''Hulkepholis'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

goniopholidid 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 (Asia, Europe and North America) between the Middle Ju ...

mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that M ...

n from the

Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma. Geology Pro ...

of southern

England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...

and eastern

Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...

. It contains two species, the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

, ''Hulkepholis willetti'', and also ''H. plotos''. ''Hulkepholis'' is most closely related to both species of ''

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. Discovery The holotype specimen of ''A ...

'' (including "Dollo's goniopholidid").

Discovery

''H. willetti'' is known from a single

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

specimen, BMNHB 001876 from the

Wealden Group The Wealden Group, occasionally also referred to as the Wealden Supergroup, is a group (a sequence of rock strata) in the lithostratigraphy of southern England. The Wealden group consists of paralic to continental (freshwater) facies sedimentary ...

of the

Isle of Wight The Isle of Wight ( ) is a county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the largest and second-most populous island of England. Referred to as 'The Island' by residents, the Isle of ...

that includes a well-preserved nearly complete

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...

. It was collected by Edgar W. Willett at Cuckfield,

West Sussex West Sussex is a county in South East England on the English Channel coast. The ceremonial county comprises the shire districts of Adur, Arun, Chichester, Horsham, and Mid Sussex, and the boroughs of Crawley and Worthing. Covering an ar ...

, from the

Valanginian In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretaceou ...

-aged Grinstead Clay Member, of the

Hastings Group The Wealden Group, occasionally also referred to as the Wealden Supergroup, is a group (a sequence of rock strata) in the lithostratigraphy of southern England. The Wealden group consists of paralic to continental (freshwater) facies sedimentary ...

, Wealden Supergroup. Willett showed the specimen to the

Geological Society of London The Geological Society of London, known commonly as the Geological Society, is a learned society based in the United Kingdom. It is the oldest national geological society in the world and the largest in Europe with more than 12,000 Fellows. Fe ...

in or around 1877 or 1878, and it was then Hulke (1878) described it as a specimen of ''

Goniopholis crassidens ''Goniopholis'' (meaning "angled scale") 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 ...

'', 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'' (meaning "angled scale") 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 ...

'', ''Goniopholis willetti'', by Salisbury and Naish in

2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrate ...

honoring its collector. A broad

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

analysis of crocodyliforms published the same year by Andrande ''et al.'' (2011), found the specimen to be the

sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

of the clade formed by ''

Anteophthalmosuchus hooleyi ''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. Discovery The holotype specimen of ''A ...

'' and the unnamed "Dollo’s goniopholidid". Thus it was reassigned to its own genus by Buscalioni ''et al.'' (2013), creating the ''

combinatio nova ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused wi ...

'', ''Hulkepholis willetti''. The generic name honors

John Whitaker Hulke John Whitaker Hulke FRCS FRS FGS (6 November 1830 – 19 February 1895) was a British surgeon, geologist and fossil collector. He was the son of a physician in Deal, who became a Huxleyite despite being deeply religious. Hulke became Huxley's ...

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 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment fact ...

. The specific name is derived from

Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the Cosmogony, origin and Cosmology#Metaphysical co ...

''πλοτός'', ''plotos'', the drifter. It is known solely from the

AR-1/56, a partial

skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...

which consists of AR-1-2045, a nearly complete but crushed skull; AR-1-2048, 4859, 4860, three

vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...

; AR-1-2046, a

rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...

; AR-1-2048, a metapodial; and AR-1-2049, 4861, 4862, three

osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...

. It was collected from the early

Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 M ...

-aged

Escucha Formation 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 bee ...

, at Santa Maria Mine located in the municipality of Ariño,

Teruel Province Teruel (Catalan: ''Terol'' ) is a province of Aragon, in the northeast of Spain. The capital is Teruel. It is bordered by the provinces of Tarragona, Castellón, Valencia (including its exclave Rincón de Ademuz), Cuenca, Guadalajara, and Zar ...

, of

Aragon Aragon ( , ; Spanish and an, Aragón ; ca, Aragó ) is an autonomous community in Spain, coextensive with the medieval Kingdom of Aragon. In northeastern Spain, the Aragonese autonomous community comprises three provinces (from north to sou ...

, 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 ''

''. Below is a

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

from that analysis:

References

{{Taxonbar, from=Q13794985 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