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Archosauriformes (
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 ...
for 'ruling lizards', and
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for 'form') is a clade of
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ag ...
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 rhynchoce ...
s that developed from
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, lizards, ...
ancestors some time in the Latest Permian (roughly 252
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
). It was defined by
Jacques Gauthier Jacques Armand Gauthier (born June 7, 1948 in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology. Life and career Gauthier is the so ...
(1994) as the clade stemming from the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of
Proterosuchidae Proterosuchidae is an early family of basal archosauriforms whose fossils are known from the Latest Permian and the Early Triassic of Europe, Asia, Africa, Australia and possibly South America. The name comes from Greek πρότερο- ("first" ...
and
Archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian ...
ia (the group that contains
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant mem ...
s,
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s and
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23  million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is ...
s ncluding_birds.html"_;"title="bird.html"_;"title="ncluding_bird">ncluding_birds">bird.html"_;"title="ncluding_bird">ncluding_birds;_Phil_Senter.html" ;"title="bird">ncluding_birds.html" ;"title="bird.html" ;"title="ncluding bird">ncluding birds">bird.html" ;"title="ncluding bird">ncluding birds; Phil Senter">bird">ncluding_birds.html" ;"title="bird.html" ;"title="ncluding bird">ncluding birds">bird.html" ;"title="ncluding bird">ncluding birds; Phil Senter (2005) defined it as the most exclusive clade containing ''Proterosuchus'' and Archosauria. These
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 rhynchoce ...
s, which include members of the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
Proterosuchidae and more advanced forms, were originally superficially crocodile-like animals with sprawling gaits and long snouts. Unlike the bulk of their
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
contemporaries, the proterosuchids survived the catastrophe at the end of the Permian. Within a few million years after the beginning of the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, the archosauriformes had diversified past the "proterosuchian"
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
. Other archosauriforms include the
Erythrosuchidae Erythrosuchidae (meaning "red crocodiles" in Greek) are a family of large basal archosauriform carnivores that lived from the later Early Triassic (Olenekian) to the early Middle Triassic (Anisian). Naming The family Erythrosuchidae was named by ...
(some of the earliest
sauropsid Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
apex predators), the
Euparkeriidae Euparkeriidae is an extinct family of small carnivorous archosauriforms which lived from the Early Triassic to the Middle Triassic (Anisian). While most other early archosauriforms walked on four limbs, euparkeriids were probably facultative b ...
(small, agile reptiles), and a variety of other strange reptiles such as proterochampsids, ''
Vancleavea ''Vancleavea'' is a genus of extinct, armoured, non- archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is ''V. campi'', named by Robert Long & Phillip A Murry in 1995. At that time, t ...
'', ''
Doswellia ''Doswellia'' is an extinct genus of archosauriform from the Late Triassic of North America. It is the most notable member of the family Doswelliidae, related to the proterochampsids. ''Doswellia'' was a low and heavily built carnivore which li ...
'', and ''
Asperoris ''Asperoris'' is an extinct genus of archosauriform reptile known from the Middle Triassic Manda Beds of southwestern Tanzania. It is the first archosauriform known from the Manda Beds that is not an archosaur. However, its relationships with ot ...
''. The most successful archosauriforms, and the only members to survive into the Jurassic, were the
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian ...
s. Archosauria includes
crocodylia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living r ...
ns, birds, and extinct relatives such as
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
s, rauisuchids, pterosaurs, and non-avian dinosaurs. Pre-''
Euparkeria ''Euparkeria'' (; meaning "Parker's good animal", named in honor of W.K. Parker) is an extinct genus of archosauriform from the Middle Triassic of South Africa. It was a small reptile that lived between 245-230 million years ago, and was close ...
'' Archosauriformes have previously been included in the suborder
Proterosuchia Proterosuchia is one of the suborders of the paraphyletic group Thecodontia; containing the most primitive and ancestral forms. These were primitive, vaguely crocodile-like, archosauriforms that mostly lived during the Early Triassic epoch. The ...
of the order
Thecodontia Thecodontia (meaning 'socket-teeth'), now considered an obsolete taxonomic grouping, was formerly used to describe a diverse "order" of early archosaurian reptiles that first appeared in the latest Permian period and flourished until the end of t ...
. Under modern
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
methodology, Proterosuchia and Thecodontia have been rejected as
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
assemblages, and the pre-archosaurian taxa are simply referred to as basal Archosauriformes. In 2016,
Martin Ezcurra Martin may refer to: Places * Martin City (disambiguation) * Martin County (disambiguation) * Martin Township (disambiguation) Antarctica * Martin Peninsula, Marie Byrd Land * Port Martin, Adelie Land * Point Martin, South Orkney Islands Aus ...
provided a name for the clade including all archosauriforms more crownward than erythrosuchids. He named the clade Eucrocopoda, which translates to "true crocodile feet" in reference to the possession of a crocodilian-style crurotarsal ankle.


