Anteophthalmosuchus
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''Anteophthalmosuchus'' (meaning "forward-pointing eye crocodile") 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 ...
of
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 ...
, 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 ...
, and western
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to th ...
.


Discovery

The holotype specimen of ''Anteophthalmosuchus'', 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 ...
, includes a well-preserved skull and partial skeleton. This specimen has been known since 1904 and was identified as the "Tie Pits specimen" or the "Hooley specimen" after
Reginald Walter Hooley Reginald Walter Hooley (5 September 1865 – 5 May 1923) was a businessman and amateur paleontologist, collecting on the Isle of Wight. He is probably best remembered for describing the dinosaur ''Iguanodon atherfieldensis'', now ''Mantellisaurus ...
, an amateur paleontologist who had described it in 1905. Hooley had originally attributed the specimen to the previously named species '' Goniopholis crassidens''. Additional referred specimens include a partial disarticulated skeleton and a partial skull that may represent a juvenile specimen. 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 ...
, Hooley's specimen was redescribed as a distinct genus and species of goniopholidid called ''Anteophthalmosuchus hooleyi''. The genus name means "forward-pointing eye crocodile" because the specimen's eye sockets are positioned high on the skull and angle forward rather than to the side as in most other flat-skulled crocodyliforms, and the species name honors Hooley. Features that distinguish ''A. hooleyi'' from ''Goniopholis crassidens'' include the lack of a hole in the lower jaw called the mandibular fenestra, very wide supratemporal fenestrae (openings) on the
skull table The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
, and a bone above the eye socket called the
palpebral An eyelid is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. The human eyel ...
that is small and does not extend over the socket as in some other goniopholidids. Two specimens from
Bernissart Bernissart (; pcd, Bernissåt) is a Municipalities of Belgium, municipality of Wallonia located in the Hainaut Province, province of Hainaut, Belgium. On January 1, 2006, Bernissart had a total population of 11,458. The total area is 43.42  ...
, Belgium, collectively referred to as "Dollo's goniopholidid", was referred to ''A. hooleyi'' in a 2016 redescription. The specimens, which consist of complete skeletons (one missing the skull) from the
Sainte-Barbe Clays Formation 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 ...
, were originally referred to ''Goniopholis simus'' by Dollo. In 2016, they were recognized as specimens of ''A. hooleyi'' due to their distinctive eye sockets, among other defining characteristics of the species. While the Dollo specimens do not possess
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
s that are pointed at the
rostral Rostral may refer to: Anatomy * Rostral (anatomical term), situated toward the oral or nasal region * Rostral bone, in ceratopsian dinosaurs * Rostral organ, of certain fish * Rostral scale, in snakes and scaled reptiles Other uses * Rostral colu ...
end, a trait in 2011 to establish ''A. hooleyi'' as a distinct species, and also differs in the relative proportions of the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
and
foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
, the authors considered these traits invalid diagnostic characteristics, as they could be a consequence of age differences or preservational artifacts. However, in 2017, Dollo's goniopholidid was moved to a new species, ''Anteophthalmosuchus epikrator'', based on comparisons with newly-discovered specimens from the Isle of Wight. A second species, ''Anteophthalmosuchus escuchae'', 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 ...
. It is known solely from the
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 ...
AR-1-1097 which consists of a partial skull. 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 ''
Hulkepholis plotos ''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'' i ...
''.


Description

At an estimated in length, ''A. hooleyi'' would have been the largest crocodyliform in the Wealden faunal assemblage, larger than the contemporaneous species ''
Hylaeochampsa vectiana ''Hylaeochampsa'' is an extinct genus of eusuchian crocodylomorphs. It is known only from a partial skull recovered from Barremian-age rocks of the Lower Cretaceous Vectis Formation ( Wealden Group) of the Isle of Wight. This skull, BMNH R 177, ...
'', '' Leiokarinosuchus brookensis'', and '' Vectisuchus leptognathus''.


Phylogeny

''Anteophthalmosuchus hooleyi'' was included in a phylogenetic analysis of goniopholidids that was published soon before the specimen was redescribed. The Hooley and Dollo specimens were found to be the closest relatives of a specimen called "Hulke's goniopholidid", now named '' Hulkepholis willetti''. 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=Q4771296 Early Cretaceous crocodylomorphs of Europe Fossil taxa described in 2011 Early Cretaceous reptiles of Europe Prehistoric pseudosuchian genera