Ornithomimosauria

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Ornithomimosaurs
Temporal range: Cretaceous, 140–66  Ma [1]
Ornithomimosauria Diversity.jpg
Collection of seven ornithomimosaurs, clockwise from top left: Gallimimus , Anserimimus , Ornithomimus , Deinocheirus , Harpymimus , Struthiomimus and "Gallimimus mongoliensis"
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Maniraptoriformes
Clade: Ornithomimosauria
Barsbold, 1976
Subgroups [2]
Synonyms
  • Deinocheirosauria Barsbold, 1976
  • Arctometatarsalia Holtz, 1994

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 America), as well as Africa and possibly Australia. [8] 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 (see classification below).

Contents

Description

The skulls of ornithomimosaurs were small, with large eyes, above relatively long and slender necks. The most basal members of the taxon (such as Pelecanimimus and Harpymimus ) had a jaw with small teeth, while the later and more derived species had a toothless beak. [9] The fore limbs ("arms") were long and slender and bore powerful claws. The hind limbs were long and powerful, with a long foot and short, strong toes terminating in hooflike claws. Ornithomimosaurs were probably among the fastest of all dinosaurs. Like other coelurosaurs, the ornithomimosaurian hide was feathered rather than scaly.

Feathers

Unambiguous evidence of feathers is known from Ornithomimus edmontonicus , of which there are multiple specimens preserving feather traces. [10] Deinocheirus and Pelecanimimus have been speculated to be feathered as well, the former due to the presence of a pygostyle, [11] and the later due to possible impressions (otherwise taken to be collagen fibers). There is a debate on whether ornithomimids possessed the pennaceous feathers seen in Pennaraptora. [12] Otherwise, a very ostrich-like plumage and feather range is known in one specimen of Ornithomimus. [13]

Classification

Named by O.C. Marsh in 1890, the family Ornithomimidae was originally classified as a group of "megalosaurs" (a "wastebasket taxon" containing any medium to large sized theropod dinosaurs), but as more theropod diversity was uncovered, their true relationships to other theropods started to resolve, and they were moved to the Coelurosauria. Recognizing the distinctiveness of ornithomimids compared to other dinosaurs, Rinchen Barsbold placed ornithomimids within their own infraorder, Ornithomimosauria, in 1976. The contents of Ornithomimidae and Ornithomimosauria varied from author to author as cladistic definitions began to appear for the groups in the 1990s.

In the early 1990s, prominent paleontologists such as Thomas R. Holtz Jr. proposed a close relationship between theropods with an arctometatarsalian foot; that is, bipedal dinosaurs in which the upper foot bones were 'pinched' together, an adaptation for running. Holtz (1994) defined the clade Arctometatarsalia as "the first theropod to develop the arctometatarsalian pes and all of its descendants." This group included the Troodontidae, Tyrannosauroidea, and Ornithomimosauria. Holtz (1996, 2000) later refined this definition to the branch-based " Ornithomimus and all theropods sharing a more recent common ancestor with Ornithomimus than with birds." Subsequently, the idea that all arctometatarsalian dinosaurs formed a natural group was abandoned by most paleontologists, including Holtz, as studies began to demonstrate that tyrannosaurids and troodontids were more closely related to other groups of coelurosaurs than they were to ornithomimosaurs. Since the strict definition of Arctometatarsalia was based on Ornithomimus, it became redundant with the name Ornithomimosauria under broad definitions of that clade, and the name Arctometatarsalia was mostly abandoned.

The paleontologist Paul Sereno, in 2005, proposed the clade "Ornithomimiformes", defining them as all species closer to Ornithomimus edmontonicus than to Passer domesticus . Because he had redefined Ornithomimosauria in a much narrower sense, a new term was made necessary within his preferred terminology to denote the clade containing the sistergroups Ornithomimosauria and Alvarezsauridae — previously the latter had been contained within the former. However, this concept only appeared on Sereno's Web site and has not yet been officially published as a valid name. [14] "Ornithomimiformes" was identical in content to Holtz's Arctometatarsalia, as it has a very similar definition. While "Ornithomimiformes" is the newer group, Sereno rejected the idea that Arctometatarsalia should take precedence, because the meaning of the former name has been changed very radically by Holtz. [14]

Phylogeny

Restoration of Beishanlong grandis Beishanlong grandis.jpg
Restoration of Beishanlong grandis

