Yixianornis
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''Yixianornis'' (meaning "
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
bird") is a
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 lightweigh ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
from the early
Cretaceous 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 th ...
period. Its remains have been found in the
Jiufotang Formation The Jiufotang Formation (Chinese: 九佛堂组, pinyin: ''jiǔfótáng zǔ'') is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see ...
at Chaoyang (
People's Republic of China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
) dated to the early
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
age, around 120 million years ago. Only one
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''Yixianornis grabaui'', is known at present. The specific name, ''grabaui'', is named after American paleontologist
Amadeus William Grabau Amadeus William Grabau (January 9, 1870 – March 20, 1946) was an American geologist who worked in China. Biography Grabau's grandfather, J.A.A. Grabau, led a group of dissident Lutheran immigrants from Germany to Buffalo, New York. His educa ...
, who surveyed China in the early 20th century.


Description

The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
(and only specimen found to date) of ''Yixianornis'', catalog number IVPP V12631 in the collections of the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
, is one of the most well-preserved bird fossils known from the
Jehol group The Jehol Biota includes all the living organisms – the ecosystem – of northeastern China between 133 and 120 million years ago. This is the Lower Cretaceous ecosystem which left fossils in the Yixian Formation and Jiufotang Formation. These ...
. 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 Ornithurae (meaning "bird tails" in Greek) is a natural group which includes the common ancestor of ''Ichthyornis'', ''Hesperornis'', and all modern birds as well as all other descendants of that common ancestor. Classification Ernst Haeckel co ...
bird specimens that preserve clear impressions of the wing and tail feathers. ''Yixianornis'' was moderately sized compared to other yanornithiformes, at about the size of a chicken ( long excluding feathers),Holtz, Thomas R. Jr. (2008) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages'
Supplementary Information
with a wingspan of about . ''Yixianornis'' was grossly similar to its close relatives ''
Yanornis ''Yanornis'' () is an extinct genus of fish-eating Early Cretaceous birds. Two species have been described, both from Liaoning province, People's Republic of China, China: ''Yanornis martini'', based on several fossils found in the 120-million-ye ...
'' and ''
Songlingornis ''Songlingornis'' is a prehistoric bird genus 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 ...
''. 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. 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. 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. ''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''). The flight feathers of ''Yixianornis'' are well preserved in the only known specimen. There were about five
primary feather Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
s on each wing, the longest measuring about . 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 long, anchored to a
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 compon ...
and ''rectrical bulb'' (see below) as in modern birds.


Biology and ecology

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. 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. In a 2006 study,
Julia Clarke Julia Allison Clarke is an American paleontologist and evolutionary biologist who studies the evolution of birds and the dinosaurs most closely related to living birds. She is the John A. Wilson Professor in Vertebrate Paleontology in the Jackson ...
, 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. Comparisons between the
scleral ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
s of ''Yixianornis'' and modern birds and reptiles indicate that it may have been diurnal, similar to most modern birds.


Classification

It was a close relative of ''
Yanornis ''Yanornis'' () is an extinct genus of fish-eating Early Cretaceous birds. Two species have been described, both from Liaoning province, People's Republic of China, China: ''Yanornis martini'', based on several fossils found in the 120-million-ye ...
'' and together with this and ''
Songlingornis ''Songlingornis'' is a prehistoric bird genus 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 ...
'' forms 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 term, ...
of early modern birds. Clarke et al. found that ''Yixianornis'' was the most primitive bird to display an essentially modern
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 compon ...
and fan of tail feathers. Later, an enantiornithine called ''
Shanweiniao ''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 Nat ...
'' was found to have a fan tail as well, though it may have evolved independently of modern birds.Jingmai K. O’connor, Xuri Wang, Luis M. Chiappe, Chunling Gao, Qingjin Meng,Xiaodong Cheng, And Jinyuan Liu (2009). "Phylogenetic support for a specialized clade of Cretaceous enantiornithine birds with information from a new species" ''Journal of Vertebrate Paleontology'' 29(1):188–204, March 2009# 2009 by the Society of Vertebrate Paleontology


References

* (2004): The Theropod Database
Phylogeny of taxa
Retrieved 2013-MAR-02. {{Taxonbar, from=Q310304 Early Cretaceous birds of Asia Bird genera Songlingornithids Fossil taxa described in 2001