Avemetatarsalia (meaning "bird metatarsals") is a clade name established by British palaeontologist Michael Benton in 1999 for all crown group archosaurs that are closer to birds than to crocodilians. An alternate name is Pan-Aves, or "all birds", in reference to its definition containing all animals, living or extinct, which are more closely related to birds than to crocodilians. Almost all avemetatarsalians are members of a similarly defined subgroup, Ornithodira. Ornithodira is defined as the last common ancestor of dinosaurs and pterosaurs, and all of its descendants.
Members of this group include the Dinosauromorpha, Pterosauromorpha, the genus Scleromochlus, and Aphanosauria. Dinosauromorpha contains more basal forms, including Lagerpeton and Marasuchus, as well as more derived forms, including dinosaurs. Birds belong to the dinosaurs as members of the theropods. Pterosauromorpha contains Pterosauria, which were the first vertebrates capable of true flight. Aphanosauria is a Triassic group of gracile carnivorous quadrupeds which was recognized in 2017.
|Clockwise from top-left:|
Tupuxuara leonardi (a pterosaur),
Alamosaurus sanjuanensis, (a sauropod),
Tsintaosaurus spinorhinus (an ornithopod),
Daspletosaurus torosus (a tyrannosaur),
Pentaceratops sternbergii (a ceratopsian),
and Grus grus (an extant avian).
The foundational characteristic is the "advanced mesotarsal" ankles, which are characterized by a large astragalus and a small calcaneum. This ankle orientation operated on a single hinge, allowing for better mobility. Probably as a result of this change, the common ancestor of the avemetatarsalians had an upright, bipedal posture, with their legs extending vertically, similar to the situation in mammals.
Feathers and other filamentary structures are known across the avemetatarsalians, from the downy pycnofibers of pterosaurs, to quill-like structures present in ornithischian dinosaurs, such as Psittacosaurus and Tianyulong, to feathers in theropod dinosaurs and their descendants, birds.
Two clades of avemetatarsalians,pterosaurs and birds, independently evolved flight. Pterosaurs are the earliest vertebrates known to have evolved powered flight. Their wings are formed by a membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger. Birds evolved flight much later. Their wings formed from elongated fingers, and their arms, all covered with flight feathers.
Bird-line archosaurs appear in the fossil record by the Anisian stage of the Middle Triassic about 245 million years ago, represented by the dinosauriform Asilisaurus. However, Early Triassic fossil footprints reported in 2010 from the Świętokrzyskie (Holy Cross) Mountains of Poland may belong to a more primitive dinosauromorph. If so, the origin of avemetatarsalians would be pushed back into the early Olenekian age, around 249 Ma. The oldest Polish footprints are classified in the ichnogenus Prorotodactylus and were made by an unknown small quadrupedal animal, but footprints called Sphingopus, found from Early Anisian strata, show that moderately large bipedal dinosauromorphs had appeared by 246 Ma. The tracks show that the dinosaur lineage appeared soon after the Permian–Triassic extinction event. Their age suggests that the rise of dinosaurs was slow and drawn out across much of the Triassic. The primitive traits found in the quadrupedal aphanosaur Teleocrater shows that the earliest avemetatarsalians had many pseudosuchian-like features, and that the traits typical for the group evolved later.
In 1986, Jacques Gauthier defined the name Ornithosuchia (previously coined by Huene) for a branch-based clade including all archosaurs more closely related to birds than to crocodiles. In the same year, Gauthier also coined and defined a slightly more restrictive node-based clade, Ornithodira, containing the last common ancestor of the dinosaurs and the pterosaurs and all of its descendants. Paul Sereno in 1991 gave a different definition of Ornithodira, one in which Scleromochlus was explicitly added. It was thus a potentially larger group than the Ornithodira of Gauthier. In 1999 Michael Benton concluded that Scleromochlus was indeed outside Gauthier's original conception of Ornithodira, so he named a new branch-based clade for this purpose: Avemetatarsalia, named after the birds (Aves), the last surviving members of the clade, and the metatarsal ankle joint that was a typical character of the group. Avemetatarsalia was defined as all Avesuchia closer to Dinosauria than to Crocodylia. In 2005, Sereno stated the opinion that Ornithodira was not a useful concept, whereas Avemetatarsalia was. In 2001, the same clade was given the name Pan-Aves (meaning "all [pan in Greek] birds" [aves in Latin]), coined by Jacques Gauthier. He defined it as the largest and most inclusive clade of archosaurs containing Aves (birds, anchored on Vultur gryphus) but not Crocodylia (anchored on Crocodylus niloticus). Gauthier referred Aves, all other Dinosauria, all Pterosauria, and a variety of Triassic archosaurs, including Lagosuchus and Scleromochlus, to this group.
