Avialae ("bird wings") is a clade of flying dinosaurs containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to modern birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used (see below).
Archaeopteryx lithographica, from the late Jurassic Period Solnhofen Formation of Germany, is the earliest known avialan which may have had the capability of powered flight. However, several older avialans are known from the late Jurassic Tiaojishan Formation of China, dated to about 160 million years ago.
|Fossil specimen of Archaeopteryx lithographica|
Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all theropods closer to birds than to Deinonychus." A nearly identical definition, "the theropod group that includes all taxa closer to Passer than to Dromaeosaurus", was used by Agnolín and Novas (2013) for their clade Averaptora.
Additionally, beginning in the late 2000s and early 2010s, several groups of researchers began adding the genus Troodon 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 physical characteristics). Jacques Gauthier, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered wings used in flapping flight, and the birds that descended from them.
Gauthier (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. Gauthier proposed a solution, number 4 below, which is to reserve the term Aves only for the crown group, the last common ancestor of all living birds and all of its descendants. He assigned other names to the other groups.
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.
|Cladogram following the results of a phylogenetic study by Wang et al., 2016.|
The earliest known avialan fossils come from the Tiaojishan Formation of China, which has been dated to the late Jurassic period (Oxfordian stage), about 160 million years ago. The avialan species from this time period include Anchiornis huxleyi and Aurornis xui. Xiaotingia zhengi used to be considered a member, but was later classified within the clade Dromaeosauridae. The well-known early avialan, Archaeopteryx, dates from slightly later Jurassic rocks (about 155 million years old) from Germany. 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.
Avialans diversified into a wide variety of forms during the Cretaceous Period. 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 Jeholornis, retained the long bony tails of their ancestors, the tails of more advanced avialans were shortened with the advent of the pygostyle 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.
Anchiornis is a genus of small, four-winged paravian dinosaur. The genus Anchiornis contains only the type species Anchiornis huxleyi, named for its similarity to modern birds. Anchiornis fossils have been only found in the Tiaojishan Formation of Liaoning, China, in rocks dated to the Late Jurassic, about 160 million years ago. Anchiornis is known from hundreds of specimens, and given the exquisite preservation of some of these fossils, it became the first Mesozoic dinosaur species for which almost the entire life appearance could be determined, and an important source of information on the early evolution of birds. Anchiornis huxleyi translates to "T.H. Huxley's near-bird" in Greek.Anchiornithidae
Anchiornithidae ("near birds") is a family of eumaniraptorans which could be the basalmost family of birds (in the general sense) in the clade Avialae. Anchiornithids have been classified at varying positions in the maniraptoran tree, with some scientists classifying them as a distinct family, a basal subfamily of Troodontidae, members of Archaeopterygidae, or an assemblage of dinosaurs that are an evolutionary grade within Avialae or Paraves.Archaeopteryx
Archaeopteryx (), meaning "old wing" (sometimes referred to by its German name Urvogel ("original bird" or "first bird")), is a genus of bird-like dinosaurs that is transitional between non-avian feathered dinosaurs and modern birds. The name derives from the ancient Greek ἀρχαῖος (archaīos) meaning "ancient", and πτέρυξ (ptéryx), meaning "feather" or "wing". Between the late 19th century and the early 21st century, Archaeopteryx had been generally accepted by palaeontologists and popular reference books as the oldest known bird (member of the group Avialae). Older potential avialans have since been identified, including Anchiornis, Xiaotingia, and Aurornis.Archaeopteryx lived in the Late Jurassic around 150 million years ago, in what is now southern Germany during a time when Europe was an archipelago of islands in a shallow warm tropical sea, much closer to the equator than it is now. Similar in size to a Eurasian magpie, with the largest individuals possibly attaining the size of a raven, the largest species of Archaeopteryx could grow to about 0.5 m (1 ft 8 in) in length. Despite their small size, broad wings, and inferred ability to fly or glide, Archaeopteryx had more in common with other small Mesozoic dinosaurs than with modern birds. In particular, they shared the following features with the dromaeosaurids and troodontids: jaws with sharp teeth, three fingers with claws, a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggest warm-bloodedness), and various features of the skeleton.These features make Archaeopteryx a clear candidate for a transitional fossil between non-avian dinosaurs and birds. Thus, Archaeopteryx plays an important role, not only in the study of the origin of birds, but in the study of dinosaurs. It was named from a single feather in 1861, though that feather would eventually prove to be non-avian. That same year, the first complete specimen of Archaeopteryx was announced. Over the years, ten more fossils of Archaeopteryx have surfaced. Despite variation among these fossils, most experts regard all the remains that have been discovered as belonging to a single species, although this is still debated.
