Caudipteryx Scale
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''Caudipteryx'' (which means "tail feather") is a
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 ...
of
peacock Peafowl is a common name for three bird species in the genera ''Pavo (genus), Pavo'' and ''Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female pea ...
-sized
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s that lived in the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
age of 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 ...
(about 124.6 million
year A year or annus is the orbital period of a planetary body, for example, the Earth, moving in its orbit around the Sun. Due to the Earth's axial tilt, the course of a year sees the passing of the seasons, marked by change in weather, the hou ...
s ago). They were
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
ed and extremely
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 ...
like in their overall appearance, to the point that some
paleontologists Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
think it was 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 ...
. Two
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 ...
have been described: ''C. zoui'' (the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
), in 1998, and ''C. dongi'', in 2000. ''Caudipteryx''
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s were first discovered in the
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 ...
of the Sihetun area of
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
Province, northeastern
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 ...
in 1997.


Description

''Caudipteryx'', like many other
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, Oviraptoros ...
ns, has a mix of reptile- and bird-like anatomical features.Witmer, L.M. (2005). “The Debate on Avian Ancestry; Phylogeny, Function and Fossils”, ''Mesozoic Birds: Above the Heads of Dinosaurs'' : 3–30. It had a short, boxy
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
with a beak-like snout that retained only a few tapered teeth in the front of the upper jaw. It had a stout trunk, long legs and was probably a swift runner. Its short tail was stiffened toward the tip, with few vertebrae, like in birds and other
oviraptorosaur Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period (geology), Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like s ...
s. It has a primitive pelvis and shoulder, and primitive skull details in the quadratojugal, squamosal, quadrate, jugal, and mandibular fenestra (in the cheek, jaw, and jaw joint). It has a hand skeleton with a reduced third finger, like that of early birds and the oviraptorid ''
Heyuannia ''Heyuannia'' ("from Heyuan") is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous epoch, in what is now China and Mongolia. It was the first oviraptorid found in China; most others were found in neighbouring Mongoli ...
''.Osmolska, H., Currie, P.J., and Barsbold, R. (2004). "Oviraptorosauria." In Weishampel, Dodson, Osmolska (eds.) ''The Dinosauria'', second edition. University of California Press, 2004. ''Caudipteryx'' had uncinate processes on the ribs, birdlike teeth, a first toe which may or may not be partially reversed and overall body proportions that are comparable to those of modern flightless birds.PDF
/ref> The size of ''C. zoui'' is estimated at 65 cm (2.1 ft) in length and 2.2 kg (4.8 lbs) in body mass, while ''C. dongi'' is thought to have been slightly larger at 80 cm (2.6 ft) in length and 2.3 kg (5 lbs) in body mass.


Paleobiology


Feathers

The hands of ''Caudipteryx'' supported symmetrical, pennaceous feathers that had vanes and barbs, measuring between long. The primary feathers were arranged in a wing-like fan along the second finger, just like primary feathers of birds and other
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, Oviraptoros ...
ns. No fossil of ''Caudipteryx zoui'' preserves any secondary feathers attached to the forearms, as found in
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
s, ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
,'' and modern birds. Either these arm feathers are not preserved, or they were not present on ''Caudipteryx'' in life. An additional fan of feathers existed on its tail. The shortness and symmetry of the feathers, and the shortness of the arms relative to the body size, indicate that ''Caudipteryx'' was
flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...
less. Its body was covered in black feathers, with a visible banding pattern preserved on tail feathers.


Diet

''Caudipteryx'' is thought to have been an
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
. In at least two specimens of ''Caudipteryx'' (NGMC 97 4 A and NGMC 97 9 A),
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In othe ...
s are preserved. As in some herbivorous dinosaurs, the
avialan Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used ...
''
Sapeornis ''Sapeornis'' is a monotypic genus of avialan which lived during the early Cretaceous period ( late Barremian to early Aptian, roughly 125-120 mya). ''Sapeornis'' contains only one species, ''Sapeornis chaoyangensis''. Description ''Sapeornis' ...
'', and modern birds, these gastroliths remain in the position where the animals'
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...
s would have been.


Paleoecology

All ''Caudipteryx''
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s were recovered from the
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 ...
in
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
,
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 ...
. Specifically, they come from a small area of the Jianshangou bed, near the town of
Zhangjiakou Zhangjiakou (; ; ) also known as Kalgan and by several other names, is a prefecture-level city in northwestern Hebei province in Northern China, bordering Beijing to the southeast, Inner Mongolia to the north and west, and Shanxi to the southw ...
. They appear to have been fairly common, though isolated to this small region. The specific region in which ''Caudipteryx'' lived was home to the other feathered dinosaurs ''
Dilong ''Dilong'' (; lit. "earth dragon") is a Chinese dragon name that is also used to mean "earthworm" in traditional Chinese medicine and ''Geosaurus'' in zoological nomenclature. Dragon In Chinese mythology, ''dilong'' "earth dragon" is one of m ...
'' and ''
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 (late Barremian) of the Yixian Formation in what is now China. It ...
''.