Ornithomimosauria has variously been used for the branch-based group of all dinosaurs closer to Ornithomimus than to birds, and in more restrictive senses. The more exclusive sense began to grow in popularity when the possibility arose that alvarezsaurids might fall under Ornithomimosauria if an inclusive definition were adopted. Another clade, Ornithomimiformes, was defined by Sereno (2005) as ( Ornithomimus velox > Passer domesticus ) and replaces the more inclusive use of Ornithomimosauria when alvarezsaurids or some other group are found to be closer relatives of ornithomimosaurs than maniraptorans, with Ornithomimosauria redefined to include dinosaurs closer to Ornithomimus than to alvarezsaurids. Gregory S. Paul has proposed that Ornithomimosauria might be a group of primitive, flightless birds, more advanced than Deinonychosauria and Oviraptorosauria. [15]

The cladogram below follows an analysis by Yuong-Nam Lee, Rinchen Barsbold, Philip J. Currie, Yoshitsugu Kobayashi, Hang-Jae Lee, Pascal Godefroit, François Escuillié & Tsogtbaatar Chinzorig. The analysis was published in 2014, and includes many ornithomimosaurian taxa. [2]

Coelurosauria

Zuolong Zuolong salleei.jpg

Tanycolagreus Tanycolagreus reconstruction.png

Tyrannoraptora

Proceratosaurus

Tyrannosauroidea Daspletosaurus torosus steveoc flipped.jpg

Ornitholestes Ornitholestes reconstruction.png

Compsognathidae Sinosauropteryx color.jpg

Maniraptoriformes
Ornithomimosauria

Nqwebasaurus Nqwebasaurus.jpg

Pelecanimimus Pelecanimimus restoration.jpeg

Shenzhousaurus Shenzhousaurus.jpg

Harpymimus Harpymimus steveoc (flipped).jpg

Deinocheiridae

Beishanlong

Garudimimus Garudimimus Restoration (flipped).png

Deinocheirus Hypothetical Deinocheirus (flipped).jpg

Ornithomimidae

Anserimimus

Gallimimus Gallimimus Steveoc86 (flipped).jpg

Ornithomimus -Ornithomimus- sp. by Tom Parker (flipped).png

Struthiomimus Struthiomimus BW.jpg

Maniraptora

Alvarezsauroidea Shuvuuia.jpg

Therizinosauroidea Nanshiungosaurus Restoration.png

Oviraptorosauria Ajancingenia reconstruction (white background).png

Paraves Confuciusornis plumage pattern.jpg

The cladogram below follows the phylogenetic analysis by Scott Hartman and colleagues in 2019, which has included a vast majority of species and uncertain specimens, resulting in a novel phylogenetic arrangement. [16]

Ornithomimosauria
Deinocheiridae

Hexing

Deinocheirus Hypothetical Deinocheirus.jpg

Shenzhousaurus Shenzhousaurus.jpg

Ornithomimidae

Aepyornithomimus

Ornithomimus "Ornithomimus" sp. by Tom Parker.png

Sinornithomimus Sinornithomimus.jpg

Garudimimidae

Arkansaurus

Archaeornithomimus Archaeornithomimus.png

"Grusimimus" GIN 960910KD

Beishanlong

Garudimimus Garudimimus Restoration.png

Nedcolbertia Nedclobertia.jpg

Angeac taxon

Harpymimus Harpymimus steveoc.jpg

"Saltillomimus" SEPCP 16/237 Saltillomimus rapidus.jpg

Tototlmimus

"Gallimimus" mongoliensis

MNAP 11762A

"Ornithomimus sedens" BHI 1266

Rativates Rativates.png

Struthiomimus Struthiomimus BW.jpg

Gallimimus bullatus Gallimimus Steveoc86.jpg

Timimus

Anserimimus

Qiupalong Qiupalong Restoration.png

Dromiceiomimus

Below is a cladogram by Serrano-Brañas et al., 2020, showing an analysis more in line with previous assumptions about ornithomimosaur classification. [17]

