Pedopenna
Encyclopedia
Pedopenna is a genus
of small, feathered, maniraptora
n dinosaur
from the Daohugou Beds
in China
. It is possibly older than Archaeopteryx
, 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
(168 million years ago) and early Late Jurassic
Period (164-152 million years ago).
The name Pedopenna refers to the long pennaceous feathers on the metatarsus; daohugouensis refers to the locality of Daohugou, where the holotype
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 maniraptora
n dinosaurs that includes both deinonychosaurs
and avialans
(the lineage including modern birds), but some scientists have classified it as a true avialan more closely related to modern birds than to deinonychosaurs.
), 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, who interpreted the Daohugou
fossil beds where Pedopenna was found as mid to late Jurassic
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
.
s on the metatarsus
(foot). Some deinonychosaurs are also known to have these 'hind wings', but those of Pedopenna differ from those of animals like Microraptor
. Pedopenna hind wings were smaller and more rounded in shape. The longest feathers 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
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.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of small, feathered, maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...
n dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
from the Daohugou Beds
Daohugou Beds
The Daohugou Beds are a series of fossil-bearing rock deposits located in northeastern China, with the type locality around Daohugou village of Ningcheng County south of Chifeng, Inner Mongolia, but extending into southwest Liaoning. The rocks are grey, finely bedded, lacustrine, sandy mudstones...
in China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
. It is possibly older than Archaeopteryx
Archaeopteryx
Archaeopteryx , sometimes referred to by its German name Urvogel , is a genus of theropod dinosaur that is closely related to birds. The name derives from the Ancient Greek meaning "ancient", and , meaning "feather" or "wing"...
, though the age of the Daohugou Beds
Daohugou Beds
The Daohugou Beds are a series of fossil-bearing rock deposits located in northeastern China, with the type locality around Daohugou village of Ningcheng County south of Chifeng, Inner Mongolia, but extending into southwest Liaoning. The rocks are grey, finely bedded, lacustrine, sandy mudstones...
where it was found is debated. A majority of studies suggest that beds probably date from between the late Middle Jurassic
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from 176-161 million years ago. In European lithostratigraphy, rocks of this Middle Jurassic age are called the Dogger....
(168 million years ago) and early Late Jurassic
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.2 ± 4.0 to 145.5 ± 4.0 million years ago , which is preserved in Upper Jurassic strata. In European lithostratigraphy, the name "Malm" indicates rocks of Late Jurassic age...
Period (164-152 million years ago).
The name Pedopenna refers to the long pennaceous feathers on the metatarsus; daohugouensis refers to the locality of Daohugou, where the holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
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
Paraves
Paraves is a branch-based clade containing birds and other closely related dinosaurs. The paravians include the Avialae, including Archaeopteryx and modern birds, and the Deinonychosauria, which includes the dromaeosaurids and troodontids....
, the group of maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...
n dinosaurs that includes both deinonychosaurs
Deinonychosauria
The Deinonychosauria were a clade of coelurosaurian theropod dinosaurs from the Late Jurassic and Cretaceous periods. These omnivores and carnivores are known for their switchblade-like second toe claws and for displaying numerous bird-like characteristics. The clade has been divided into two...
and avialans
Avialae
Avialae is a clade of dinosaurs containing their only living representatives, birds , and the most immediate extinct relatives of birds.-Competing definitions:...
(the lineage including modern birds), but some scientists have classified it as a true avialan more closely related to modern birds than to deinonychosaurs.
Description
The feet of Pedopenna resembled those of the related troodontids and dromaeosaurids (which together form the group DeinonychosauriaDeinonychosauria
The Deinonychosauria were a clade of coelurosaurian theropod dinosaurs from the Late Jurassic and Cretaceous periods. These omnivores and carnivores are known for their switchblade-like second toe claws and for displaying numerous bird-like characteristics. The clade has been divided into two...
), 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, who interpreted the Daohugou
Daohugou Beds
The Daohugou Beds are a series of fossil-bearing rock deposits located in northeastern China, with the type locality around Daohugou village of Ningcheng County south of Chifeng, Inner Mongolia, but extending into southwest Liaoning. The rocks are grey, finely bedded, lacustrine, sandy mudstones...
fossil beds where Pedopenna was found as mid to late Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
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
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...
.
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 featherFeather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...
s on the metatarsus
Metatarsus
The metatarsus or metatarsal bones 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 medial side : the first, second, third, fourth, and fifth...
(foot). Some deinonychosaurs are also known to have these 'hind wings', but those of Pedopenna differ from those of animals like Microraptor
Microraptor
Microraptor 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 feathers 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
Covert (feather)
A covert feather on a bird is one of a set of feathers, called coverts, which as the name implies, cover other feathers. The coverts help to smooth airflow over the wings and tail.- Wing-coverts :...
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.