Yixianornis Temporal range: Early Cretaceous, | |
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Type specimen, Paleozoological Museum of China | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Avialae |
Family: | † Songlingornithidae |
Genus: | † Yixianornis |
Species: | †Y. grabaui |
Binomial name | |
†Yixianornis grabaui Zhou & Zhang, 2001 | |
Yixianornis (meaning "Yixian Formation bird" [1] ) is a bird genus from the early Cretaceous period. Its remains have been found in the Jiufotang Formation at Chaoyang (People's Republic of China) dated to the early Aptian age, around 120 million years ago. [2] Only one species, Yixianornis grabaui, is known at present. The specific name, grabaui, is named after American paleontologist Amadeus William Grabau, who surveyed China in the early 20th century.
The type specimen (and only specimen found to date) of Yixianornis, catalog number IVPP V12631 in the collections of the Institute of Vertebrate Paleontology and Paleoanthropology, is one of the most well-preserved bird fossils known from the Jehol group. It is nearly complete and, unlike many other fossils, the bones are mostly uncrushed and were not split in half when the stone slabs were initially separated. It is also one of the few known Mesozoic ornithuran bird specimens that preserve clear impressions of the wing and tail feathers. [3]
Yixianornis was moderately sized compared to other yanornithiformes, at about the size of a chicken. IVPP V12631 was measured approximately 19 to 20 centimetres (7.5 to 7.9 in) long (excluding feathers), [1] [4] 11.5 centimetres (4.5 in) tall at the hips, [4] with a wingspan of about 40 cm (16 in). [3] Its body weight has been estimated to have been 200 to 350 grams (0.44 to 0.77 lb). [4] [5] Yixianornis was grossly similar to its close relatives Yanornis and Songlingornis . All three had teeth, though in Yixianornis the tips of the jaws were toothless and pockmarked with small pits and grooves, possibly indicating the presence of a beak. The teeth were small and peg-like, and lacked any serrations. The lower jaw was thin and delicate. [3]
The breastbone bore a strong keel for the attachment of flight muscles, and contained a distinct opening or fenestra, a unique characteristic of yanornithiformes. The upper and lower arm were about the same length. Like other ornithurines, Yixianornis had a highly fused hand, with many wrist bones joined that are free in more primitive birds. [3]
The hips and hind limbs preserved more primitive features than the forelimbs, supporting the idea that the modern adaptations in the locomotion seen in modern birds evolved after the many specializations needed for flight. The hallux, or reversed, perching toe of modern birds, is noticeably pointed backward in the fossil. However, there is no clear evidence that the bone itself was twisted, which would indicate that the toe was permanently reversed. [3]
Yixianornis differed from related birds only in small skeletal characteristics. For example, the teeth in the lower jaw of Yixianornis were confined to a smaller space, while they lined more of the jaw in related species. The wishbone (furcula) of Yixianornis was more narrow than in its relatives, and its shoulder blade was much shorter, only half the length of the upper arm bone (humerus). [3]
The flight feathers of Yixianornis are well preserved in the only known specimen. There were about five primary feathers on each wing, the longest measuring about 6.7 cm (2.6 in). Unlike in many more basal species, the feathers did not become narrower or more pointed at the tip. Rather, the feathers were very broad and rounded. There were eight tail feathers, up to 9.2 cm (3.6 in) long, anchored to a pygostyle and rectrical bulb (see below) as in modern birds. [3]
The feathers of Yixianornis are unique among those preserved in other Mesozoic bird specimens, and have allowed scientists to infer its probable lifestyle. The wings were broad and rounded, with similarly large, rounded feathers. The tail feathers were arranged in a graduated series, with the outer feathers anchored closer to the base of the tail. This would have given the tail a slightly rounded silhouette. Additionally, carbonized tissue impressions show that it had a rectrical bulb, the muscles around the tail which allow the tail feathers to be fanned out during flight, and retraced when at rest. [3]
These adaptations are rare among Mesozoic birds, many of which are known to have had long, pointed wings and very few, if any, long tail feathers. The rectrical bulb and plough-shaped pygostyle allowing for tail fanning is also a unique characteristic of ornithurine birds, of which Yixianornis is among the earliest known. [3] In a 2006 study, Julia Clarke, Zhou Zhonghe and Zhang Fucheng found that the ability to fan the tail, along with the broad wings, show that it probably preferred environments with dense vegetation, where high maneuverability in flight would be necessary. [3]
Comparisons between the scleral rings of Yixianornis and modern birds and reptiles indicate that it may have been diurnal, similar to most modern birds. [6]
It was a close relative of Yanornis and together with this and Songlingornis forms a clade of early modern birds. Clarke et al. found that Yixianornis was the most primitive bird to display an essentially modern pygostyle and fan of tail feathers. [7] Later, an enantiornithine called Shanweiniao was found to have a fan tail as well, though it may have evolved independently of modern birds. [8]
Confuciusornis is a genus of basal crow-sized avialan from the Early Cretaceous Period of the Yixian and Jiufotang Formations of China, dating from 125 to 120 million years ago. Like modern birds, Confuciusornis had a toothless beak, but closer and later 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 was thought to be the oldest known bird to have a beak, though this title now belongs to an earlier relative Eoconfuciusornis. It was named after the Chinese moral philosopher Confucius. Confuciusornis is one of the most abundant vertebrates found in the Yixian Formation, and several hundred complete specimens have been found.
The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans, the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and clawed fingers on each wing, but otherwise looked much like modern birds externally. Over eighty species of Enantiornithes have been named, but some names represent only single bones, so it is likely that not all are valid. The Enantiornithes became extinct at the Cretaceous–Paleogene boundary, along with Hesperornithes and all other non-avian dinosaurs.
