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Confuciusornis is a genus of primitive crow-sized birds from the Early Cretaceous Yixian and Jiufotang Formations of China, dating from 125 to 120 million years ago. Like modern birds, Confuciusornis had a toothless beak, but close relatives of modern birds such as Hesperornis and Ichthyornis were toothed, indicating that the loss of teeth occurred convergently in Confuciusornis and living birds. It is the oldest known bird to have a beak.[1] It was named after the Chinese moral philosopher Confucius (551–479 BC). Confuciusornis is one of the most abundant vertebrates found in the Yixian Formation, and several hundred complete, articulated specimens have been found.[

Discovery and namingEdit

In November 1993 Chinese paleontologists Zhou Zhonge, Hu Yoaming and Hou Lianhai, of the Institute of Vertebrate Paleontology and Paleoanthropology at Beijing, were presented a bird fossil bought at a flea market in Sihetun by amateur paleontologist Zhang He. It showed a partial skeleton of a bird new to science of which even some feather remains had been preserved. In December 1993 two further specimens were acquired from a farmer, Yang Yushan. Soon afterwards, it was discovered that local farmers were in the process of collecting hundreds of specimens that were provisionally prepared by them to be illegally sold to commercial fossil dealers.[3] Many hundreds have since also been added to the official collections of Chinese institutions. In 2010 the Shandong Tianyu Museum of Nature alone possessed 536 specimens.[4] The fossils are compressed but otherwise typically very complete often showing the entire feathering of the animal.

Based on the abundant material, six species have been formally named and described: C. sanctus (the type species), C. dui, C. feducciai, C. jianchangensis, C. chuonzhous and C. suniae. The latter two are usually considered synonymous with C. sanctus.[5] Most species lived in the early Aptian 125 million year old Jianshangou Beds of the Yixian Formation, though C. jianchangnsis is found in the later (120Ma) middle Aptian Jiufotang Formation. C. sanctus is known from the former, and is one of the most common vertebrate species found in the Yixian, often present in dense concentrations. At one time forty were discovered on a surface of about 100 m². This has been explained as the result of entire flocks of birds being simultaneously killed by ash, heat or poisonous gas following the volcanic eruptions that caused the tuff stone in which the fossils were found to be deposited as lake sediments.[6]

In 1995 Zhou, Hou, Zhang and Gu Youcai named Confuciusornis sanctus. The generic name combines the philosopher Confucius with a Greek ὄρνις, (ornis), "bird". The specific name means "holy one" in Latin and is a translation of Chinese 圣贤, shèngxián, "sage", again in reference to Confucius. The holotype specimen is IVPP V10918.[7] In 1997 Hou named two species. The first was Confuciusornis chuonzhous, based on specimen IVPP V10919, originally a paratype of C. sanctus. The specific name refers to Chuanzhou, an ancient name for Beipiao. The second species was Confuciusornis suniae, based on specimen IVPP V11308. The specific name honours madam Sun, the wife of Shikuan Liang who donated the fossil to the IVPP.[8] In 1999 Hou, Zhou, Zhang, Larry Martin and Alan Feduccia named Confuciusornis dui, based on specimen IVPP V11553. The specific name again honours a donating collector: Du Wengya.[9] In 2009 Zhang Fucheng et al. named Confuciusornis feducciai, based on specimen D2454, the specific name honouring Feduccia.[10] In 2010 Li Li, Wang Jingqi and Hou Shilin named Confuciusornis jianchangensis, based on specimen PMOL-AB00114 found at Toudaoyingzi.[11]

In 2002 Hou named the genus Jinzhouornis. Luis Chiappe later concluded that this is probably a junior synonym of Confuciusornis.

DescriptionEdit

Confuciusornis was about the size of a modern pigeon, with a wingspan of up to 0.7 meters (2.3 ft), and its body weight has been estimated to have been as much as 1.5 kilograms,[12] or less than 0.2 kilograms.[13] C. feducciai was about a third longer than average specimens of C. sanctus.[10]

Confuciusornis shows a mix of basal and derived traits. It was more "advanced" or derived than Archaeopteryx in possessing a short tail with a pygostyle (a bone formed from a series of short, fused tail vertebrae) and a bony sternum, but more basal or "primitive" than modern birds in retaining large claws on the forelimbs, having a primitive skull with a closed eye-socket, and a relatively small breastbone. At first the number of basal characteristics was exaggerated: Hou assumed in 1995 that a long tail was present and mistook grooves in the jaw bones for small degenerated teeth.[14]

The skull of Confuciusornis was equipped with a pointed toothless beak. It was relatively heavy-built and immobile, incapable of the kinesis of modern birds that can raise the snout relative to the back of the skull. This immobility was caused by the presence of a triradiate postorbital separating the eye-socket from the lower temporal opening, as with more basal theropods, and the premaxillae of the snout reaching all the way to the frontals, forcing the nasals to the sides of the snout.[15]

