Thescientific question of within which larger group of animals birds evolved has traditionally been called the "origin of birds". The present scientific consensus is that birds are a group of maniraptoran theropod dinosaurs that originated during the Mesozoic Era.
A close relationship between birds and dinosaurs was first proposed in the nineteenth century after the discovery of the primitive bird Archaeopteryx in Germany. Birds and extinct non-avian dinosaurs share many unique skeletal traits.[1] Moreover, fossils of more than thirty species of non-avian dinosaur with preserved feathers have been collected. There are even very small dinosaurs, such as Microraptor and Anchiornis, which have long, vaned arm and leg feathers forming wings. The Jurassic basal avialan Pedopenna also shows these long foot feathers. Paleontologist Lawrence Witmer concluded in 2009 that this evidence is sufficient to demonstrate that avian evolution went through a four-winged stage.[2] Fossil evidence also demonstrates that birds and dinosaurs shared features such as hollow, pneumatized bones, gastroliths in the digestive system, nest-building, and brooding behaviors.
Although the origin of birds has historically been a contentious topic within evolutionary biology, only a few scientists still dispute the dinosaurian origin of birds, suggesting descent from other types of archosaurian reptiles. Within the consensus that supports dinosaurian ancestry, the exact sequence of evolutionary events that gave rise to the early birds within maniraptoran theropods is disputed. The origin of bird flight is a separate but related question for which there are also several proposed answers.
Scientific investigation into the origin of birds began shortly after the 1859 publication of Charles Darwin's On the Origin of Species.[3] In 1860, a fossilized feather was discovered in Germany's Late Jurassic Solnhofen limestone. Christian Erich Hermann von Meyer described this feather as Archaeopteryx lithographica the next year.[4] Richard Owen described a nearly complete skeleton in 1863, recognizing it as a bird despite many features reminiscent of reptiles, including clawed forelimbs and a long, bony tail.[5]
Biologist Thomas Henry Huxley, known as "Darwin's Bulldog" for his tenacious support of the new theory of evolution by means of natural selection, almost immediately seized upon Archaeopteryx as a transitional fossil between birds and reptiles. Starting in 1868, and following earlier suggestions by Carl Gegenbaur,[6] and Edward Drinker Cope,[7] Huxley made detailed comparisons of Archaeopteryx with various prehistoric reptiles and found that it was most similar to dinosaurs like Hypsilophodon and Compsognathus.[8][9] The discovery in the late 1870s of the iconic "Berlin specimen" of Archaeopteryx, complete with a set of reptilian teeth, provided further evidence. Like Cope, Huxley proposed an evolutionary relationship between birds and dinosaurs. Although Huxley was opposed by the very influential Owen, his conclusions were accepted by many biologists, including Baron Franz Nopcsa,[10] while others, notably Harry Seeley,[11] argued that the similarities were due to convergent evolution.
A turning point came in the early twentieth century with the writings of Gerhard Heilmann of Denmark. An artist by trade, Heilmann had a scholarly interest in birds and from 1913 to 1916, expanding on earlier work by Othenio Abel,[12] published the results of his research in several parts, dealing with the anatomy, embryology, behavior, paleontology, and evolution of birds.[13] His work, originally written in Danish as Vor Nuvaerende Viden om Fuglenes Afstamning, was compiled, translated into English, and published in 1926 as The Origin of Birds.
