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").[1][2][3][4]

Hulkepholis
Temporal range: Early Cretaceous, 135–112 Ma
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Family: Goniopholididae
Genus: Hulkepholis
Buscalioni et al., 2013
Type species
Goniopholis willetti
Salisbury & Naish, 2011
Species
  • H. plotos Buscalioni et al., 2013
  • H. willetti Salisbury & Naish, 2011

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

References

  1. ^ a b 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. ^ a b c d 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.
  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.
  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. ^ a b 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.
Aegyptosuchidae

Aegyptosuchidae is an extinct family of eusuchian crocodyliforms from the Cretaceous period of Africa. They are characterized by their large size and flat heads. The family includes two genera, Aegyptosuchus and Aegisuchus.

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.

Anthracosuchus

Anthracosuchus (meaning "coal crocodile" in Greek) is an extinct genus of dyrosaurid crocodyliform from the Paleocene of Colombia. Remains of Anthracosuchus balrogus, the only known species, come from the Cerrejón Formation in the Cerrejón mine, and include four fossil specimens with partial skulls. Anthracosuchus differs from other dyrosaurids in having an extremely short (brevirostrine) snout, widely spaced eye sockets with bony protuberances around them, and osteoderms that are smooth and thick. It is one of the most basal dyrosaurids along with Chenanisuchus and Cerrejonisuchus. The species name is a reference to the Balrog, a creature in J. R. R. Tolkien's fantasy novel The Lord of the Rings that could, like the remains of Anthracosuchus, be found in a mine.

Atoposauridae

Atoposauridae is a family of crocodile-line archosaurs belonging to Neosuchia. The majority of the family are known from Late Jurassic to Early Cretaceous marine deposits in France, Portugal, and Bavaria in southern Germany. The discovery of the genus Aprosuchus, however, extends the duration of the lineage to the end of the Cretaceous in Romania.

Brillanceausuchus

Brillanceausuchus is an extinct genus of atoposaurid crocodylomorph. Fossils have been found in Early Cretaceous–age rocks of Cameroon. The genus is notable for the position of the secondary choana within its palate. Parts of the pterygoid bones make up the rostral margin of the choana and thus separate it from the palatines, a feature also seen in the more advanced neosuchian suborder Eusuchia. This characteristic was once thought to be characteristic of Eusuchia, but its presence in Brillanceausuchus suggests that the trait is homoplasic, thus making the evolution of the position of the choana within crocodilians more complex than previously thought.

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.

Coelosuchus

Coelosuchus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils have been found from the Graneros Shale of the Benton Group in Wyoming, and are of Cenomanian age. It was slightly over 1 meter in length.

Eusuchia

The Eusuchia ("true crocodiles") are a clade of crocodylomorphs that first appears in the Early Cretaceous with Hylaeochampsa. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-T extinction. Since the other two clades died out 35 and 11 million years ago, all living crocodilian species are eusuchians, as are many extinct forms.

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.

Goniopholis

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.

Karatausuchus

Karatausuchus is an extinct genus of atoposaurid crocodylomorph. It is known from a single specimen discovered in the Late Jurassic (Oxfordian - Kimmeridgian) Karabastau Svita from the vicinity of Mikhailovka in the Karatau Mountains of southern Kazakhstan. The type specimen is PIN 25858/1, a complete but poorly preserved juvenile skeleton with some possible soft tissue preservation. It is notable for having over 90 teeth, but its other anatomical details are difficult to discern. The length of this individual is estimated at 160 millimetres (6.3 in). Karatausuchus was described in 1976 by Mikhail Efimov, and the type species is K. sharovi.

Khoratosuchus

Khoratosuchus is an extinct genus of neosuchian crocodylomorph which existed in northeast Thailand during the early Cretaceous period. Its type species is Khoratosuchus jintasakuli. Khoratosuchus is the youngest and most advanced Mesozoic crocodyliform yet known from Thailand. It possesses several distinctive features that help determine its phylogenetic position among crocodylomorphs, including secondary choanae relatively posterior and almost encircled by the pterygoid bones on the palate and a smooth dorsal surface of the skull.

Laganosuchus

Laganosuchus is an extinct genus of stomatosuchid crocodyliform. Fossils have been found from Niger and Morocco and date back to the Upper Cretaceous.

Nannosuchus

Nannosuchus (meaning "dwarf crocodile") is an extinct genus of goniopholidid mesoeucrocodylian from the Berriasian of England.

Shamosuchus

Shamosuchus is an extinct genus of neosuchian crocodile that lived during the Late Cretaceous (Santonian-Campanian) period in what is now the Gobi desert of Mongolia, approximately 85 to 74 million years ago.

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.

Symptosuchus

Symptosuchus is an extinct genus of goniopholidid mesoeucrocodylian. It is known from the Late Cretaceous of Argentina. Argentine paleontologist Florentino Ameghino named the genus in 1899, along with the type species S. contortidens. It was formally described by Carlos Rusconi in 1934.

Wahasuchus

Wahasuchus is a genus of extinct mesoeucrocodylian of the Middle Campanian age found in the Quseir Formation, Egypt. The generic name derives from the Arabic word واحة (waha), which means "oasis", and souchos from the Greek in honor of crocodile-headed god of ancient Egypt. The specific egyptensis (Lat.) means from Egypt.Fossils of skull and jaw fragments, dorsal vertebrae, and fragmentary appendicular remains have been recovered.

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