Maniraptoriformes

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Maniraptoriformes
Temporal range:
Middle JurassicPresent, 165–0  Ma [1]
O
S
D
C
P
T
J
K
Pg
N
Possible Early Jurassic record [2]
Struthiomimus ROM.jpg
Ornithomimid ( Struthiomimus altus ) fossil cast
House sparrow04.jpg
House sparrow (Passer domesticus)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Neocoelurosauria
Clade: Maniraptoriformes
Holtz, 1995
Subgroups
Synonyms
  • Bullatosauria Holtz, 1993
  • Avepectora Paul, 2002

Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings [3] that contains ornithomimosaurs and maniraptorans. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of Ornithomimus 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 belong to this clade. In particular, a fossil of the Alvarezsauridae Shuvuuia has a version of keratin consistent with that of avian feathers. [4]

Many nearly complete fossil maniraptoriforms are known from the late Jurassic. Archaeopteryx is known from Bavaria at 155–150 Ma. Ornitholestes , the troodontid Hesperornithoides , Coelurus fragilis and Tanycolagreus topwilsoni are all known from the Morrison Formation in Wyoming at about 150 Ma. The Daohugou biota, including Anchiornis and Epidexipteryx , is the earliest record of maniraptoriformes, dating to about 160 Ma. One possible maniraptoriform, Eshanosaurus , lived even earlier, during the Early Jurassic, though its identification is controversial. [5]

The wide range of fossils in the early Cretaceous 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 were thought to be maniraptoriforms, [6] but as of the first decade of the 21st century, this seems unlikely. [7] [8]

History of study

In 1994, a study by paleontologist Thomas Holtz found a close relationship between the Ornithomimosauria 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 birds 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 and all its descendants. [9] 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. [10] In 2010, Paul used Avepectora for a smaller clade, excluding ornithomimosaurs, compsognathids and alvarezsauroids. [11]

Classification

The relationships among coelurosaurs shown below were found in a phylogenetic analysis by Godefroit and colleagues in 2013. [12]

Maniraptoriformes

Ornithomimosauria Hypothetical Deinocheirus (flipped).jpg

Maniraptora

Alvarezsauria Patagonykuspuertai (flipped).jpg

Therizinosauria Therizinosaurus Restoration.png

Pennaraptora

Oviraptorosauria Gigantoraptor Restoration.png

Paraves

Scansoriopterygidae Ambopteryx restoration.png

unnamed

See also

Related Research Articles

<span class="mw-page-title-main">Theropoda</span> Clade of dinosaurs

Theropoda, whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally carnivorous, although a number of theropod groups evolved to become herbivores and omnivores. Theropods first appeared during the Carnian age of the late Triassic period 231.4 million years ago (Ma) and included the majority of large terrestrial carnivores from the Early Jurassic until at least the close of the Cretaceous, about 66 Ma. In the Jurassic, birds evolved from small specialized coelurosaurian theropods, and are today represented by about 10,500 living species.

<span class="mw-page-title-main">Troodontidae</span> Extinct family of bird-like dinosaurs

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 discoveries of complete and articulated specimens, have helped to increase understanding about this group. Anatomical studies, particularly studies of the most primitive troodontids, like Sinovenator, demonstrate striking anatomical similarities with Archaeopteryx and primitive dromaeosaurids, and demonstrate that they are relatives comprising a clade called Paraves.

<span class="mw-page-title-main">Coeluridae</span> Extinct family of dinosaurs

Coeluridae is a historically unnatural group of generally small, carnivorous dinosaurs from the late Jurassic Period. For many years, any small Jurassic or Cretaceous theropod that did not belong to one of the more specialized families recognized at the time was classified with the coelurids, creating a confusing array of 'coelurid' theropods that were not closely related. Although they have been traditionally included in this family, there is no evidence that any of these primitive coelurosaurs form a natural group with Coelurus, the namesake of Coeluridae, to the exclusion of other traditional coelurosaur groups.