Ornithomimosauria

Nqwebasaurus Nqwebasaurus.jpg

Beishanlong

Pelecanimimus Pelecanimimus restoration.jpeg

Shenzhousaurus Shenzhousaurus.jpg

Deinocheiridae

Harpymimus Harpymimus steveoc.jpg

Deinocheirus Hypothetical Deinocheirus.jpg

Garudimimus Garudimimus Restoration.png

Paraxenisaurus

Ornithomimidae

Archaeornithomimus Archaeornithomimus.png

Bissekty Taxon

Sinornithomimus Sinornithomimus.jpg

Gallimimus Gallimimus Steveoc86.jpg

Struthiomimus Struthiomimus BW.jpg

Qiupalong Qiupalong Restoration.png

Anserimimus

Ornithomimus "Ornithomimus" sp. by Tom Parker.png

Palaeobiology

Ornithomimosaurs probably acquired most of their calories from plants. Many ornithomimosaurs, including primitive species, have been found with numerous gastroliths in their stomachs, characteristic of herbivores. Henry Fairfield Osborn suggested that the long, sloth-like "arms" of ornithomimosaurs may have been used to pull down branches on which to feed, an idea supported by further study of their strange, hook-like hands. [18] The sheer abundance of ornithomimids — they are the most common small dinosaurs in North America — is consistent with the idea that they were plant eaters, as herbivores usually outnumber carnivores in an ecosystem. However, they may have been omnivores that ate both plants and small animal prey.

Comparisons between the scleral rings of two ornithomimosaur genera ( Garudimimus and Ornithomimus ) and modern birds and reptiles indicate that they may have been cathemeral, active throughout the day at short intervals. [19]

Social behavior

Block containing eight specimens of Sinornithomimus Sinornithomimus dongi.jpg
Block containing eight specimens of Sinornithomimus

Ornithomimosaurs are fairly well known for their gregarious life-styles. Some of the first findings of ornithomimosaur bonebeds were reported from the Iren Dabasu Formation in 1993 by Charles W. Gilmore. The bonebed consisted of numerous individuals of Archaeornithomimus ranging from young to adult remains. [20] Multiple specimens of Sinornithomimus were collected from a single monospecific bonebed with a considerable density of juvenile individuals—out of 14, 11 were juveniles—, suggesting a gregarious behavior for an increased protection from predators. The notable abundance of juveniles indicates a high mortality in them or that a large mass-mortality event of an entire group occurred, with more susceptibility in juveniles. Additionally, the increase in the tibia-femur ratio through the ontogeny of Sinornithomimus may indicate higher cursorial capacities in adults than in juveniles. [21] Moreover, and also contrary to the Sinornithomimus bonebed, a high concentration of ornithomimosaur specimens from the Bayshi Tsav locality was collected in a single multitaxic bonebed that is composed of at least five individuals at different ontogenetic stages. It is unlikely that the individuals of this bonebed represent a strategical social behaviour of a single species given the identification of at least two different taxa. Under this consideration, it is possible that a small pack of more than 10 individuals of different ornithomimosaurian herds was travelling together in optimal areas to find food resources, nesting sites or something else. [22] [23]

See also

Related Research Articles

<i>Gallimimus</i> Ornithomimid dinosaur genus from the Late Cretaceous Period

Gallimimus is a genus of theropod dinosaur that lived in what is now Mongolia during the Late Cretaceous period, about seventy million years ago (mya). Several fossils in various stages of growth were discovered by Polish-Mongolian expeditions in the Gobi Desert of Mongolia during the 1960s; a large skeleton discovered in this region was made the holotype specimen of the new genus and species Gallimimus bullatus in 1972. The generic name means "chicken mimic", referring to the similarities between its neck vertebrae and those of the Galliformes. The specific name is derived from bulla, a golden capsule worn by Roman youth, in reference to a bulbous structure at the base of the skull of Gallimimus. At the time it was named, the fossils of Gallimimus represented the most complete and best preserved ornithomimid material yet discovered, and the genus remains one of the best known members of the group.

<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.

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

Anserimimus is a genus of ornithomimid theropod dinosaur, from the Late Cretaceous Period of what is now Mongolia. It was a lanky, fast-running animal, possibly an omnivore. From what fossils are known, it probably closely resembled other ornithomimids, except for its more powerful forelimbs.

<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, if at all. 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.

