Goniopholis Sp
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''Goniopholis'' (meaning "angled scale") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 J ...
crocodyliform Crocodyliformes is a clade of Crurotarsi, 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 ...
that lived in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
during the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
and
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. Like other goniopholidids, it resembled living crocodilians, and probably had a similar ecology as semi-aquatic ambush predators.


Discovery and species

The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
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 organism ...
. 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 143.1 ±0.6 Ma and 137.05 ± 0.2 (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 154.8 ±0.8 Ma and 149.2 ±0.7 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 journalist, novelist, poet, and short-story writer. He was born in British Raj, British India, which inspired much ...
, "in recognition for his enthusiasm for natural sciences". ''G. kiplingi'' had skull reaching , it is one of the largest goniopholidid along with ''
Amphicotylus ''Amphicotylus'' is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was described in 1878.E. D. Cope. 1878. Descriptions of new extinct Vertebrata from the Upper Tertiary and Dakota Fo ...
milesi'' which had skull reaching . Based on skull length, total body length of ''G. kiplingi'' is estimated at . 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 asse ...
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 (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Daven ...
on the basis of bone
histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
and stable isotope analysis. The taxon ''Macellodus brodei'' was named in 1854 by Sir Richard Owen for a partial maxilla and referred jaws, with Owen interpreting the material as that of a lizard. The maxilla was considered missing my Hoffstetter in 1967, who designated a neotype, though this neotype was then removed from ''Macellodus'' and referred to the lacertilian '' Becklesisaurus''. Review by Richard Estes in 1983 rediscovered the type of ''Macellodus'' among crocodilian remains in the
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum (Lo ...
, recognizing that it belonged to the premaxilla of a crocodilian. Estes considered that ''Macellodus'' should be a synyonym of ''Goniopholis'', and ''G. brodei'' would have priority over ''G. simus'', but instead of advocating for synonymy Estes found that ''G. brodei'' is undiagnostic.


Formerly assigned species

Two species were referred to ''Goniopholis'' from
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. '' Goniopholis hartti'' from the
Lower Cretaceous Lower may refer to: * ''Lower'' (album), 2025 album by Benjamin Booker * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also * Nizhny {{Disambiguation ...
of Brazil is in fact a member of the genus ''
Sarcosuchus ''Sarcosuchus'' (), from Ancient Greek σάρξ (''sárx''), meaning "flesh", and σούχος (''soúkhos''), meaning "crocodile", is an extinct genus of crocodyliform a that lived during the Early Cretaceous, from the late Hauterivian to the ...
''. ''G. paulistanus'', based on two tooth crowns and a disassociated fragment of the right tibia from the Upper Cretaceous Bauru Group, has been reassigned to Itasuchidae and given its own genus ''
Roxochampsa ''Roxochampsa'' is an extinct genus of crocodylomorph from the Late Cretaceous of Brazil belonging to the sebecosuchian clade Itasuchidae. The type species is ''R. paulistanus''. Discovery and naming ''Roxochampsa'' was originally described ...
''. From
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, ''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 described in 1878.E. D. Cope. 1878. Descriptions of new extinct Vertebrata from the Upper Tertiary and Dakota Fo ...
'' and '' Denazinosuchus'', respectively, while ''G. felix'', ''G. gilmorei'', and ''G. stovalli'', all from the
Morrison Formation The Morrison Formation is a distinctive sequence of 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, sandstone, siltston ...
, are referable to ''Amphicotylus'' and closely related to '' Eutretauranosuchus'' which are known from the same formation. ''G. phuwiangensis'' is known from NE
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
, 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 t ...
''. '' Nannosuchus'' from the Early Cretaceous (
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 143.1 ±0.6 Ma and 137.05 ± 0.2 (million years ago) ...
stage) of England and
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
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'', respectively. Dollo's goniopholidid has also been assigned to ''Anteophthalmosuchus''. ''
Koumpiodontosuchus aprosdokiti ''Koumpiodontosuchus'' is an extinct genus of neosuchian crocodyliform from the Early Cretaceous Wessex Formation of England. The only species is ''K. aprosdokiti'', named in 2014. Discovery The first fossilised fragment of a skull was discover ...
'' from
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
was initially identified as a juvenile ''Goniopholis''.


Description

Like other goniophoilids, ''Goniopholis'' bears a superficial resemblance to modern crocodilians. However, unlike modern crocodilians and like other goniopholidids, the dermal armour covering the back was composed of two rows of large rectangular
scutes A scute () or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterio ...
running parallel down each side of the midline, with a "peg and groove" mechanism articulating the sets of plates together, with the outer edge of the plates deflected downwards.


Ecology

Goniopholidids likely had a similar ecology to modern crocodilians as semi-aquatic
ambush predators Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey usi ...
.


Classification

Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
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 Goniopholididae Early Cretaceous crocodylomorphs of Europe Prehistoric reptiles of Africa Late Jurassic crocodylomorphs of Europe Late Jurassic crocodylomorphs of North America Taxa named by Richard Owen Fossil taxa described in 1841 Prehistoric pseudosuchian genera