<span class="mw-page-title-main">Alvarezsauridae</span> Family of extinct long-legged dinosaurs

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 specialized. They had tiny but stout forelimbs, with compact, bird-like hands. Their skeletons suggest that they had massive breast and arm muscles, possibly adapted for digging or tearing. They had long, tube-shaped snouts filled with tiny teeth. They have been interpreted as myrmecophagous, adapted to prey on colonial insects such as termites, with the short arms acting as effective digging instruments to break into nests.

<span class="mw-page-title-main">Maniraptora</span> Clade of dinosaurs

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, Dromaeosauridae, Troodontidae, Oviraptorosauria, and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period, and survive today as living birds.

<span class="mw-page-title-main">Coelurosauria</span> Clade of dinosaurs

Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.

<span class="mw-page-title-main">Tetanurae</span> Clade containing most theropod dinosaurs

Tetanurae is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans. Tetanurans are defined as all theropods more closely related to modern birds than to Ceratosaurus and contain the majority of predatory dinosaur diversity. Tetanurae likely diverged from its sister group, Ceratosauria, during the late Triassic. Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.

<span class="mw-page-title-main">Ornithomimosauria</span> Extinct clade of theropod dinosaurs

Ornithomimosauria 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, as well as Africa and possibly Australia. The group first appeared in the Early Cretaceous and persisted until the Late Cretaceous. Primitive members of the group include Nqwebasaurus, Pelecanimimus, Shenzhousaurus, Hexing and Deinocheirus, the arms of which reached 2.4 m (8 feet) in length. More advanced species, members of the family Ornithomimidae, include Gallimimus, Struthiomimus, and Ornithomimus. Some paleontologists, like Paul Sereno, consider the enigmatic alvarezsaurids to be close relatives of the ornithomimosaurs and place them together in the superfamily Ornithomimoidea.

<i>Struthiomimus</i> Extinct genus of reptile

Struthiomimus, meaning "ostrich-mimic", is a genus of ornithomimid dinosaurs from the late Cretaceous of North America. Ornithomimids were long-legged, bipedal, ostrich-like dinosaurs with toothless beaks. The type species, Struthiomimus altus, is one of the more common, smaller dinosaurs found in Dinosaur Provincial Park; their overall abundance—in addition to their toothless beak—suggests that these animals were mainly herbivorous or omnivorous, rather than purely carnivorous. Similar to the modern extant ostriches, emus, and rheas, ornithomimid dinosaurs likely lived as opportunistic omnivores, supplementing a largely plant-based diet with a variety of small mammals, reptiles, amphibians, insects, invertebrates, and anything else they could fit into their mouth, as they foraged.

<span class="mw-page-title-main">Therizinosauria</span> Extinct clade of dinosaurs

Therizinosaurs are an extinct group of large herbivorous theropod dinosaurs whose fossils have been found across the Middle Jurassic to Late Cretaceous deposits in Europe, Asia and North America. Various features of the forelimbs, skull and pelvis unite these finds as both theropods and maniraptorans, making them relatives of birds. The name of the representative genus, Therizinosaurus, is derived from the Greek θερίζω and σαῦρος. The older representative, Segnosaurus, is derived from the Latin sēgnis ('slow') and the Greek σαῦρος.

<i>Proceratosaurus</i> Extinct genus of dinosaurs

Proceratosaurus is a genus of carnivorous theropod dinosaur from the Middle Jurassic (Bathonian) of England. It contains a single species. P. bradleyi, known from a mostly complete skull and lower jaws. Proceratosaurus was a small dinosaur, estimated to measure around 3 m (9.8 ft) in length. Its name refers to how it was originally thought to be an ancestor of Ceratosaurus, due to the partially preserved portion of the crest of Proceratosaurus superficially resembling the small crest of Ceratosaurus. Now, however, it is considered a coelurosaur, specifically a member of the family Proceratosauridae, and amongst the earliest known members of the clade Tyrannosauroidea.

<span class="mw-page-title-main">Tyrannosauroidea</span> Extinct superfamily of dinosaurs

Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period. By the end of the Cretaceous Period, tyrannosauroids were the dominant large predators in the Northern Hemisphere, culminating in the gigantic Tyrannosaurus. Fossils of tyrannosauroids have been recovered on what are now the continents of North America, Europe and Asia, with fragmentary remains possibly attributable to tyrannosaurs also known from South America and Australia.