<i>Ornithomimus</i> Ornithomimid dinosaur genus from the Late Cretaceous Period

Ornithomimus is a genus of ornithomimid theropod dinosaurs from the Campanian and Maastrichtian ages of Late Cretaceous Western North America. Ornithomimus was a swift, bipedal dinosaur which fossil evidence indicates was covered in feathers and equipped with a small toothless beak that may indicate an omnivorous diet. It is usually classified into two species: the type species, Ornithomimus velox, and a referred species, Ornithomimus edmontonicus. O. velox was named in 1890 by Othniel Charles Marsh on the basis of a foot and partial hand from the Denver Formation of Colorado. Another seventeen species have been named since then, though almost all of them have been subsequently assigned to new genera or shown to be not directly related to Ornithomimus velox. The best material of species still considered part of the genus has been found in Alberta, representing the species O. edmontonicus, known from several skeletons from the Horseshoe Canyon Formation. Additional species and specimens from other formations are sometimes classified as Ornithomimus, such as Ornithomimus samueli from the earlier Dinosaur Park Formation.

<i>Dromiceiomimus</i> Extinct genus of reptiles

Dromiceiomimus is a genus of ornithomimid theropod from the Late Cretaceous of Alberta, Canada. The type species, D. brevitertius, is considered a synonym of Ornithomimus edmontonicus by some authors, while others consider it a distinct and valid taxon. It was a small ornithomimid that weighed about 135 kilograms (298 lb).

<i>Deinocheirus</i> Genus of theropod dinosaurs

Deinocheirus is a genus of large ornithomimosaur that lived during the Late Cretaceous around 70 million years ago. In 1965, a pair of large arms, shoulder girdles, and a few other bones of a new dinosaur were first discovered in the Nemegt Formation of Mongolia. In 1970, this specimen became the holotype of the only species within the genus, Deinocheirus mirificus; the genus name is Greek for "horrible hand". No further remains were discovered for almost fifty years, and its nature remained a mystery. Two more complete specimens were described in 2014, which shed light on many aspects of the animal. Parts of these new specimens had been looted from Mongolia some years before, but were repatriated in 2014.

<i>Avimimus</i> Extinct genus of reptiles

Avimimus, meaning "bird mimic", is a genus of oviraptorosaurian theropod dinosaur, named for its bird-like characteristics, that lived in the late Cretaceous in what is now Mongolia, around 85 to 70 million years ago.

<i>Garudimimus</i> Ornithomimosaur genus from the Late Cretaceous

Garudimimus is a genus of ornithomimosaur that lived in Asia during the Late Cretaceous. The genus is known from a single specimen found in 1981 by a Soviet-Mongolian paleontological expedition in the Bayan Shireh Formation and formally described in the same year by Rinchen Barsbold; the only species is Garudimimus brevipes. Several interpretations about the anatomical traits of Garudimimus were made in posterior examinations of the specimen, but most of them were criticized during its comprehensive redescription in 2005. Extensive undescribed ornithomimosaur remains at the type locality of Garudimimus may represent additional specimens of the genus.

<span class="mw-page-title-main">Philip J. Currie</span> Canadian paleontologist and curator

Philip John Currie is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the 1980s, he became the director of the Canada-China Dinosaur Project, the first cooperative palaeontological partnering between China and the West since the Central Asiatic Expeditions in the 1920s, and helped describe some of the first feathered dinosaurs. He is one of the primary editors of the influential Encyclopedia of Dinosaurs, and his areas of expertise include theropods, the origin of birds, and dinosaurian migration patterns and herding behavior. He was one of the models for palaeontologist Alan Grant in the film Jurassic Park.

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

Archaeornithomimus is a genus of ornithomimosaurian theropod dinosaur that lived in Asia during the Late Cretaceous period, around 96 million years ago in the Iren Dabasu Formation.

<span class="mw-page-title-main">Ornithomimidae</span> Group of theropod dinosaurs

Ornithomimidae is a family of theropod dinosaurs which bore a superficial resemblance to modern ostriches. Ornithomimids were fast, omnivorous or herbivorous dinosaurs known mainly from the Late Cretaceous Period of Laurasia, though they have also been reported from the Lower Cretaceous Wonthaggi Formation of Australia.

