Allokotosauria is a

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

of early

archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...

s from the Middle to

Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...

known from

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

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 ...

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 ...

and

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 ...

. Allokotosauria was first described and named when a new

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

grouping of specialized herbivorous archosauromorphs was recovered by Sterling J. Nesbitt, John J. Flynn, Adam C. Pritchard, J. Michael Parrish, Lovasoa Ranivoharimanana and André R. Wyss in

2015 File:2015 Events Collage new.png, From top left, clockwise: Civil service in remembrance of November 2015 Paris attacks; Germanwings Flight 9525 was purposely crashed into the French Alps; the rubble of residences in Kathmandu following the Apri ...

. The name Allokotosauria is derived from

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

meaning "strange reptiles" in reference to unexpected grouping of early archosauromorph with a high disparity of features typically associated with

herbivory A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

History

Nesbitt ''et al.'' (2015) defined the group as a

stem-based taxon Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...

containing ''

Azendohsaurus madagaskarensis ''Azendohsaurus'' is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, ''Azendohsaurus laaroussii'', was described and named by Jean- ...

'' and '' Trilophosaurus buettneri'' and all taxa more closely related to them than to ''

Tanystropheus longobardicus ''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and tail combined. T ...

'', ''

Proterosuchus fergusi ''Proterosuchus'' is an extinct genus of archosauriform reptiles that lived during the Early Triassic. It contains three valid species: the type species ''P. fergusi'' and the referred species ''P. alexanderi'' and ''P. goweri''. All three sp ...

'', ''

Protorosaurus speneri ''Protorosaurus'' ("first lizard") is a genus of lizard-like early reptiles. Members of the genus lived during the late Permian period in what is now Germany and Great Britain. Once believed to have been an ancestor to lizards, ''Protorosaurus' ...

'' or ''

Rhynchosaurus articeps ''Rhynchosaurus'' (''beaked lizard'') is a genus of rhynchosaur that lived during the Middle Triassic Period (geology), period. It lived in Europe. It was related to the archosauria, archosaurs, but not within that group. The type species of ''R ...

''. Therefore, Allokotosauria includes the families

Azendohsauridae Azendohsauridae is a family of allokotosaurian archosauromorphs that lived during the Middle to Late Triassic period, around 242-216 million years ago. The family was originally named solely for the eponymous ''Azendohsaurus'', marking out its ...

and

Trilophosauridae Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is ''Trilophosaurus'', a herbivore up to long. It had a short, unusually heavily built skull, equipped with massive, broad flattened che ...

by definition, as well as the potentially more basal ''

Pamelaria ''Pamelaria'' is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, ''Pamelaria dolichotrachela'', from the Middle Triassic of India. ''Pamelaria'' has sprawling legs, a long neck, and a pointed skull with no ...

'' which is closer to them than to other early archosauromorphs. ''Pamelaria'' is the earliest known allokotosaur, dating to the

Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic Ep ...

India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...

. Azendohsauridae is currently represented by a single genus ''

Azendohsaurus ''Azendohsaurus'' is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, ''Azendohsaurus laaroussii'', was described and named by Jean-Mi ...

'' known from the

Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic). ...

Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...

, while trilophosaurids are mostly known from the

Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...

stages of

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 potentially

European Russia European Russia (russian: Европейская Россия, russian: европейская часть России, label=none) is the western and most populated part of Russia. It is geographically situated in Europe, as opposed to the cou ...

, though one member of the latter group, '' Variodens inopinatus'', is known from

Rhaetian The Rhaetian is the latest age of the Triassic Period (in geochronology) or the uppermost stage of the Triassic System (in chronostratigraphy). It was preceded by the Norian and succeeded by the Hettangian (the lowermost stage or earliest age ...

. According to studies of ''

Arctosaurus ''Arctosaurus'' is an extinct genus of archosauromorph, possibly an allokotosaurian, but was often classified as a sauropodomorph dinosaur between 1900 and 1976. Although it has also been classified as a theropod, recent review finds that the ...

'' material from

Cameron Island Cameron Island is one of the uninhabited members of the Queen Elizabeth Islands in the Canadian arctic islands in Nunavut, Canada. Located in the Arctic Ocean, close to Bathurst Island, it has an area of , long and wide. Île Vanier lies imme ...

in Canada, the latter may have been an allokotosaurian because of the similarities with ''

'' due to the presence of a posterior ridge from the centrum to the diapophyses which extends from the diapophysis all the way to the posterior ventrolateral corner of the centrum. This ridge overhangs a deep groove in the lateral surface of the centrum.

Description and phylogeny

Allokotosauria is most notably characterized by wrinkled side surface of orbital border of the

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 ...

, expanded and hooked

quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms upper ...

head on the posterior side, and a prominent tubercle developed above to the glenoid fossa of the

scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...

, although there are other unambiguous traits that differentiate it from other early archosauromorphs. 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 ...

showing the phylogenetic relationships of Allokotosauria within

Archosauromorpha Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...

as recovered by Nesbitt ''et al.'' (2015). Ezcurra (2016) also recovered a highly supported Allokotosauria with the same topology (including only ''

'', ''

'' and '' Trilophosaurus buettneri'' in his analysis), but noted that ''Pamelaria'' is nearly as likely to represent a basal

azendohsaurid Azendohsauridae is a family of allokotosaurian archosauromorphs that lived during the Middle to Late Triassic period, around 242-216 million years ago. The family was originally named solely for the eponymous '' Azendohsaurus'', marking out its ...

instead. Sengupta ''et al.'' (2017) described a new azendohsaurid and recovered ''Pamelaria'' as an azendohsaurid. Pritchard and Nesbitt (2017) found support for the inclusion of

Kuehneosauridae Kuehneosauridae is an extinct family of small, lizard-like gliding diapsids known from the Triassic period of Europe and North America. They are distinguished from other diapsids by their 'wings' formed by elongated ribs. These allowed the animal ...

within Allokotosauria.

References

{{Taxonbar, from=Q25095775 Triassic reptiles Triassic reptiles of Africa Triassic reptiles of Asia Triassic reptiles of Europe Triassic reptiles of North America Anisian first appearances Rhaetian extinctions Taxa named by Sterling Nesbitt