Most of these eleven fossils include impressions of feathers. Because these feathers are of an advanced form (flight feathers), these fossils are evidence that the evolution of feathers began before the Late Jurassic. The type specimen of Archaeopteryx was discovered just two years after Charles Darwin published On the Origin of Species. Archaeopteryx seemed to confirm Darwin's theories and has since become a key piece of evidence for the origin of birds, the transitional fossils debate, and confirmation of evolution.
In March 2018, scientists reported that Archaeopteryx was likely capable of flight, but in a manner distinct and substantially different from that of modern birds.Bird
Birds, also known as Aves or avian dinosaurs, are a group of endothermic vertebrates, characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the 5 cm (2 in) bee hummingbird to the 2.75 m (9 ft) ostrich. They rank as the world's most numerically-successful class of tetrapods, with approximately ten thousand living species, more than half of these being passerines, sometimes known as perching birds. Birds have wings which are more or less developed depending on the species; the only known groups without wings are the extinct moa and elephant birds. Wings, which evolved from forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in flightless birds, including ratites, penguins, and diverse endemic island species of birds. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming.
The fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier feathered dinosaurs within the theropod group, which are traditionally placed within the saurischian dinosaurs. The closest living relatives of birds are the crocodilians. Primitive bird-like dinosaurs that lie outside class Aves proper, in the broader group Avialae, have been found dating back to the mid-Jurassic period, around 170 million years ago. Many of these early "stem-birds", such as Archaeopteryx, retained primitive characteristics such as teeth and long bony tails. DNA-based evidence finds that birds diversified dramatically around the time of the Cretaceous–Palaeogene extinction event 66 million years ago, which killed off the pterosaurs and all the non-avian dinosaur lineages. But birds, especially those in the southern continents, survived this event and then migrated to other parts of the world while diversifying during periods of global cooling. This makes them the sole surviving dinosaurs according to cladistics.
Some birds, especially corvids and parrots, are among the most intelligent animals; several bird species make and use tools, and many social species pass on knowledge across generations, which is considered a form of culture. Many species annually migrate great distances. Birds are social, communicating with visual signals, calls, and bird songs, and participating in such social behaviours as cooperative breeding and hunting, flocking, and mobbing of predators. The vast majority of bird species are socially monogamous (referring to social living arrangement, distinct from genetic monogamy), usually for one breeding season at a time, sometimes for years, but rarely for life. Other species have breeding systems that are polygynous (arrangement of one male with many females) or, rarely, polyandrous (arrangement of one female with many males). Birds produce offspring by laying eggs which are fertilised through sexual reproduction. They are usually laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching. Some birds, such as hens, lay eggs even when not fertilised, though unfertilised eggs do not produce offspring.
Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds (poultry and game) being important sources of eggs, meat, and feathers. Songbirds, parrots, and other species are popular as pets. Guano (bird excrement) is harvested for use as a fertiliser. Birds prominently figure throughout human culture. About 120–130 species have become extinct due to human activity since the 17th century, and hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them. Recreational birdwatching is an important part of the ecotourism industry.Chongmingia
Chongmingia is a genus of primitive avialan belonging to Pygostylia that lived during the Aptian. It was found in the Jiufotang Formation in Chaoyang, China, and was described by Wang et al., 2016. The name comes from the word Chongming, referring to a Chinese mythological bird, and the specific epithet is in honor of Mr. Xiaoting Zheng.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, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaur group alive today.Most feathered dinosaurs discovered so far have been coelurosaurs. Philip J. Currie considers it likely and probable that all coelurosaurs were feathered. In the past, Coelurosauria was used to refer to all small theropods, but this classification has since been abolished.Epidexipteryx
Epidexipteryx is a genus of small paravian dinosaurs, known from one fossil specimen in the collection of the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. Epidexipteryx represents the earliest known example of ornamental feathers in the fossil record. The type specimen is catalog number IVPP V 15471. It has been reported to be a maniraptoran dinosaur from the Middle Jurassic or Upper Jurassic age Daohugou Beds of Inner Mongolia, China (about 160 or 168 mya).The specific name, Epidexipteryx hui ("Hu's display feather"), and its Chinese name Hushi Yaolong ("Hu Yaoming's dragon") were coined in memory of paleomammologist Hu Yaoming.Evolution of birds
The evolution of birds began in the Jurassic Period, with the earliest birds derived from a clade of theropoda dinosaurs named Paraves. Birds are categorized as a biological class, Aves. For more than a century, the small theropod dinosaur Archaeopteryx lithographica from the Late Jurassic period was considered to have been the earliest bird. Modern phylogenies place birds in the dinosaur clade Theropoda. According to the current consensus, Aves and a sister group, the order Crocodilia, together are the sole living members of an unranked "reptile" clade, the Archosauria. Four distinct lineages of bird survived the Cretaceous–Paleogene extinction event 66 million years ago, giving rise to ostriches and relatives (Paleognathae), ducks and relatives (Anseriformes), ground-living fowl (Galliformes), and "modern birds" (Neoaves).