Implications

The discovery of ''Caudipteryx'' has led to many intensive studies and debate over the relationship of birds and dinosaurs. The possible positions in the debate can be summarized as follows: ''Caudipteryx'' is either a member of the
Oviraptorosauria Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wit ...
, or a bird, or both, and birds are either dinosaurs or they are not. (See the rest of this section and Phylogeny, below). Because ''Caudipteryx'' has clear and unambiguously pennaceous feathers, like modern birds, and because several cladistic analyses have consistently recovered it as a non-avian oviraptorid dinosaur, it provided, at the time of its description, the clearest and most succinct evidence that birds evolved from dinosaurs.
Lawrence Witmer Lawrence M. Witmer (born October 10, 1959, at Rochester, New York) is an American paleontologist and paleobiologist. He is a Professor of Anatomy and a Chang Ying-Chien Professor of Paleontology at the Department of Biomedical Sciences at the Her ...
stated: “The presence of unambiguous feathers in an unambiguously non-avian theropod has the rhetorical impact of an atomic bomb, rendering any doubt about the theropod relationships of birds ludicrous.” However, not all scientists agreed that ''Caudipteryx'' was unambiguously non-avian, and some of them continued to doubt that general consensus. Paleornithologist Alan Feduccia sees ''Caudipteryx'' as a flightless bird evolving from earlier
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
ian dinosaurs rather than from late theropods.Feduccia, A. (1999). ''The Origin and Evolution of Birds''. 420 pp. Yale University Press, New Haven. . Jones ''et al.'' (2000) found that ''Caudipteryx'' was a bird based on a mathematical comparison of the body proportions of flightless birds and non-avian theropods. Dyke and Norell (2005) criticized this result for flaws in their mathematical methods, and produced results of their own which supported the opposite conclusion. Other researchers not normally involved in the debate over bird origins, such as Zhou, acknowledged that the true affinities of ''Caudipteryx'' were debatable.


Classification

The consensus view, based on several
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analyses, is that ''Caudipteryx'' is a basal (primitive) member of the
Oviraptorosauria Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wit ...
, and the oviraptorosaurians are non-avian theropod dinosaurs. ''
Incisivosaurus ''Incisivosaurus'' ("incisor lizard") is a genus of small, probably herbivorous theropod dinosaurs from the early Cretaceous Period of what is now the People's Republic of China. The first specimen to be described (by Xu ''et al.'' in 2002), IVPP ...
'' is the only oviraptorosaur that is more primitive. Halszka Osmólska ''et al.'' (2004) ran a cladistic analysis that came to a different conclusion. They found that the most birdlike features of oviraptorids actually place the whole clade within Aves itself, meaning that ''Caudipteryx'' is both an oviraptorid and a bird. In their analysis, birds evolved from more primitive theropods, and one lineage of birds became flightless, re-evolved some primitive features, and gave rise to the oviraptorids. This analysis was persuasive enough to be included in paleontological textbooks like Benton's ''Vertebrate Paleontology'' (2005).Osmólska, Halszka, Currie, Philip J., Barsbold, Rinchen (2004) ''The Dinosauria'' Weishampel, Dodson, Osmolska. "Chapter 8 Oviraptorosauria" University of California Press. The view that ''Caudipteryx'' was secondarily flightless is also preferred by
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
,Paul, G.S. (2002). ''Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds.'' Johns Hopkins University Press, Baltimore. ''et al.'',Lü, J., Dong, Z., Azuma, Y., Barsbold, R., and Tomida, Y. (2002). "Oviraptorosaurs compared to birds." In Zhou, Z., and Zhang, F. (eds.), ''Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution'', 175–189. Beijing Science Press. and Maryańska ''et al.'' Others, such as Stephen Czerkas and
Larry Martin Larry Dean Martin (December 8, 1943 – March 9, 2013) was an American vertebrate paleontologist and curator of the Natural History Museum and Biodiversity Research Center at the University of Kansas. Among Martin's work is research on the Trias ...
have concluded that ''Caudipteryx'' is not a
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur at all. They believe that ''Caudipteryx'', like all
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, Oviraptoros ...
ns, is a flightless bird, and that birds evolved from non-dinosaurian archosaurs.


See also

*
Timeline of oviraptorosaur research This timeline of oviraptorosaur research is a chronological listing of events in the history of paleontology focused on the oviraptorosaurs, a group of beaked, bird-like theropod dinosaurs. The early history of oviraptorosaur paleontology is cha ...


References


External links

* Australian Museum
Chinese Dinosaurs: ''Caudipteryx zoui''
Retrieved 2007-FEB-19. * Natural History Museum of Los Angeles County

Retrieved 2007-FEB-19. * Research Casting International
Life-size model of ''Caudipteryx zoui''
Retrieved 2007-FEB-19. * CNN
Scientists: Fossils prove that birds evolved from dinosaurs
Retrieved 2007-AUG-10 {{Taxonbar, from=Q131069 Oviraptorosaurs Feathered dinosaurs Early Cretaceous dinosaurs of Asia Fossils of China Fossil taxa described in 1998 Taxa named by Philip J. Currie Taxa named by Mark Norell Yixian fauna