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 s ...

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 exp ...

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, Ovirapt ...

ns. This group was named by Thomas Holtz, 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 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 ...

, 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 s ...

belong to this clade. In particular, a fossil of the Alvarezsauridae '' Shuvuuia'' 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). "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 troo ...

'', '' Coelurus fragilis'' and '' Tanycolagreus topwilsoni'' are all known from the

Morrison Formation The Morrison Formation is a distinctive sequence of 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, sandstone, siltstone, ...

in Wyoming at about 150 Ma. The Daohugou biota, 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 "n ...

'' 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 (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma ...

, 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 foss ...

Thomas Holtz 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 disco ...

, 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 least ...

'' and ''

'' 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.

References

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

Fossil evidence

History of study

Classification

See also

References