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Avialae ("bird wings") 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 ter ...
containing the only living
dinosaurs 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 the ...
, the
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. It is usually defined as all
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaurs more closely related to birds (Aves) than to
deinonychosaurs Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica,Case, J.A., Marti ...
, though alternative definitions are occasionally used (see below). '' Archaeopteryx lithographica'', from the late
Jurassic Period The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
Solnhofen Formation of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
, is possibly the earliest known avialan which may have had the capability of powered flight,Supplementary info
/ref> though it might have been a deinonychosaur instead. Several older (but non flight-capable) avialans are known from the late Jurassic
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian- Oxfordian stages). It is known for its exceptionally preserved fossils, including those of ...
of
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, dated to about 160 million years ago.


Definition

Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s closer to birds than to ''
Deinonychus ''Deinonychus'' ( ; ) is a genus of dromaeosaurid theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period, about 115–108 million y ...
''."Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.) (2004). ''The Dinosauria'', Second Edition. University of California Press., 861 pp. A nearly identical definition, "the theropod group that includes all taxa closer to ''
Passer ''Passer'' is a genus of sparrows, also known as the true sparrows. The genus contains 28 species and includes the house sparrow and the Eurasian tree sparrow, two of the most common birds in the world. They are small birds with thick bills for ...
'' than to '' Dromaeosaurus''", was used by Agnolín and Novas (2013) for their clade Averaptora, operating under the assumption that
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil disco ...
s and birds were more closely related to each other than to dromaeosaurs. They also redefine Avialae as the smallest clade containing ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'' and modern birds. Additionally, beginning in the late 2000s and early 2010s, several groups of researchers began adding the genus ''
Troodon ''Troodon'' ( ; ''Troödon'' in older sources) is a wastebasket taxon and a dubious genus of relatively small, bird-like dinosaurs known definitively from the Campanian age of the Late Cretaceous period (about 77  mya). It includes at leas ...
'' as an additional specifier in the definition of Avialae. ''Troodon'' had long been considered a close relative of the dromaeosaurids in the larger group Deinonychosauria, though some contemporary studies found it and other troodontids more closely related to modern birds, and so it has been specifically excluded from Avialae in more recent studies. Avialae is also occasionally defined as an apomorphy-based clade (that is, one based on derived characteristics that were not present among lineage predecessors).
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 ...
, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
s used in flapping
flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...
, and the birds that descended from them.Gauthier, J. (1986). "Saurischian monophyly and the origin of birds." In: K. Padian, ed. ''The origin of birds and the evolution of flight.'' San Francisco: California, Acad.Sci. pp.1–55. (Mem.Calif.Acad.Sci.8.)Gauthier, J., and de Queiroz, K. (2001).
Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name Aves
" Pp. 7-41 in ''New perspectives on the origin and early evolution of birds: proceedings of the International Symposium in Honor of John H. Ostrom'' (J. A. Gauthier and L. F. Gall, eds.). Peabody Museum of Natural History, Yale University, New Haven, Connecticut, U.S.A.


Differentiation from Aves

Gauthier and de Queiroz (page 34) identified four conflicting ways of defining the term "Aves", which is a problem since the same biological name is being used four different ways. They proposed a solution, number 4 below, which is to reserve the term Aves only for the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
, the last common ancestor of all living birds and all of its descendants. Other definitions of Aves found in literature were reassigned to other clade names. # Aves can mean all reptiles closer to birds than to
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 me ...
s (alternatively
Avemetatarsalia Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. ...
Panaves # Aves can mean those advanced
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-avia ...
s with feathers (alternatively Avifilopluma) # Aves can mean those
feathered dinosaur A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been s ...
s that can fly (alternately Avialae) # Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "crown group"). (alternatively
Neornithes 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 ...
) Under the fourth definition ''Archaeopteryx'' is an avialan, and not a member of Aves. Gauthier's proposals have been adopted by many researchers in the field of paleontology and bird evolution, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is sometimes used synonymously with the vernacular term "bird" by these researchers.


Evolution

The earliest known avialans come from the
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian- Oxfordian stages). It is known for its exceptionally preserved fossils, including those of ...
of
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, which has been dated to the late
Jurassic The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
period ( Oxfordian stage), about 160 million years ago. The avialan species from this time period include ''
Anchiornis huxleyi ''Anchiornis'' is a genus of small, four-winged paravian dinosaurs, with only one known species, the type species ''Anchiornis huxleyi'', named for its similarity to modern birds. The Latin name ''Anchiornis'' derives from a Greek word meaning " ...
'' and ''
Aurornis xui ''Aurornis'' is an extinct genus of anchiornithid theropod dinosaurs from the Jurassic period of China. The genus ''Aurornis'' contains a single known species, ''Aurornis xui'' (). ''Aurornis xui'' may be the most basal ("primitive") avialan d ...
''. ''
Xiaotingia zhengi ''Xiaotingia'' is a genus of anchiornithid theropod dinosaur from Middle Jurassic or early Late Jurassic deposits of western Liaoning, China, containing a single species, ''Xiaotingia zhengi''. Discovery ''Xiaotingia'' is known from the holoty ...
'' used to be considered a member, but was later classified within the clade
Dromaeosauridae Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
. The well-known ''Archaeopteryx'' dates from slightly later Jurassic rocks (about 155 million years old) from
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
. Many of these early avialans shared unusual anatomical features that may be ancestral to modern birds, but were later lost during bird evolution. These features include enlarged claws on the second toe which may have been held clear of the ground in life, and long feathers or "hind wings" covering the hind limbs and feet, which may have been used in aerial maneuvering. It is also thought that early avialans were either cranially akinetic or had otherwise limited
cranial kinesis Cranial kinesis is the term for significant movement of skull bones relative to each other in addition to movement at the joint between the upper and lower jaw. It is usually taken to mean relative movement between the upper jaw and the braincase. ...
. Avialans diversified into a wide variety of forms during the
Cretaceous Period The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of t ...
. Many groups retained primitive characteristics, such as clawed wings and teeth, though the latter were lost independently in a number of avialan groups, including modern birds (Aves). While the earliest forms, such as ''Archaeopteryx'' and '' Shenzhouraptor'', retained the long bony tails of their ancestors, the tails of more advanced avialans were shortened with the advent of the
pygostyle Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compone ...
bone in the group Pygostylia. In the late Cretaceous, around 95 million years ago, the ancestor of all modern birds also evolved a better sense of smell. The following cladogram is based on the analysis by Hartman ''et al''. (2019), which found flight likely evolved five separate times among paravian dinosaurs, two of those among Avialae (in Scansoriopterygids and other avialans). ''Archaeopteryx'' and "anchiornithids" were placed in Deinonychosauria, Avialae's sister group. In a study conducted in 2020, ''Archaeopteryx'' was recovered as an avialan.


See also

*
Bird flight Bird flight is the primary mode of locomotion used by most bird species in which birds take off and fly. Flight assists birds with feeding, breeding, avoiding predators, and migrating. Bird flight is one of the most complex forms of locomo ...
*
Feathered dinosaur A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been s ...


References

{{Taxonbar, from=Q782930 Oxfordian first appearances Extant Late Jurassic first appearances Taxa named by Jacques Gauthier