Avemetatarsalia

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Avemetatarsalians
Temporal range:
Ma
(possible Early Triassic record if Prorotodactylus is part of the clade[1])
Clockwise from top-left:
Dorygnathus banthensis (a pterosaur),
Alamosaurus sanjuanensis (a sauropod),
Edmontosaurus sp. (an ornithopod),
Daspletosaurus torosus (a theropod),
Styracosaurus albertensis (a ceratopsian) & Scolosaurus cutleri (an ankylosaurian),
and Grus grus (an extant avian).
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Sauropsida
Clade: Archosauria
Clade: Avemetatarsalia
Benton
, 1999
Subgroups
Synonyms
  • Dracones Haeckel, 1895
  • Ornithosuchia Huene, 1908
  • Ornithotarsi Gauthier, 1986
  • Pan-Aves Gauthier and de Queiroz, 2001

Avemetatarsalia (meaning "bird

Michael Benton in 1999. 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.[4]

Although dinosaurs and pterosaurs were the only avemetatarsalians to survive past the end of the Triassic, other groups flourished during the Triassic. The most

plesiomorphic ("primitive") known avemetatarsalians were the aphanosaurs. Aphanosaurs were rare, four-legged carnivores which were only properly distinguished as a group in 2017.[5] The split between dinosaurs and pterosaurs occurred just after aphanosaurs branched off the archosaur family tree. This split corresponds to the subgroup Ornithodira, defined as the last common ancestor of dinosaurs and pterosaurs, and all of its descendants. Until the discovery of aphanosaurs, Ornithodira and Avemetatarsalia were considered roughly equivalent concepts.[3]

Pterosauromorpha includes all avemetatarsalians closer to pterosaurs than to dinosaurs. True non-pterosaur pterosauromorphs have been historically difficult to determine. Small, insectivorous archosaurs of the family Lagerpetidae may potentially be examples, alongside the similar genus Scleromochlus.[6] Dinosauromorpha, on the other hand, includes all avemetatarsalians closer to dinosaurs than to pterosaurs. Probable non-dinosaur dinosauromorphs include the diverse and widespread silesaurids, as well as more controversial and fragmentary taxa such as Marasuchus, Lagosuchus, Nyasasaurus, and Saltopus. Lagerpetids were also traditionally considered dinosauromorphs,[7][3] though this has been more recently debated.[8][6][9]

Description

The "advanced mesotarsal" ankle present in most avemetatarsalians.

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 that of 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

flight feathers
.

Avemetatarsalians were generally more lightly built than crocodile-line archosaurs. They had smaller heads and usually a complete lack of osteoderms.

Origin

Bird-line archosaurs appear in the fossil record by the

rise of dinosaurs was slow and drawn out across much of the Triassic.[1] 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.[11]

Classification

In 1986,

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 "Panaves" (lit.'all birds', from Greek pan- + Latin aves), 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.[13]

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.[14] 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.[15] In fact, the name Ornithosuchia may be "illegal" under the PhyloCode because it does not include its eponymous taxon as part of its definition.[15]

Cladogram after Nesbitt et al. (2017),[16] with clade names from Cau (2018).[17]

Archosauria

Pseudosuchia (crocodile-line archosaurs)

Avemetatarsalia

Aphanosauria

Ornithodira 
†Pterosauromorpha

Pterosauria

 Dinosauromorpha 

Lagerpetidae

 
Dinosauriformes
 

Marasuchus

 
Dracohors
 

Kammerer et al. (2020) and Ezcurra et al. (2020) supported an alternative hypothesis regarding the relationships of lagerpetids. They were interpreted as non-pterosaur pterosauromorphs. This phylogeny would shorten the morphological and chronological gap perceived between pterosaurs and other stem-birds, and explain the origin of this group.[8][6] Bennett (2020) argued that Scleromochlus, a genus historically considered a relative of ornithodirans or even a basal pterosauromorph, was instead a non-archosaur archosauriform (possibly a doswelliid).[18]

In 2023, Nesbitt et al. described

phylogenetic analyses of Nesbit et al. (2023) are shown in the cladogram below:[19]

Archosauria

Pseudosuchia

Avemetatarsalia

References

  1. ^ .
  2. ^ .
  3. ^ .
  4. .
  5. .
  6. ^ .
  7. ^ a b Gauthier, Jacques (1986). "Saurischian monophyly and the origin of birds". In Padian, Kevin (ed.). The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Sciences. Vol. 8. San Francisco: California Academy of Sciences. pp. 1–55.
  8. ^
    PMID 32631980
    .
  9. .
  10. .
  11. ^ Dinosaur Evolution: Crocodile-Like Ancient Cousin, Teleocrater Rhadinus, Confuses Scientists
  12. ^ Sereno, P. C. 1991. Basal archosaurs: phylogenetic relationships and functional implications. Journal of Vertebrate Paleontology Memoir 2, 11(4, Supplement):1–53.
  13. .
  14. ^ Senter, P. (2005). "Phylogenetic taxonomy and the names of the major archosaurian (Reptilia) clades". PaleoBios. 25 (3): 1–7.
  15. ^ a b Taylor (2007). "Phylogenetic definitions in the pre-PhyloCode era; implications for naming clades under the PhyloCode" (PDF). PaleoBios. 27 (1): 1–6.
  16. S2CID 9095072
    .
  17. .
  18. .
  19. .

Sources