Beipiaosaurus is a genus of therizinosauroid theropod dinosaurs that lived in Asia during the Early Cretaceous in the Yixian Formation. The first remains were found in 1996 and formally described in 1999. Before the discovery of Yutyrannus, Beipiaosaurus were among the heaviest dinosaurs known from direct evidence to be feathered. Beipiaosaurus is known from three reported specimens. Numerous impressions of feather structures were preserved that allowed researchers to determine the feathering color which turned out to be brownish.
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 component of the uropygium, a structure colloquially known as the bishop's nose, parson's nose, pope's nose, or sultan's nose. This is the fleshy protuberance visible at the posterior end of a bird that has been dressed for cooking. It has a swollen appearance because it also contains the uropygial gland that produces preen oil.
Yanornis is an extinct genus of fish-eating Early Cretaceous birds. Two species have been described, both from Liaoning province, China: Yanornis martini, based on several fossils found in the 120-million-year-old Jiufotang Formation at Chaoyang, and Yanornis guozhangi, from the 124-million-year-old Yixian Formation.
Longipteryx is a genus of prehistoric bird which lived during the Early Cretaceous. It contains a single species, Longipteryx chaoyangensis. Its remains have been recovered from the Jiufotang Formation at Chaoyang in Liaoning Province, China. Apart from the holotype IVPP V 12325 - a fine and nearly complete skeleton — another entire skeleton and some isolated bones are known to date.
Sapeornis is a monotypic genus of avialan dinosaurs which lived during the early Cretaceous period. Sapeornis contains only one species, Sapeornis chaoyangensis.
Hongshanornis is a genus of ornithuromorph birds known from early Cretaceous lake deposits of the Yixian Formation, Inner Mongolia, China. The holotype specimen, recovered in 2005, is currently held by the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. It was found in the Jianshangou fossil beds, dated to 124.6 million years ago. Three additional specimens have been reported, though only one of those has been definitively identified as belonging to Hongshanornis. This latter specimen was found in the Dawangzhangzi fossil beds, which are about 122 million years old.
Archaeorhynchus is a genus of beaked avialan stem-birds from the early Cretaceous period. A fossil of its only known species, Archaeorhynchus spathula, was first reported in 2005 by Zhou & Zhang to have been found in Yixian Formation rocks at Yixian, Liaoning province, China, showing a well-preserved and essentially complete skeleton. Two more complete specimens were found in Lower Cretaceous deposits of Jianchang, Liaoning, northeastern China, preserving new anatomical information. These deposits are 120 million years old, whereas the original specimen was 125 million years old, meaning the age range for this species is 125-120Ma.
Songlingornis is an extinct genus of ornithuromorph dinosaurs from the Early Cretaceous. Its fossils have been found in the Jiufotang Formation of Liaoning (PRC). The age of these rocks is somewhat disputed, but probably around the early Aptian, 125-120 million years ago. Only one species, Songlingornis linghensis, is known at present.
Pygostylia is a group of avialans which includes the Confuciusornithidae and all of the more advanced species, the Ornithothoraces.
Protopteryx is an extinct bird and possibly the basalmost enantiornithean, from the Cretaceous period. The type species is P. fengningensis. It was first discovered in the Sichakou Member of the Yixian Formation or Huajiying Formation of Hebei Province, northern China, dating from 131 Ma ago. Protopteryx has been found in the Daibeigou formation, as well. The name Protopteryx means "primitive feather": "proto-" meaning "the first of" and "-pteryx" meaning "feather" or "wing." The name comes from the fact that Protopteryx feathers are more primitive than those of modern birds, such as the two elongated tail feathers that lack barbs and rami.
Similicaudipteryx, meaning "similar to Caudipteryx", is a genus of theropod dinosaur of the family Caudipteridae.
Shanweiniao is a genus of long-snouted enantiornithean birds from Early Cretaceous China. One species is known, Shanweiniao cooperorum. There is one known fossil, a slab and counterslab. The fossil is in the collection of the Dalian Natural History Museum, and has accession number DNHM D1878/1 and DNHM1878/2. It was collected from the Lower Cretaceous Dawangzhengzi Beds, middle Yixian Formation, from Lingyuan in the Liaoning Province, China.
Songlingornithidae is a family of basal euornithean dinosaurs from the Early Cretaceous of China. All known specimens come from the Jiufotang Formation and the Yixian Formation, dating to the early Barremian and Aptian ages, 125–120 million years ago.
Yanornithiformes is an order of ornithuromorph dinosaurs from the early Cretaceous Period of China. All known specimens come from the Yixian Formation and Jiufotang Formation, dating to the early Aptian age, 124.6 to 120 million years ago.
Longipterygidae is a family of early enantiornithean avialans from the Early Cretaceous epoch of China. All known specimens come from the Jiufotang Formation and Yixian Formation, dating to the early Aptian age, 125-120 million years ago.
Dinosaur coloration is generally one of the unknowns in the field of paleontology, as skin pigmentation is nearly always lost during the fossilization process. However, recent studies of feathered dinosaurs and skin impressions have shown the colour of some species can be inferred through the use of melanosomes, the colour-determining pigments within the feathers.
Euornithes is a natural group which includes the most recent common ancestor of all avialans closer to modern birds than to Sinornis.
Chiappeavis is a genus of enantiornithean bird from Early Cretaceous of northeastern China. The only species is Chiappeavis magnapremaxillo. Chiappeavis is classified within the family Pengornithidae. It is known from a single, almost complete skeleton including feather impressions discovered in the Jiufotang Formation of the Jehol Group. Long feathers formed a fan-shaped tail that was probably employed in flight.
Winter 2011 Appendix