Phylogenetically, Aves is usually defined as all descendants of the most recent common ancestor of a specific modern bird species (such as the house sparrow, Passer domesticus), and either Archaeopteryx, or some prehistoric species closer to Neornithes (to avoid the problems caused by the unclear relationships of Archaeopteryx to other theropods). If the latter classification is used then the larger group is termed Avialae. Currently, the relationship between dinosaurs, Archaeopteryx, and modern birds is still under debate.Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period (see Eshanosaurus), and are regarded as surviving today as living birds.Palaeopteryx
Palaeopteryx (meaning "ancient wing") is a genus of theropod dinosaur now considered a nomen dubium. It was named and misidentified by J. A. Jensen in 1981, then redescribed by Jensen and K. Padian in 1989. At that time the binomial Palaeopteryx thomsoni was deemed invalid by Jensen. The only referred specimen is a single bone fragment (BYU 2022).
Palaeopteryx (BYU 2022) has been the subject of much confusion on the internet, in the popular scientific press, and among creationist writers. It has been described as a possible bird older than Archaeopteryx, but it cannot be clearly assigned to Avialae, and its horizon is younger than that of Archaeopteryx, though it is still from the Jurassic.
BYU 2022 is about 45 millimeters long. It was described by Jensen in 1981 as an "avian – like" proximal left tibiotarsus. It was then listed by R. E. Molnar in 1985 in a survey of the earliest known birds. Jensen and Padian reidentified it as the distal right radius of "a small deinonychosaur or bird" in 1989.
BYU 2022 was collected in the 1970s by paleontological expeditions from Brigham Young University directed by J. A. Jensen. It was found in Late Jurassic deposits in the "Dry Mesa" quarry on the Uncompahgre Upwarp in western Colorado (Brushy Basin Member, Morrison Formation). It was found among mixed fossil remains that included pterosaur and dinosaur material. One notable specimen found with it is the right femur of a derived maniraptoran theropod (BYU 2023). BYU 2023 is missing the distal end and is about 63 mm long. It is probably too small to be from the same individual as BYU 2022. BYU 2023 shows apomorphies known only in advanced maniraptorans, including Microvenator, Microraptor, and Archaeopteryx.
BYU 2022 and 2023 are important because they are samples of small – bodied maniraptorans from Jurassic North America.Paraves
Paraves are a widespread group of theropod dinosaurs that originated in the Late Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and scansoriopterygids, the group also contains the avialans, among which are the over ten thousand species of living birds. Primitive members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species.Rahonavis
Rahonavis is a genus of bird-like theropods from the Late Cretaceous (Maastrichtian, about 70 mya) of what is now northwestern Madagascar. It is known from a partial skeleton (UA 8656) found by Catherine Forster and colleagues in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province. Rahonavis was a small predator, at about 70 centimetres (2.3 ft) long, with the typical Velociraptor-like raised sickle claw on the second toe.
The name Rahonavis means, approximately, "cloud menace bird", from Malagasy rahona (RA-hoo-na, "cloud" or "menace") + Latin avis "bird". The specific name, R. ostromi, was coined in honor of John Ostrom.Saurischia
Saurischia ( saw-RIS-kee-ə, meaning "reptile-hipped" from the Greek sauros (σαῦρος) meaning 'lizard' and ischion (ἴσχιον) meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia). ‘Saurischia’ translates to lizard-hipped.