<i>Nqwebasaurus</i> Extinct genus of dinosaur

Nqwebasaurus is a basal coelurosaur and is the basal-most member of the coelurosaurian clade Ornithomimosauria from the Early Cretaceous of South Africa. The name Nqwebasaurus is derived from the Xhosa word Nqweba which is the local name for the Kirkwood district, and thwazi is ancient Xhosa for "fast runner". Currently it is the oldest coelurosaur in Africa and shows that basal coelurosaurian dinosaurs inhabited Gondwana 50 million years earlier than previously thought. The type specimen of Nqwebasaurus was discovered by William J. de Klerk who is affiliated with the Albany Museum in Grahamstown. It is the only fossil of its species found to date and was found in the Kirkwood Formation of the Uitenhage Group. Nqwebasaurus has the unofficial nickname "Kirky", due to being found in the Kirkwood.

<span class="mw-page-title-main">Nemegt Formation</span> Geological formation in Mongolia

The Nemegt Formation is a geological formation in the Gobi Desert of Mongolia, dating to the Late Cretaceous. The formation consists of river channel sediments and contains fossils of fish, turtles, crocodilians, and a diverse fauna of dinosaurs, including birds.

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

Deinocheiridae is a family of ornithomimosaurian dinosaurs, living in Asia and the Americas from the Albian until the Maastrichtian. The family was originally named by Halszka Osmólska and Roniewicz in 1970, including only the type genus Deinocheirus. In a 2014 study by Yuong-Nam Lee and colleagues and published in the journal Nature, it was found that Deinocheiridae was a valid family. Lee et al. found that based on a new phylogenetic analysis including the recently discovered complete skeletons of Deinocheirus, the type genus, as well as Garudimimus and Beishanlong, could be placed as a successive group, with Beishanlong as the most primitive and Deinocheirus as most derived. The family Garudimimidae, named in 1981 by Rinchen Barsbold, is now a junior synonym of Deinocheiridae as the latter family includes the type genus of the former. The group existed from 115 to 69 million years ago, with Beishanlong living from 115 to 100 mya, Garudimimus living from 98 to 83 mya, and Deinocheirus living from 71 to 69 mya. Other genera included are Paraxenisaurus, and possibly Harpymimus and Hexing.

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

Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings 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."

<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.

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

Aepyornithomimus is a genus of ornithomimid theropod dinosaur from the Late Cretaceous Djadokhta Formation in Mongolia. It lived in the Campanian, around 75 million years ago, when the area is thought to have been a desert. The type and only species is A. tugrikinensis.

<i>Oksoko avarsan</i> Extinct species of dinosaur

Oksoko is a genus of oviraptorid dinosaur from the Late Cretaceous of Asia, that lived in what is now the Nemegt Formation in Mongolia. It includes the type species Oksoko avarsan.

