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Maniraptoriformes is 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
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 with
pennaceous feather 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 ...
s and
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...
s that contains
ornithomimosaurs Ornithomimosauria ("bird-mimic lizards") are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia (now Asia, Europe and North ...
and
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. This group was named by
Thomas Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomorp ...
, who defined it as "the most recent common ancestor of ''
Ornithomimus ''Ornithomimus'' (; "bird mimic") is a genus of ornithomimid dinosaurs from the Late Cretaceous Period of what is now North America. ''Ornithomimus'' was a swift bipedal theropod which fossil evidence indicates was covered in feathers, equipped w ...
'' and
birds 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 ...
, and all descendants of that common ancestor."


Fossil evidence

Many fossils have been discovered in recent years, particularly in China. Many of the feathered dinosaurs belong to this clade. In particular, a fossil of the
Alvarezsauridae Alvarezsauridae is a family of small, long-legged dinosaurs. Although originally thought to represent the earliest known flightless birds, they are now thought to be an early diverging branch of maniraptoran theropods. Alvarezsaurids were highly ...
''
Shuvuuia ''Shuvuuia'' is a genus of bird-like theropod dinosaur from the late Cretaceous period of Mongolia. It is a member of the family Alvarezsauridae, small coelurosaurian dinosaurs which are characterized by short but powerful forelimbs specialized f ...
'' has a version of keratin consistent with that of avian feathers. Many nearly complete
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 ...
maniraptoriforms are known from the late Jurassic. ''
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" ...
'' is known from Bavaria at 155–150 Ma. ''
Ornitholestes ''Ornitholestes'' (meaning "bird robber") is a small theropod dinosaur of the late Jurassic (Brushy Basin Member of the Morrison Formation, middle Kimmeridgian age, about 154 million years agoTurner, C.E. and Peterson, F., (1999). "Biostratigraph ...
'', the troodontid ''
Hesperornithoides ''Hesperornithoides'' (meaning "western bird form"; nicknamed "Lori") is a genus of troodontid theropod dinosaur that lived in North America during the Late Jurassic period. Discovery Although several Troodontid teeth were found, with the trood ...
'', ''
Coelurus fragilis ''Coelurus'' ( ) is a genus of coelurosaurian dinosaur from the Late Jurassic period (mid-late Kimmeridgian faunal stage, 155–152 million years ago). The name means "hollow tail", referring to its hollow tail vertebrae (Greek ÎºÎ¿á¿–Î»Î¿Ï ...
'' and '' Tanycolagreus topwilsoni'' are all known from the
Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...
in Wyoming at about 150 Ma. The
Daohugou biota The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian- Oxfordian stages). It is known for its exceptionally preserved fossils, including those of ...
, including '' Anchiornis'' and '' Epidexipteryx'', is the earliest record of maniraptoriformes, dating to about 160 Ma. One possible maniraptoriform, ''
Eshanosaurus ''Eshanosaurus'' is a genus of a dinosaur from the early Jurassic Period. It is known only from a fossil partial lower jawbone, found in China. It may be a therizinosaurian, and if so the earliest known coelurosaur. Discovery and naming The typ ...
'', lived even earlier, during the Early Jurassic, though its identification is controversial. The wide range of fossils in 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 ...
and morphological evidence suggests that the main branches of maniraptoriform differentiation were separate before the end of the Jurassic. Until recently, the relatives of ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosa ...
'' were thought to be maniraptoriforms, but as of the first decade of the 21st century, this seems unlikely.


History of study

In 1994, a study by
paleontologist 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 ...
Thomas Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomorp ...
found a close relationship between the
Ornithomimosauria Ornithomimosauria ("bird-mimic lizards") are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period (geology), Period of Laurasia (now Asia ...
and Troodontidae, and named this group Bullatosauria. Holtz rejected this hypothesis in 1999, and most paleontologists now consider troodontids to be much more closely related to either
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 ...
s or Dromaeosauridae than they are to ornithomimosaurs, causing the Bullatosauria to be abandoned. The name referred to the inflated (bulbous) sphenoid both groups shared. Holtz defined the group as the clade containing the most recent common ancestor of '' Troodon'' and ''
Ornithomimus ''Ornithomimus'' (; "bird mimic") is a genus of ornithomimid dinosaurs from the Late Cretaceous Period of what is now North America. ''Ornithomimus'' was a swift bipedal theropod which fossil evidence indicates was covered in feathers, equipped w ...
'' and all its descendants. The concept is now considered redundant, and the clade Bullatosauria is now viewed as synonymous with Maniraptoriformes. In 2002, Gregory S. Paul named an apomorphy-based clade Avepectora, defined to include all theropods with a bird-like arrangement of the pectoral bones, where the angled shoulder girdle (coracoids) come in contact with the breastbone (sternum). According to Paul, ornithomimosaurs are the most basal members of this group.Paul, G.S. (2002). ''Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds''. Baltimore: Johns Hopkins University Press. . In 2010, Paul used Avepectora for a smaller clade, excluding ornithomimosaurs, compsognathids and alvarezsauroids.Paul, G. S. (2010). The Princeton field guide to dinosaurs Princeton University Press. ''Princeton, New Jersey''.


Classification

The relationships among coelurosaurs shown below were found in a phylogenetic analysis by Godefroit and colleagues in 2013.


See also

* Coelurosauria *
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 ...


References

{{Taxonbar, from=Q134143 Extant Late Jurassic first appearances Oxfordian first appearances