Thomas Richard Holtz Jr. 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, ecomorphology, and locomotion of terrestrial predators, especially on tyrannosaurids and other theropod dinosaurs. He wrote the book Dinosaurs and is the author or co-author of the chapters "Saurischia", "Basal Tetanurae", and "Tyrannosauroidea" in the second edition of The Dinosauria. He has also been consulted as a scientific advisor for the Walking with Dinosaurs BBC series as well as the Discovery special When Dinosaurs Roamed America, and has appeared in numerous documentaries focused on prehistoric life, such as Jurassic Fight Club on History and Monsters Resurrected, Dinosaur Revolution and Clash of the Dinosaurs on Discovery.

<span class="mw-page-title-main">Compsognathidae</span> Extinct family of dinosaurs

Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other dinosaurs such as Archaeopteryx inspired the idea for the connection between dinosaur reptiles and modern-day avian species. Compsognathid fossils preserve diverse integument — skin impressions are known from four genera commonly placed in the group, Compsognathus, Sinosauropteryx, Sinocalliopteryx, and Juravenator. While the latter three show evidence of a covering of some of the earliest primitive feathers over much of the body, Juravenator and Compsognathus also show evidence of scales on the tail or hind legs. "Ubirajara jubatus", informally described in 2020, had elaborate integumentary structures on its back and shoulders superficially similar to the display feathers of a standardwing bird-of-paradise, and unlike any other non-avian dinosaur currently described.

<span class="mw-page-title-main">Avialae</span> Clade including all birds and their ancestors

Avialae 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.

<span class="mw-page-title-main">Paraves</span> Clade of all dinosaurs that are more closely related to birds than to oviraptorosaurs

Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avialans, which include diverse extinct taxa as well as the over 10,000 species of living birds. Basal members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species. A number of differing scientific interpretations of the relationships between paravian taxa exist. New fossil discoveries and analyses make the classification of Paraves an active subject of research.

<i>Epidexipteryx</i> Extinct genus of dinosaurs

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 ornamental feathers in the fossil record.

<span class="mw-page-title-main">Averostra</span> Clade of dinosaurs

Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs, namely Ceratosauria and Tetanurae, and represent the only group of post-Early Jurassic theropods. Both survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians, megaraptorans an incertae sedis group within Tetanurae, and two groups of tetanurans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of Maniraptoriformes, Aves, survived the extinction event and persisted to the present day.

<span class="mw-page-title-main">Timeline of ornithomimosaur research</span>

This timeline of ornithomimosaur research is a chronological listing of events in the history of paleontology focused on the ornithomimosaurs, a group of bird-like theropods popularly known as the ostrich dinosaurs. Although fragmentary, probable, ornithomimosaur fossils had been described as far back as the 1860s, the first ornithomimosaur to be recognized as belonging to a new family distinct from other theropods was Ornithomimus velox, described by Othniel Charles Marsh in 1890. Thus the ornithomimid ornithomimosaurs were one of the first major Mesozoic theropod groups to be recognized in the fossil record. The description of a second ornithomimosaur genus did not happen until nearly 30 years later, when Henry Fairfield Osborn described Struthiomimus in 1917. Later in the 20th century, significant ornithomimosaur discoveries began occurring in Asia. The first was a bonebed of "Ornithomimus" asiaticus found at Iren Debasu. More Asian discoveries took place even later in the 20th century, including the disembodied arms of Deinocheirus mirificus and the new genus Gallimimus bullatus. The formal naming of the Ornithomimosauria itself was performed by Rinchen Barsbold in 1976.

<span class="mw-page-title-main">Anchiornithidae</span> Extinct family of dinosaurs

Anchiornithidae is a family of small paravian dinosaurs. Anchiornithids have been classified at varying positions in the paravian tree, with some scientists classifying them as a distinct family, a basal subfamily of Troodontidae, members of Archaeopterygidae, or an assemblage of dinosaurs that are an evolutionary grade within Avialae or Paraves.

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