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

crocodyliform Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...

that lived in

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

and

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

during the

Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...

and

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 ...

. 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 The American alligator (''Alligator mississippiensis''), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the g ...

Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed throughout sub-Saharan Africa, occurring mostly in the central, eastern ...

Discovery and 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 ...

of the genus is ''G. crassidens'' which is known from the Berriasian of England, and the referable species ''G simus'' from the Berriasian of NW Germany, might be

conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...

. Other species that are referable to ''Goniopholis'' include ''G. kiplingi'' from the

Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...

of England, and ''G. baryglyphaeus'' from the Late Jurassic (

Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...

) of Portugal making it the oldest known ''Goniopholis'' species. The species ''G. kiplingi'' honors the author

Rudyard Kipling Joseph Rudyard Kipling ( ; 30 December 1865 – 18 January 1936)''The Times'', (London) 18 January 1936, p. 12. was an English novelist, short-story writer, poet, and journalist. He was born in British India, which inspired much of his work. ...

, "in recognition for his enthusiasm for natural sciences". Eggs attributed to ''Goniopholis'' were found in the Late Jurassic of Portugal. A partial skeleton of an indeterminate species of ''Goniopholis'' has been recovered from the Berriasian aged

Angeac-Charente bonebed The Angeac-Charente bonebed is a fossil deposit located near Angeac-Charente in western France. It dates to the Berriasian stage of the Early Cretaceous, and is coeval with the Purbeck Group of Southern England. It has amongst the most diverse assem ...

of France.Ronan Allain, Romain Vullo, Lee Rozada, Jérémy Anquetin, Renaud Bourgeais, et al.
Vertebrate paleobiodiversity of the Early Cretaceous (Berriasian) Angeac-Charente Lagerstätte (southwestern France): implications for continental faunal turnover at the J/K boundary
Geodiversitas, Museum National d’Histoire Naturelle Paris, In press. ffhal-03264773f ''Goniopholis'' have been inferred to have been

ectothermic An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...

on the basis of bone

histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...

and stable isotope analysis.

Formerly assigned species

Two species were referred to ''Goniopholis'' from

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

. ''

Goniopholis hartti ''Sarcosuchus'' (; ) is an extinct genus of crocodyliform and distant relative of living crocodilians that lived during the Early Cretaceous, from the late Hauterivian to the early Albian, 133 to 112 million years ago of what is now Africa and ...

'' from the

Lower Cretaceous Lower may refer to: *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is situated about five miles south west of Dursley, eight ...

of Brazil is in fact a member of the genus ''

Sarcosuchus ''Sarcosuchus'' (; ) is an extinct genus of crocodyliform and distant relative of living crocodilians that lived during the Early Cretaceous, from the late Hauterivian to the early Albian, 133 to 112 million years ago of what is now Africa and ...

''. ''G. paulistanus'', based on two tooth crowns and a disassociated fragment of the right tibia from the Upper Cretaceous

Bauru Group The Bauru Group is a geological group of the Bauru Sub-basin, Paraná Basin in Minas Gerais, São Paulo, General Salgado, Itapecuru-Mirim, Mato Grosso, Brazil whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils t ...

, has been reassigned to Itasuchidae and given its own genus '' Roxochampsa''. From

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

, ''G. lucasii'' and ''G. kirtlandicus'' are currently placed in their own genera ''

Amphicotylus ''Amphicotylus'' is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was species description, described in 1878.E. D. Cope. 1878. Descriptions of new extinct Vertebrata from the Upper Te ...

'' and ''

Denazinosuchus ''Denazinosuchus'' is a genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Upper Cretaceous Fruitland Formation and Kirtland Formation (late Campanian-early Maastrichtian) of the San Juan Basin, New Mexico. It is ...

'', respectively, while ''G. felix'', ''G. gilmorei'', and ''G. stovalli'', all from the

Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...

, are referable to ''Amphicotylus'' and closely related to ''

Eutretauranosuchus ''Eutretauranosuchus'' is an extinct genus of goniopholidid crocodyliform. ''E. delfsi'' is the only known species within the genus. Discovery and history The holotype, ''Eutretauranosuchus delfsi'', was discovered by Edwin Delfs in 1957 amo ...

'' which are known from the same formation. ''G. phuwiangensis'' is known from NE

Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...

, but this species is fragmentary and was recently reassigned to ''

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 ...

''. ''

Nannosuchus ''Nannosuchus'' (meaning "dwarf crocodile") is an extinct genus of goniopholidid mesoeucrocodylian from the Berriasian Middle Purbeck Formation of England that was originally named as a species of ''Goniopholis''. The type species, ''N. gracili ...

'' from the Early Cretaceous (

stage) of England and

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 ...

currently considered to be valid, was referred to as ''G. gracilidens'' by some authors. Willett's / Hulke's, Hooley's and Dollo's goniopholidids represent several complete specimens previously classified as either ''G. simus'' or ''G. crassidens'', and one of them was recently re-described as the new species, ''G. willetti''. More recently these specimens were removed from ''Goniopholis'', and two of them, Hooley's and Hulke's goniopholidids, have been already reassigned to their own genera ''

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 ...

'' and ''

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' ...

'', respectively. Dollo's goniopholidid has also been assigned to ''Anteophthalmosuchus''.

Classification

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 ...

including several ''Goniopholis'' species:

References

Sources

* * * * Owen, R. 1878. Monograph on The Fossil Reptilia of the Wealden and Purbeck Formations, Supplement no. VII. Crocodilia (''Goniopholis'', ''Pterosuchus'', and ''Suchosaurus''). ''Palaeontological Society Monograph'', p. 1-15. * * * {{Taxonbar, from=Q1011864 Early Cretaceous crocodylomorphs of Europe Prehistoric reptiles of Africa Late Jurassic crocodylomorphs of Europe Early Cretaceous reptiles of Europe Taxa named by Richard Owen Fossil taxa described in 1841 Prehistoric pseudosuchian genera