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''Pedopenna'' (meaning "foot feather") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of small, feathered, maniraptoran
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 ...
from the Daohugou Beds in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. It is possibly older than ''
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" ...
'', though the age of the Daohugou Beds where it was found is debated. A majority of studies suggest that beds probably date from between the late
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively ...
(168 million years ago) and early
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Period (164-152 million years ago). The name ''Pedopenna'' refers to the long
pennaceous The pennaceous feather is a type of feather present in most modern birds and in some other species of maniraptoriform dinosaurs. Description A pennaceous feather has a stalk or quill. Its basal part, called a ''calamus'', is embedded in the skin ...
feathers on the metatarsus; ''daohugouensis'' refers to the locality of Daohugou, where the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
was found. ''Pedopenna daohugouensis'' probably measured 1 meter (3 ft) or less in length, but since this species is only known from the hind legs, the actual length is difficult to estimate. ''Pedopenna'' was originally classified as a paravian, the group of maniraptoran dinosaurs that includes both deinonychosaurs and
avialans 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 ...
(the lineage including modern birds), but some scientists have classified it as a true avialan more closely related to modern birds than to deinonychosaurs. More recently, it has been recovered as a scansoriopterygid.Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. .


Description

The feet of ''Pedopenna'' resembled those of the related troodontids and
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 (which together form the group Deinonychosauria), though were overall more primitive. In particular, the second toe of ''Pedopenna'' was not as specialized as in deinonychosaurs. While ''Pedopenna'' did have an enlarged claw and slightly shortened second toe, it was not as highly developed as the strongly curved, sickle-like claws of its relatives. Xu Xing and
Zhang Fucheng Zhang may refer to: Chinese culture, etc. * Zhang (surname) (張/张), common Chinese surname ** Zhang (surname 章), a rarer Chinese surname * Zhang County (漳县), of Dingxi, Gansu * Zhang River (漳河), a river flowing mainly in Henan * ...
, who interpreted the
Daohugou The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerates ...
fossil beds where ''Pedopenna'' was found as mid to late
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
in age, used the presence of such a primitive member of the avian lineage, in combination with many primitive members of closely related lineages there, to support the idea that birds originated in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
.


Feathers

The bird-like characteristics of ''Pedopenna'' are further evidence of the dinosaur-bird evolutionary relationship. Apart from having a very bird-like skeletal structure in its legs, ''Pedopenna'' was remarkable due to the presence of long pennaceous feathers on the
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
(foot). Some deinonychosaurs are also known to have these 'hind wings', but those of ''Pedopenna'' differ from those of animals like ''
Microraptor ''Microraptor'' ( Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. ...
''. ''Pedopenna'' hind wings were smaller and more rounded in shape. The longest
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 premie ...
s were slightly shorter than the metatarsus, at about 55 mm (2 in) long. Additionally, the feathers of ''Pedopenna'' were symmetrical, unlike the asymmetrical feathers of some deinonychosaurs and birds. Since asymmetrical feathers are typical of animals adapted to flying, it is likely that ''Pedopenna'' represents an early stage in the development of these structures. While many of the feather impressions in the fossil are weak, it is clear that each possessed a rachis and barbs, and while the exact number of foot feathers is uncertain, they are more numerous than in the hind-wings of ''Microraptor''. ''Pedopenna'' also shows evidence of shorter feathers overlying the long foot feathers, evidence for the presence of
covert Secrecy is the practice of hiding information from certain individuals or groups who do not have the "need to know", perhaps while sharing it with other individuals. That which is kept hidden is known as the secret. Secrecy is often controvers ...
s as seen in modern birds. Since the feathers show fewer aerodynamic adaptations than the similar hind wings of ''Microraptor'', and appear to be less stiff, suggests that if they did have some kind of aerodynamic function, it was much weaker than in deinonychosaurs and birds. Xu and Zhang, in their 2005 description of ''Pedopenna'', suggested that the feathers could be ornamental, or even vestigial. It is possible that a hind wing was present in the ancestors of deinonychosaurs and birds, and later lost in the bird lineage, with ''Pedopenna'' representing an intermediate stage where the hind wings are being reduced from a functional gliding apparatus to a display or insulatory function. this theory may be confirmed by the recent discovery of ''
Serikornis ''Serikornis'' is a genus of small, feathered anchiornithid dinosaur from the Upper Jurassic Tiaojishan Formation of Liaoning, China. It is represented by the type species ''Serikornis sungei''. Discovery ''Serikornis'', first described in Aug ...
'', an anchiornithid with soft, symmetrical feathers and filaments on parts of the body that were similar to Pedopenna's.


References

{{Taxonbar, from=Q220431 Prehistoric paravians Jurassic dinosaurs of Asia Fossil taxa described in 2005