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 ...
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 ...
reptiles 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, li ...
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 s ...
(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-avi ...
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 ...
s,
pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the Order (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...
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 t ...
s
ncluding birds">bird.html" ;"title="ncluding bird">ncluding birds; Phil Senter">bird">ncluding_bird<_a>s.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
reptiles, which include members of the Family (biology), family 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 mor ...
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. 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 s ...
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 ...
(small, agile reptiles), and a variety of other strange reptiles such as
proterochampsids, ''
Vancleavea'', ''
Doswellia'', and ''
Asperoris''. The most successful archosauriforms, and the only members to survive into the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
, 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-avi ...
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 ...
ns,
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, 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 t ...
'' Archosauriformes have previously been included in the suborder
Proterosuchia 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 ...
. 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 ch ...
methodology, Proterosuchia and Thecodontia have been rejected as
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
assemblages, and the pre-archosaurian taxa are simply referred to as basal Archosauriformes. In 2016,
Martin Ezcurra 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 spec ...
''-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 gr ...
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'' 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 (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...
,
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
coelurosauria
Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.
Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, ty ...
n 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'' 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 conver ...
historical group including proterosuchids 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
*
*
References
External links
Paleos
{{Taxonbar, from=Q282487
Lopingian first appearances
Extant Permian first appearances
Taxa named by Jacques Gauthier
Fossil taxa described in 1986