In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure, though today most paleontologists classify Saurischia as an unranked clade rather than an order.Sauriurae
Sauriurae (meaning "lizard tails" in Greek) is a now-deprecated subclass of birds created by Ernst Haeckel in 1866. It was intended to include Archaeopteryx and distinguish it from all other birds then known, which he grouped in the sister-group Ornithurae ("bird tails"). The distinction Haeckel referred to in this name is that Archaeopteryx possesses a long, reptile-like tail, while all other birds known to him had short tails with few vertebrae, fused at the end into a pygostyle. The unit was not much referred to, and when Hans Friedrich Gadow in 1893 erected Archaeornithes for basically the same fossils, this became the common name for the early reptile-like grade of birds.
Ji Qiang and Larry Martin have continued to refer to the Sauriurae as a valid natural group. However, researchers like Jacques Gauthier (2001) and Julia Clarke (2002) have found that fossils found after Haeckel's time have bridged the gap between long and short-tailed Avialae. In their view, any grouping of avialans with long tails must exclude some of their descendants—making Sauriurae a paraphyletic and, thus, an invalid group under current systems of phylogenetic nomenclature.Scansoriopterygidae
Scansoriopterygidae (meaning "climbing wings") is an extinct family of climbing and gliding maniraptoran dinosaurs. Scansoriopterygids are known from five well-preserved fossils, representing four species, unearthed in the Tiaojishan Formation fossil beds (dating to the mid-late Jurassic Period) of Liaoning and Hebei, China.
Scansoriopteryx heilmanni (and its likely synonym Epidendrosaurus ninchengensis) was the first non-avian dinosaur found that had clear adaptations to an arboreal or semi-arboreal lifestyle–it is likely that they spent much of their time in trees. Both specimens showed features indicating they were juveniles, which made it difficult to determine their exact relationship to other non-avian dinosaurs and birds. It was not until the description of Epidexipteryx hui in 2008 that an adult specimen was known. In 2015, the discovery of another, larger adult specimen belonging to the species Yi qi showed that scansoriopterygids were not only climbers but also had adaptations that could have been used for gliding flight.Sinornithosaurus
Sinornithosaurus (derived from a combination of Latin and Greek, meaning 'Chinese bird-lizard') is a genus of feathered dromaeosaurid dinosaur from the early Cretaceous Period (early Aptian) of the Yixian Formation in what is now China. It was the fifth non–avian feathered dinosaur genus discovered by 1999. The original specimen was collected from the Sihetun locality of western Liaoning. It was found in the Jianshangou beds of the Yixian Formation, dated to 124.5 million years ago. Additional specimens have been found in the younger Dawangzhangzi bed, dating to around 122 million years ago.Xu Xing described Sinornithosaurus and performed a phylogenetic analysis which demonstrated that it is basal, or primitive, among the dromaeosaurs. He has also demonstrated that features of the skull and shoulder are very similar to Archaeopteryx and other Avialae. Together these two facts demonstrate that the earliest dromaeosaurs were more like birds than the later dromaeosaurs were.
Sinornithosaurus was among the smallest dromaeosaurids, with a length of about 90 centimetres (3.0 ft). In 2010, Gregory S. Paul gave higher estimations of 1.2 metres and three kilogrammes.Xiaotingia
Xiaotingia is a genus of anchiornithid theropod dinosaur from early Late Jurassic deposits of western Liaoning, China, containing a single species, Xiaotingia zhengi.Zhongornis
Zhongornis (meaning "intermediate bird") is a genus of primitive avialan that lived during the Early Cretaceous. It was found in rocks of the Yixian Formation in Lingyuan City (China), and described by Gao et al. in 2008.Zhongornis has only one described species, Zhongornis haoae. The only specimen is a fossil slab and counterslab numbered D2455/6. It is in the collection of the Dalian Natural History Museum. It is a fairly complete skeleton about eight centimeters in length. Pores in the bones and unfused sutures in the skeleton indicate that the specimen was a juvenile, but the authors believe that it was developed enough to erect a new taxon on the basis of its unique morphological characters. There are feather impressions preserved on the right hand and also probable tail feathers preserved near the left foot. Zhongornis had a beaked mouth with no teeth. The tail is proportionately short, has thirteen vertebrae, and no pygostyle. The third finger has only two phalangeal bones, unlike non - avian dinosaurs and Confuciusornis, and more like Enantiornithes and more advanced avialans. These features and a cladistic analysis indicate that Zhongornis is the sister group to all pygostylia, meaning that it is intermediate between long - tailed Avialae like Archaeopteryx and more advanced taxa like Confuciusornis.