Metabolism

Vascular density and
osteocyte An osteocyte, an oblate shaped type of bone cell with dendritic processes, is the most commonly found cell in mature bone. It can live as long as the organism itself. The adult human body has about 42 billion of them. Osteocytes do not divide and ...
density, shape and area have been used to estimate the bone growth rate of archosaurs, leading to the conclusion that this rate had a tendency to grow in ornithodirans and decrease in pseudosuchians. The same method also supports the existence of high resting metabolical rates similar to those of living endotherms (mammals and birds) in the ''
Prolacerta ''Prolacerta'' is a genus of archosauromorph from the lower Triassic of South Africa and Antarctica. The only known species is ''Prolacerta broomi''. The generic name ''Prolacerta'' is derived from Latin meaning “before lizard” and its speci ...
''-Archosauriformes clade that were retained by most subgroups, though decreased in ''Proterosuchus,''
Phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent ...
ia and Crocodilia. Erythrosuchids and ''Euparkeria'' are basal archosauriforms showing signs of high growth rates and elevated metabolism, with ''Erythrosuchus'' possessing a rate similar of the fastest-growing dinosaurs. Sexual maturity in those Triassic taxa was probably reached quickly, providing advantage in a habitat with unpredictable variation from heavy rainfall to drought and high mortality. ''
Vancleavea ''Vancleavea'' is a genus of extinct, armoured, non- archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is ''V. campi'', named by Robert Long & Phillip A Murry in 1995. At that time, t ...
'' and ''Euparkeria'', which show slower growth rates compared to ''Erythrosuchus'', lived after the climatic stabilization. Early crown archosaurs possessed increased growth rates, which were retained by ornithodirans. Ornithosuchians and poposaurs are stem-crocodilians that show high growth rates similar to those of basal archosauriforms. Developmental, physiological, anatomical and palaeontological lines of evidence indicate that crocodilians evolved from endothermic ancestors. Living crocodilians are ambush predators adapted to a semi-aquatic lifestyle that benefits from ectothermy due to the lower oxygen intake that allows longer diving time. The mixing of oxygenated and deoxygenated blood in their circulatory system is apparently an innovation that benefits ectothermic life. Earlier archosaurs likely lacked those adaptations and instead had completely separated blood as birds and mammals do. A similar process occurred in phytosaurs, which were also semi-aquatic. The similarities between
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
,
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n and coelurosaurian integument suggest a common origin of thermal insulation (feathers) in ornithodirans at least 250 million years ago. Erythrosuchids living in high latitudes might have benefited from some sort of insulation. If ''
Longisquama ''Longisquama'' is a genus of extinct reptile. There is only one species, ''Longisquama insignis'', known from a poorly preserved skeleton and several incomplete fossil impressions from the Middle to Late Triassic Madygen Formation in Kyrgyzsta ...
'' was an archosauromorph, it could be associated with the origin of feathers.


Relationships

Below is a cladogram from Nesbitt (2011): Below is a cladogram from Sengupta ''et al.'' (2017), based on an updated version of Ezcurra (2016) that reexamined all historical members of the "Proterosuchia" (a
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conve ...
historical group including
proterosuchid Proterosuchidae is an early family of basal archosauriforms whose fossils are known from the Latest Permian and the Early Triassic of Europe, Asia, Africa, Australia and possibly South America. The name comes from Greek πρότερο- ("first" ...
s and erythrosuchids). The placement of fragmentary taxa that had to be removed to increase tree resolution are indicated by dashed lines (in the most derived position that they can be confidently assigned to). Taxa that are ''
nomina dubia In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'' are indicated by the note "dubium". Bold terminal taxa are collapsed.


Sources

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References


External links


Paleos
{{Taxonbar, from=Q282487 Lopingian first appearances Extant Permian first appearances Taxa named by Jacques Gauthier Fossil taxa described in 1986