''Caudipteryx'' (meaning "tail feather") is a
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of small
oviraptorosaur dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s that lived in
Asia
Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
during the
Early Cretaceous
The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
, around 124.6 million years 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 an exa ...
ed and extremely
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
like in their overall appearance, to the point that some
paleontologists suggested it was a
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
. Two
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
have been described: ''C. zoui'' (the
type species
In International_Code_of_Zoological_Nomenclature, 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 spe ...
), in 1998, and ''C. dongi'', in 2000.
It had a stout trunk, long legs and was probably a swift runner. The discovery of ''Caudipteryx'' has led to many intensive studies and debate over the relationship of birds and dinosaurs.
History
In 1997, several well-preserved
dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
skeletons were recovered from the Jiulongsong Member of the
Chaomidianzi Formation (now Jianshangou Bed of the
Yixian Formation
The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
), at the Sihetun locality of
Liaoning province
)
, image_skyline =
, image_alt =
, image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong
, image_map = Liaoning in China (+all claims hatched).svg
, ...
,
China
China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The
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 preserve ...
s were later described in 1998 and used as the type specimens for the new dinosaur taxa ''Caudipteryx'' and ''
Protarchaeopteryx''. ''Caudipteryx'' was erected with the type species ''C. zoui'' and the
holotype
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
is NGMC 97-4-A, a nearly complete individual preserving conspicuous feather impressions and
gastroliths. The
paratype
In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
is NGMC 97-9-A, another relatively complete individual with feather impressions. The
generic name, ''Caudipteryx'', means "tail feather", and the
specific name, ''zoui'', is in honor of
Zou Jiahua for his prominent support to the scientific community as the
vice premier of China.
Around the summer of 1988, a partially complete skeleton of ''Caudipteryx'' lacking the skull was found in sediments of the "Layer 6" of the Yixian Formation, at the Zhangjiagou locality, which is set apart from Sihetun. This specimen, IVPP V 12344, was in 2000 described and designed as the holotype for new species ''Caudipteryx dongi'', and in a similar fashion to previous specimens of the genus, it preserves exquisite traces of feather integument. The specific name ''dongi'' honors
Zhiming Dong, a Chinese
paleontologist
Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
.
Additional specimens

During 2000, Zhong-He Zhou and colleagues described two additional specimens of ''Caudipteryx'', BPM 0001 and IVPP V 12430, referred to ''C. zoui'' and ''C.'' sp. respectively. Both individuals preserve nearly complete skulls and have feather impressions.
Further analyses to IVPP V 12430 have revealed the preservation of propatagium on its left arm.
In 2021 Xiaoting Zheng and team described STM4-3 representing an articulated individual lacking the skull and tail tip, including abundant integument, gastroliths, but also a cartilage fragment that was reported to preserve
chondrocyte
Chondrocytes (, ) are the only cells found in healthy cartilage. They produce and maintain the cartilaginous matrix, which consists mainly of collagen and proteoglycans. Although the word '' chondroblast'' is commonly used to describe an immatu ...
s. The specimen was collected from outcrops of the Yixian Formation at Dapingfang Town near
Chaoyang city, west
Liaoning
)
, image_skyline =
, image_alt =
, image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong
, image_map = Liaoning in China (+all claims hatched).svg
, ...
.
Description
Size

''Caudipteryx'' was a small theropod, measuring long and weighing about based on femur length.
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, Dromaeosauridae, Troodontidae, ...
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. ]
Skull
It had a short, boxy
skull
The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate.
In the human, the skull comprises two prominent ...
with a beak-like snout that retained only a few tapered teeth in the front of the upper jaw.
Postcranial skeleton
Its short tail was stiffened toward the tip, with few vertebrae, like in birds and other
oviraptorosaurs. 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''.
[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>
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, Dromaeosauridae, Troodontidae, ...
ns. An additional fan of feathers existed on its tail. The body of ''C. zoui'' was covered in black feathers, with a visible banding pattern preserved on tail feathers.
A study on the number of flight feathers has concluded that ''Caudipteryx'' was secondarily flightless.
Classification
The consensus view, based on several cladistic
Cladistics ( ; from Ancient Greek 'branch') 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 ...
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 (geology), Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like s ...
, and the oviraptorosaurians are non-avian theropod dinosaurs. '' Incisivosaurus'' 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. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
,[Paul, G.S. (2002). ''Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds.'' Johns Hopkins University Press, Baltimore. ] Lü ''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 have concluded that ''Caudipteryx'' is not a theropod
Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
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, Dromaeosauridae, Troodontidae, ...
ns, is a flightless bird, and that birds evolved from non-dinosaurian archosaurs
Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
.
A weighted cladogram from 2014, using TNT, is shown below.
Relationship with birds
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 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 () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
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.
Paleobiology
Diet
''Caudipteryx'' is thought to have been an omnivore
An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
. 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 ...
s are preserved. As in some herbivorous dinosaurs, the avialan
Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative defi ...
'' 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 (birds and other dinosaurs, crocodiles, alligators, pterosaurs), earthworms, some gast ...
s would have been.
Paleoenvironment
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 preserve ...
s were recovered from the Yixian Formation
The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
in Liaoning
)
, image_skyline =
, image_alt =
, image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong
, image_map = Liaoning in China (+all claims hatched).svg
, ...
, China
China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. 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 southwest ...
. 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'' and ''Sinornithosaurus
''Sinornithosaurus'' (derived from a combination of Latin and Ancient Greek, Greek, meaning 'Chinese bird-lizard') is a genus of Feathered dinosaurs, feathered dromaeosaurid dinosaur from the early Cretaceous Period (geology), Period (late Barrem ...
''.
See also
* Dinosaur coloration
* Timeline of oviraptorosaur research
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
Oviraptorosauria
Dinosaur genera
Barremian dinosaurs
Yixian Formation
Fossil taxa described in 1998
Taxa named by Philip J. Currie
Taxa named by Mark Norell
Dinosaurs of China
Feathered dinosaurs