Like Huxley, Heilmann compared Archaeopteryx and other birds to an exhaustive list of prehistoric reptiles, and also came to the conclusion that theropod dinosaurs like Compsognathus were the most similar. However, Heilmann noted that birds had clavicles (collar bones) fused to form a bone called the furcula ("wishbone"), and while clavicles were known in more primitive reptiles, they had not yet been recognized in dinosaurs. Since he was a firm believer in an interpretation of Dollo's law that stated that evolution was not "reversible", Heilmann could not accept that clavicles were lost in dinosaurs and re-evolved in birds. He was therefore forced to rule out dinosaurs as bird ancestors and ascribe all of their similarities to convergent evolution. Heilmann stated that bird ancestors would instead be found among the more primitive "thecodont" grade of reptiles.[14] Heilmann's extremely thorough approach ensured that his book became a classic in the field, and its conclusions on bird origins, as with most other topics, were accepted by nearly all evolutionary biologists for the next four decades.[15]
Clavicles are relatively delicate bones and therefore in danger of being destroyed or at least damaged beyond recognition. Nevertheless, some fossil theropod clavicles had actually been excavated before Heilmann wrote his book, but these had been misidentified.[16]The absence of clavicles in dinosaurs became the orthodox view despite the discovery of clavicles in the primitive theropod Segisaurus in 1936.[17] The next report of clavicles in a dinosaur was in a Russian article in 1983.[18]
Contrary to what Heilmann believed, paleontologists now accept that clavicles and in most cases furculae are a standard feature not just of theropods but of saurischian dinosaurs. Up to late 2007 ossified furculae (i.e. made of bone rather than cartilage) have been found in all types of theropods except the most basal ones, Eoraptor and Herrerasaurus.[19] The original report of a furcula in the primitive theropod Segisaurus (1936) was confirmed by a re-examination in 2005.[20] Joined, furcula-like clavicles have also been found in Massospondylus, an Early Jurassic sauropodomorph.[21]
The tide began to turn against the 'thecodont' hypothesis after the 1964 discovery of a new theropod dinosaur in Montana. In 1969, this dinosaur was described and named Deinonychus by John Ostrom of Yale University.[22] The next year, Ostrom redescribed a specimen of Pterodactylus in the Dutch Teylers Museum as another skeleton of Archaeopteryx.[23] The specimen consisted mainly of a single wing and its description made Ostrom aware of the similarities between the wrists of Archaeopteryx and Deinonychus.[24]
In 1972, British paleontologist Alick Walker hypothesized that birds arose not from 'thecodonts' but from crocodile ancestors like Sphenosuchus.[25] Ostrom's work with both theropods and early birds led him to respond with a series of publications in the mid-1970s in which he laid out the many similarities between birds and theropod dinosaurs, resurrecting the ideas first put forth by Huxley over a century before.[26][27][28] Ostrom's recognition of the dinosaurian ancestry of birds, along with other new ideas about dinosaur metabolism,[29] activity levels, and parental care,[30] began what is known as the dinosaur renaissance, which began in the 1960s and, according to some, continues to this day.[31]
Ostrom's revelations also coincided with the increasing adoption of phylogenetic systematics (cladistics), which began in the 1960s with the work of Willi Hennig.[32] Cladistics is an exact method of arranging species based strictly on their evolutionary relationships, which are calculated by determining the evolutionary tree implying the least number of changes in their anatomical characteristics. In the 1980s, cladistic methodology was applied to dinosaur phylogeny for the first time by Jacques Gauthier and others, showing unequivocally that birds were a derived group of theropod dinosaurs.[33] Early analyses suggested that dromaeosaurid theropods like Deinonychus were particularly closely related to birds, a result that has been corroborated many times since.[34][35]
The early 1990s saw the discovery of spectacularly preserved bird fossils in several Early Cretaceous geological formations in the northeastern Chinese province of Liaoning.[36][37] In 1996, Chinese paleontologists described Sinosauropteryx as a new genus of bird from the Yixian Formation,[38] but this animal was quickly recognized as a more basal theropod dinosaur closely related to Compsognathus. Surprisingly, its body was covered by long filamentous structures. These were dubbed 'protofeathers' and considered homologous with the more advanced feathers of birds,[39] although some scientists disagree with this assessment.[40] Chinese and North American scientists described Caudipteryx and Protarchaeopteryx soon after. Based on skeletal features, these animals were non-avian dinosaurs, but their remains bore fully formed feathers closely resembling those of birds.[41] "Archaeoraptor", described without peer review in a 1999 issue of National Geographic,[42] turned out to be a smuggled forgery,[43] but authentic remains continue to pour out of the Yixian, both legally and illegally. Feathers or "protofeathers" have been found on a wide variety of theropods in the Yixian.[44][45] The morphological gap between non-avian theropods and birds is further closed by the discoveries of extremely bird-like non-avian dinosaurs,[46] as well as non-avian dinosaur-like basal birds.[47]
Embryologists and some paleontologists who oppose the bird-dinosaur link have long numbered the digits of birds II-III-IV on the basis of multiple studies of the development in the egg.[48] This is based on the fact that in most amniotes, the first digit to form in a 5-fingered hand is digit IV, which develops a primary axis. Therefore, embryologists have identified the primary axis in birds as digit IV, and the surviving digits as II-III-IV. The fossils of advanced theropod (Tetanurae) hands appear to have the digits I-II-III (some genera within Avetheropoda also have a reduced digit IV[49]). If this is true, then the II-III-IV development of digits in birds is an indication against theropod (dinosaur) ancestry. However, with no ontogenical (developmental) basis to definitively state which digits are which on a theropod hand (because no non-avian theropods can be observed growing and developing today), the labelling of the theropod hand is not absolutely conclusive.[citation needed]
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