References

  1. 1 2 Holtz, Thomas R. Jr. (2012) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
  2. 1 2 Lee, Y.-N.; Barsbold, R.; Currie, P.J.; Kobayashi, Y.; Lee, H.-J.; Godefroit, P.; Escuillié, F.; Chinzorig, T. (2014). "Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus". Nature. 515 (7526): 1–4. Bibcode:2014Natur.515..257L. doi:10.1038/nature13874. PMID   25337880. S2CID   2986017.
  3. Brownstein CD. (2016) Redescription of Arundel formation Ornithomimosaur material and a reinterpretation of Nedcolbertia justinhofmanni as an "Ostrich Dinosaur": Biogeographic implications. PeerJ Preprints 4:e2308v1 https://doi.org/10.7287/peerj.preprints.2308v1
  4. Choiniere, J. N.; Forster, C. A.; De Klerk, W. J. (2012). "New information on Nqwebasaurus thwazi, a coelurosaurian theropod from the Early Cretaceous (Hauteriverian?) Kirkwood Formation in South Africa". Journal of African Earth Sciences . 71–72: 1–17. Bibcode:2012JAfES..71....1C. doi:10.1016/j.jafrearsci.2012.05.005.
  5. Cuesta E, Vidal D, Ortega F, Shibata M, Sanz JL (2021). "Pelecanimimus (Theropoda: Ornithomimosauria) postcranial anatomy and the evolution of the specialized manus in Ornithomimosaurs and sternum in maniraptoriforms". Zoological Journal of the Linnean Society. 194 (2): 553–591. doi:10.1093/zoolinnean/zlab013.
  6. Sereno, P. (2017). "Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa". Ameghiniana. 54 (5): 576–616. doi:10.5710/AMGH.23.10.2017.3155. S2CID   134718338.
  7. Jin Liyong, Chen Jun & Pascal Godefroit (2012). "A New Basal Ornithomimosaur (Dinosauria: Theropoda) from the Early Cretaceous Yixian Formation, Northeast China". In Godefroit, P. (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. pp. 467–487. Bibcode:2012bdec.book.....G.
  8. Choiniere, Jonah N.; Forster, Catherine A.; De Klerk, William J. (2012). "New information on Nqwebasaurus thwazi, a coelurosaurian theropod from the Early Cretaceous Kirkwood Formation in South Africa". Journal of African Earth Sciences. 71–72: 1–17. Bibcode:2012JAfES..71....1C. doi:10.1016/j.jafrearsci.2012.05.005.
  9. Last of the Dinosaurs: The Cretaceous Period
  10. van der Reest, A.J.; Wolfe, A.P.; Currie, P.J. (2016). "A densely feathered ornithomimid (Dinosauria: Theropoda) from the Upper Cretaceous Dinosaur Park Formation, Alberta, Canada". Cretaceous Research. 58: 108–117. Bibcode:2016CrRes..58..108V. doi:10.1016/j.cretres.2015.10.004.
  11. Lee, Yuong-Nam; Barsbold, Rinchen; Currie, Philip J.; Kobayashi, Yoshitsugu; Lee, Hang-Jae; Godefroit, Pascal; Escuillié, François; Chinzorig, Tsogtbaatar (2014). "Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus". Nature. 515 (7526): 257–260. Bibcode:2014Natur.515..257L. doi:10.1038/nature13874. PMID   25337880. S2CID   2986017.
  12. Foth, Christian; Tischlinger, Helmut; Rauhut, Oliver W. M. (2014). "New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers". Nature. 511 (7507): 79–82. Bibcode:2014Natur.511...79F. doi:10.1038/nature13467. PMID   24990749. S2CID   4464659.
  13. Van Der Reest, Aaron J.; Wolfe, Alexander P.; Currie, Philip J. (2016). "A densely feathered ornithomimid (Dinosauria: Theropoda) from the Upper Cretaceous Dinosaur Park Formation, Alberta, Canada". Cretaceous Research. 58: 108–117. Bibcode:2016CrRes..58..108V. doi:10.1016/j.cretres.2015.10.004.
  14. 1 2 Sereno, P. C. (2005). Stem Archosauria—TaxonSearch Archived 2009-01-15 at the Wayback Machine [version 1.0, 2005 November 7]
  15. Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
  16. Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi: 10.7717/peerj.7247 . PMC   6626525 . PMID   31333906.
  17. Claudia Inés Serrano-Brañas; Belinda Espinosa-Chávez; S. Augusta Maccracken; Cirene Gutiérrez-Blando; Claudio de León-Dávila; José Flores Ventura (2020). "Paraxenisaurus normalensis, a large deinocheirid ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous), Coahuila, Mexico". Journal of South American Earth Sciences. 101: Article 102610. Bibcode:2020JSAES.10102610S. doi:10.1016/j.jsames.2020.102610. S2CID   218968100.
  18. Nicholls and Russell (1985).
  19. Schmitz and Motani (2011)
  20. Gilmore, C. W. (1933). "On the dinosaurian fauna of the Iren Dabasu Formation". Bulletin of the American Museum of Natural History. 67 (2): 23−78. hdl: 2246/355 .
  21. Kobayashi, Y.; Lü, J.-C. (2003). "A new ornithomimid dinosaur with gregarious habits from the Late Cretaceous of China" (PDF). Acta Palaeontologica Polonica. 48 (2): 235−259.
  22. Chinzorig, T.; Kobayashi, Y.; Saneyoshi, M.; Tsogtbaatar, K.; Batamkhatan, Z.; Ryuji, T. (2017). "Multitaxic bonebed of two new ornithomimids (Theropoda, Ornithomimosauria) from the Upper Cretaceous Bayanshiree Formnation of southeastern Gobi desert, Mongolia". Journal of Vertebrate Paleontology. Program and Abstracts: 97.
  23. Tsogtbaatar, K. (2019). Evolution, diversity, and disparity of ornithomimosaurs (Dinosauria: Theropoda) from the Upper Cretaceous of Mongolia (PDF) (PhD thesis). Hokkaido University. hdl:2115/74432.

Further reading