In a 2005 review of archosaur classification, Phil Senter attempted to resolve this conflicting set of terminology by applying strict priority to names based on when and how they were first defined. Senter noted that Ornithosuchia, the earliest name used for the total group of archosaurs closer to birds than to crocodiles, should be the valid name for that group, and have precedence over later names with identical definitions, such as Avemetatarsalia and Pan-Aves. While this has been followed by some researchers, others have either continued to use Avemetatarsalia or Ornithodira, or have followed Senter only reluctantly. Mike Taylor (2007) for example noted that, while Senter is correct in stating that Ornithosuchia has priority, this is "undesirable" because it probably excludes the eponymous family Ornithosuchidae, and questioned the utility of using priority before the PhyloCode is implemented to govern it. In fact, the name Ornithosuchia may be "illegal" under the PhyloCode because it does not include its eponymous taxon as part of its definition.
Aphanosauria ("hidden lizards") is group of reptiles distantly related to dinosaurs (including birds). They were at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchia, includes modern crocodilians. Aphanosaurs possessed features from both groups, indicating that they are the oldest and most primitive known clade of avemetatarsalians, at least in terms of their position on the archosaur family tree. Other avemetatarsalians include the flying pterosaurs, small bipedal lagerpetids, herbivorous silesaurids, and the incredibly diverse dinosaurs, which survive to the present day in the form of birds. Aphanosauria is formally defined as the most inclusive clade containing Teleocrater rhadinus and Yarasuchus deccanensis but not Passer domesticus (House sparrow) or Crocodylus niloticus (Nile crocodile). This group was first recognized during the description of Teleocrater. Although only known by a few genera, Aphanosaurs had a widespread distribution across Pangaea in the Middle Triassic.They were fairly slow quadrupedal long-necked carnivores, a biology more similar to basal archosaurs than to advanced avemetatarsalians such as pterosaurs, lagerpetids, and early dinosaurs. In addition, they seemingly possess 'crocodile-normal' ankles (with a crurotarsal joint), showing that 'advanced mesotarsal' ankles (the form acquired by many dinosaurs, pterosaurs, lagerpetids, and advanced silesaurids) were not basal to the whole clade of Avemetatarsalia. Nevertheless, they possessed elevated growth rates compared to their contemporaries, indicating that they grew quickly, more like birds than modern reptiles. Despite superficially resembling lizards, the closest modern relatives of aphanosaurs are birds.Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of birds and crocodilians. This group also includes all extinct dinosaurs, extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most recent common ancestor of living birds and crocodilians and all of its descendants. It includes two main clades: Pseudosuchia, which includes crocodilians and their extinct relatives, and Avemetatarsalia, which includes birds and their extinct relatives (such as non-avian dinosaurs and pterosaurs).Asilisaurus
Asilisaurus ( ah-SEE-lee-SOR-əs; from Swahili, asili ("ancestor" or "foundation"), and Greek, σαυρος (sauros, "lizard") is an extinct genus of silesaurid archosaur. It is one of the oldest known animals on the dinosaur/pterosaur side of the archosaurian tree (the Avemetatarsalia), dating to about 245 million years ago.
Fossils were uncovered in Tanzania and date back to the Anisian stage of the Middle Triassic. It was described in 2010 by a team of researchers from the United States, Germany, and South Africa, in the journal Nature; the type species is A. kongwe. It is the first example of an avian-line radiation during the Anisian, with the diversification of archosaurs during this time previously only documented from crocodylian-line archosaurs. It was the first non-dinosaurian dinosauriform recovered from Africa.Asilisaurus measured from 1 to 3 metres (3 to 10 ft) long and 0.5 to 1 metre (2 to 3 ft) high at the hip, and weighed 10 to 30 kilograms (20 to 70 lb).In a recently published research conducted by paleontologists from Virginia Tech, Asilisaurus was thought to be ontogenetically of polymorphism, just like a big family with siblings and cousins but differing in height or body mass. Researchers suggested that the early dinosaurs were also the case because of their close affinities. Through examining the muscle scars of femora of enough Asilisaurus specimens as well as histological studies, they proposed that instead of considering variations as sexual differences, they are better interpreted as individual differences.Averostra
Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.Avetheropoda
Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.Cerapoda
Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.Crurotarsi
Crurotarsi is a group of archosauriform reptiles that includes the archosaurs (represented today by birds and crocodilians) and the extinct, crocodile-like phytosaurs. The name is derived from the Latin word crus and the Greek word tarsos; it refers to the specialized articulation between crus and tarsus—specifically between fibula and calcaneum—present in the skeletons of suchians and phytosaurs, with a hemicylindrical condyle on the calcaneum articulating against fibula.Dinosauriformes
Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.Dongusuchus
Dongusuchus (meaning Donguz River crocodile in Greek, for the area where the type specimen was found) is an extinct genus of archosauriform. Fossils have been found from the Donguz Formation outcropping on the banks of the Donguz River in the Orenburg Oblast of Russia. They are associated with a fossil assemblage called the Eryosuchus Fauna, named after the capitosaurid Eryosuchus, the most common organism found from the assemblage. The locality dates back to the Anisian and early Ladinian stages of the Middle Triassic.Sennikov (1988) and Gower and Sennikov (2000) suggested that Dongusuchus was a gracile rausuchian with a long, sigmoidally curved neck, unlike the more typical robust short-necked rauisuchians that appear later in the Triassic. More recently, Nesbitt (2009) argued that Dongusuchus most probably represents a non-archosaurian archosauriform. According to Nesbitt (2009), the poorly-defined crista tibiofibularis and the absence of a distinct anteromedial tuber of the proximal portion in Dongusuchus suggest that it is not a member of Archosauria. Although Gower and Sennikov (2000) suggested that the distinct sigmoidal shape of Dongusuchus femur is unique, a similar shape is present in the femora of some phytosaurs. A paratype tibia was also found to be more closely related to Euparkeria and phytosaurs, on the basis of its convex and rounded distal surface. Additionally, the proximal surface of the tibia lacks a trait present in nearly all pseudosuchians, a depression on its lateral condyle. Nesbitt assigned Dongusuchus to Archosauriformes on the basis of the following traits: its femur has a low fourth trochanter, and the distal condyles do not expand markedly beyond the shaft. These traits suggest that Dongusuchus was an archosauriform more derived than Erythrosuchus.Dongusuchus was also excluded from Archosauria by Niedźwiedzki et al. (2014) and a new large cladistic analysis of archosauromorphs by Ezcurra (2016) found Dongusuchus to be the sister taxon to the Indian Yarasuchus. Both Dongusuchus and Yarasuchus were recovered in a clade with Spondylosoma and Teleocrater by Nesbitt et al. (2017) at the base of Avemetatarsalia, making them more closely related to bird-line archosaurs.Eopterosauria
Eopterosauria is a group of basal pterosaurs from the Triassic, which form their own clade. The term was first used in Andres et al. (2014) to include Preondactylus, Austriadactylus, Peteinosaurus and Eudimorphodontidae. Inside the group were two other new clades, Preondactylia, which included Preondactylus and Austriadactylus, and Eudimorphodontoidea, to include Eudimorphodontidae and Raeticodactylidae. Eopterosauria was defined as "the least inclusive clade containing Preondactylus buffarinii and Eudimorphodon ranzii". The specimen BSP 1994, previously assigned to Eudimorphodon, was named the separate taxon Austriadraco in 2015, and assigned to the new family Austriadraconidae, but further classification was not described. The following phylogenetic analysis follows the topology of Andres et al. (2014).Jeholosauridae
Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.Lutungutali
Lutungutali (meaning "high hip" in the Bemba language) is an extinct genus of silesaurid dinosauriform from the Middle Triassic of Zambia. The single type species of the genus is Lutungutali sitwensis. Lutungutali was named in 2013 and described from a fossil specimen, holotype NHCC LB32, including hip bones and tail vertebrae. The specimen was collected in 2009 from the upper Ntawere Formation, which dates to the Anisian stage of the Middle Triassic. Lutungutali is the first known silesaurid from Zambia and, along with the Tanzanian silesaurid Asilisaurus and dinosauriform Nyasasaurus, the oldest bird-line archosaur known from body fossils (i.e. parts of the skeleton).Melanorosauridae
The Melanorosauridae were a family of sauropodomorph dinosaurs which lived during the Late Triassic and Early Jurassic. The name Melanorosauridae was first coined by Friedrich von Huene in 1929. Huene assigned several families of dinosaurs to the infraorder "Prosauropoda": the Anchisauridae, the Plateosauridae, the Thecodontosauridae, and the Melanorosauridae. Since then, these families have undergone numerous revisions. Galton and Upchurch (2004) considered Camelotia, Lessemsaurus, and Melanorosaurus members of the family Melanorosauridae. A more recent study by Yates (2007) indicates that the melanorosaurids were instead early sauropods.Neotheropoda
Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.Orionides
Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.Riojasauridae
Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).Scleromochlus
Scleromochlus (Greek for "hard fulcrum") is an extinct genus of small avemetatarsalians from the Late Triassic period.Spondylosoma
Spondylosoma (meaning "vertebra body") is a genus of avemetatarsalian archosaur belonging to the clade Aphanosauria from the late Ladinian-age Middle Triassic Lower Santa Maria Formation in Geopark of Paleorrota, Brazil.
Avemetatarsalians (Dinosauromorpha and relatives)