an assemblage of successively more basal taxa within Coelognathosuchia. Coelognathosuchia itself was positioned near the base of the larger clade Neosuchia as the sister group to a clade containing the Early Cretaceous neosuchian Bernissartia and Eusuchia, the group that includes all modern crocodilians and their closest extinct relatives.

The Sainte-Barbe Clays Formation is a geological formation in Belgium. It is found in localised areas of the northern margin of the Mons Basin, alongside the equivalently aged Hautrage and Baudour Clay Formations. It is Upper Barremian-Lower Aptian in age. It predominantly consists of laminated clay, with some lignite. It is well known for the "Iguanodon sinkhole" locality near Bernissart where many specimens of Iguanodon bernissartensis were described by Louis Dollo in the late 19th century.

References

  1. 1 2 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. 1 2 3 4 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. 1 2 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 .