Fossils of Confuciusornis show that it had an exceptionally large humerus (upper arm bone). Near its shoulder-end this was equipped with a prominent deltopectoral crest. Characteristically this crista deltopectoralis was with Confuciusornis pierced by an oval hole which may have reduced the bone's weight or enlarged the attachment area of the flight miscles. The furcula or wishbone, like that of Archaeopteryx, was a simple curved bar lacking a pointed process at the back, a hypocleidum. The sternum (breastbone) was relatively broad and had a low keel which was raised at the back end. This bony keel may or may not have anchored a larger, cartilaginous, keel for enlarged pectoral muscles.[15] The scapulae (shoulder blades) were fused to the strut-like coracoid bones and may have formed a solid base for the attachment of wing muscles. The orientation of the shoulder joint was sideways, instead of angled upward as in modern birds; this means that Confuciusornis was unable to lift its wings high above its back. According to a study by Phil Senter in 2006, the joint was even pointed largely downwards meaning that the humerus could not be lifted above the horizontal. This would make Confuciusornis incapable of the upstroke required for flapping flight; the same would have been true for Archaeopteryx.

The wrist of Confuciusornis shows fusion, forming a carpometacarpus. The second and third metacarpals were also partially fused but the first was unfused, however, and also the fingers could freely move relative to each other. The second metacarpal, supporting the flight feathers, was very heavily built; its finger carries a small claw. The claw of the first finger to the contrary was very large and curved, that of the third intermediate in size.[15] The formula of the finger phalanges was 2-3-4-0-0.

The pelvis was connected to a sacrum formed by seven sacral vertebrae. The pubis was strongly pointing backwards. The left and right ischia were not fused. The femur was straight; the tibia only slightly longer. The metatarsals of the foot were relatively short and fused to each other and to the lower ankle bones, forming a tarsometatarsus. A rudimentary fifth metatarsal is present. The first metatarsal was attached to the lower shaft of the second and supported a first toe or hallux, pointing to the back.[15] The formula of the toe phalanges was 2-3-4-5-0. The proportions of the toes suggest that they were used for both walking and perching, while the large claws of the thumb and third finger were probably used for climbing.

FeathersEdit

The wing feathers of Confuciusornis were long and modern in appearance. The primary wing feathers of a 0.5 kilogram individual reached 20.7 centimeters in length. The five longest primary feathers (remiges primarii) were more than 3.5 times the length of the hand and relatively longer than those of any living bird, while the secondary feathers of the lower arm were rather short by comparison. Thus, the wing shape was very unlike that of living birds, being long and narrow.[12] The primary feathers were asymmetrical to varying degree, and especially so in the outermost primaries.[15] It is unclear whether the upper arm carried tertiaries. Covert feathers are preserved covering the upper part of the wing feathers in some specimens, and some specimens have preserved the contour feathers of the body.[15] Unlike some more advanced birds, Confuciusornis lacked an alula, or "bastard wing". In modern birds this is formed by feathers anchored to the first digit of the hand, but this digit appears to have been free of feathers and independent of the body of the wing in Confuciusornis.[15] According to Dieter Stefan Peters to compensate for the lack of an alula, the third finger might have formed a separate winglet below the main wing, functioning like the flap of an aircraft.[17] Despite the relatively advanced and long wing feathers, the forearm bones lacked any indication of quill knobs (papillae ulnares), or bony attachment points for the feather ligaments.[15]

Many specimens preserve a pair of long, narrow tail feathers, which grew longer than the entire length of the rest of the body. Unlike the feathers of most modern birds, these feathers were not differentiated into a central quill and barbs for most of their length. Rather, most of the feather formed a ribbon-like sheet, about six millimetres wide. Only at the last one quarter of the feather, towards the rounded tip, does the feather become differentiated into a central shaft with interlocking barbs. Many individuals of Confuciusornis lacked even these two tail feathers, possibly due to sexual dimorphism. The rest of the tail around the pygostyle was covered in short, non-aerodynamic feather tufts similar to the contour feathers of the body, rather than the familiar feather fan of modern bird tails.

ColorationEdit

In early 2010, a group of scientists led by Zhang Fucheng examined fossils with preserved melanosomes (organelles which contain colors). By studying such fossils with an electron microscope, they found melanosomes preserved in a fossil Confuciusornis specimen, IVPP V13171.[18] The melanosomes were of two types, eumelanosomes and pheomelanosomes. This indicates that Confuciusornis had hues of grey, red/brown and black, possibly something like the modern zebra finch. It was also the first time an early bird fossil has been shown to contain preserved pheomelanosomes.

PaleobiologyEdit

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