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Maniraptoriformes is a clade 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 e ...
s that contains ornithomimosaurs 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, equippe ...
'' and birds, 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 A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been su ...
belong to this clade. In particular, a fossil of the Alvarezsauridae ''
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'' 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). "Biostratigrap ...
'', 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'' and '' Tanycolagreus topwilsoni'' are all known from the Morrison Formation 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 ''Anchiornis'' is a genus of small, four-winged paravian dinosaurs, with only one known species, the type species ''Anchiornis huxleyi'', named for its similarity to modern birds. The Latin name ''Anchiornis'' derives from a Greek word meaning " ...
'' and ''
Epidexipteryx ''Epidexipteryx'' is a genus of small paravian dinosaurs, known from one fossil specimen in the collection of the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. ''Epidexipteryx'' represents the earliest known example of o ...
'', 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 t ...
'', lived even earlier, during the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
, 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 ...
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 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. ''Tyrannosaurus'' live ...
'' 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 of Laurasia (now Asia, Europe and Nor ...
and
Troodontidae Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, 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 lightweig ...
s or
Dromaeosauridae 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 ...
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 ''Troodon'' ( ; ''Troödon'' in older sources) is a wastebasket taxon and a dubious genus of relatively small, bird-like dinosaurs known definitively from the Campanian age of the Late Cretaceous period (about 77  mya). It includes at leas ...
'' 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, equippe ...
'' 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 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 ...
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 Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, t ...
*
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