Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings[3] that contains ornithomimosaurs and maniraptors. 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."

Temporal range:
Late JurassicPresent, 167–0 Ma
Possible Early Jurassic record
Struthiomimus ROM
Ornithomimid (Struthiomimus altus) fossil cast
House sparrow04
House sparrow (Passer domesticus)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Clade: Maniraptoromorpha
Clade: Maniraptoriformes
Holtz, 1995
  • Bullatosauria Holtz, 1993
  • Ornithomimoidea Sereno, 1999
  • Avepectora Paul, 2002

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 this now 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]


The relationships among coelurosaurs shown below were found in a phylogenetic analysis by Godefroit and colleagues in 2013.[11]


Ornithomimosauria Hypothetical Deinocheirus (flipped)


Alvarezsauria Patagonykuspuertai (flipped)


Therizinosauria F. utahensis reconstruction (flipped)


Oviraptorosauria Gigantoraptor BW white background


Scansoriopterygidae Scansor chick.png


Eosinopteryx Eosinopteryx


DromaeosauridaeDeinonychus ewilloughby (flipped)


Troodontidae Zanabazar (flipped)

Avialae Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg

See also


  1. ^ Zichuan Qin; James Clark; Jonah Choiniere; Xing Xu (2019). A new alvarezsaurian theropod from the Upper Jurassic Shishugou Formation of western China. Scientific Reports. 9: Article number 11727. doi:10.1038/s41598-019-48148-7.
  2. ^ Azuma, Y., Xu X., Shibata, M., Kawabe, S., Miyata, K., and Imai, T. 2016. "A bizarre theropod from the Early Cretaceous of Japan highlighting mosaic evolution among coelurosaurians." Scientific Reports, 6(20478). doi:10.1038/srep20478
  3. ^ Zelenitsky, D. K.; Therrien, F.; Erickson, G. M.; Debuhr, C. L.; Kobayashi, Y.; Eberth, D. A.; Hadfield, F. (2012). "Feathered Non-Avian Dinosaurs from North America Provide Insight into Wing Origins". Science. 338 (6106): 510–514. Bibcode:2012Sci...338..510Z. doi:10.1126/science.1225376. PMID 23112330.
  4. ^ Schweitzer, M.H.; Watt, J.A.; Avci, R.; Knapp, L.; Chiappe, L.; Norell, M.; Marshall, M. (1999). "Beta-keratin specific immunological reactivity in feather-like structures of the Cretaceous Alvarezsaurid, Shuvuuia deserti". Journal of Experimental Zoology. 285 (2): 146–157. doi:10.1002/(SICI)1097-010X(19990815)285:2<146::AID-JEZ7>3.0.CO;2-A. PMID 10440726.
  5. ^ Barrett, P.M. (2009). "The affinities of the enigmatic dinosaur Eshanosaurus deguchiianus from the Early Jurassic of Yunnan Province, People's Republic of China". Palaeontology. 52 (4): 681–688. doi:10.1111/j.1475-4983.2009.00887.x.
  6. ^ Benton, Michael J. (2004) Vertebrate Palaentology. 3rd ed. Wiley. ISBN 978-0-632-05637-8
  7. ^ Weishampel, David B., Dodson, Peter, and Osmólska, Halszka (2004) The Dinosauria. University of California Press. ISBN 0-520-25408-2
  8. ^ Senter, Phil (2007). "A new look at the phylogeny of coelurosauria (dinosauria: theropoda)". Journal of Systematic Palaeontology. 5 (4): 429. doi:10.1017/S1477201907002143.
  9. ^ Holtz, T.R. Jr (1994). "The phylogenetic position of the Tyrannosauridae. Implications for theropod systematics". Journal of Paleontology. 68: 1100–1117.
  10. ^ Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press. ISBN 0801867630.
  11. ^ Godefroit, Pascal; Cau, Andrea; Hu, Dong-Yu; Escuillié, François; Wu, Wenhao; Dyke, Gareth (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. 498 (7454): 359–362. Bibcode:2013Natur.498..359G. doi:10.1038/nature12168. PMID 23719374.

Australovenator (meaning "southern hunter") is a genus of megaraptorid theropod dinosaur from Cenomanian (Late Cretaceous)-age Winton Formation (dated to 95 million years ago) of Australia. It is known from partial cranial and postcranial remains which were described in 2009 by Scott Hocknull and colleagues, although additional descriptions and analyses continue to be published. It is the most complete predatory dinosaur discovered in Australia.


Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans 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.


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.


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, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaur group alive today.Most feathered dinosaurs discovered so far have been coelurosaurs. Philip J. Currie considers it likely and probable that all coelurosaurs were feathered. In the past, Coelurosauria was used to refer to all small theropods, but this classification has since been abolished.


Deinocheiridae is a family of ornithomimosaurian dinosaurs, living in Asia 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.

Dinosaur classification

Dinosaur classification began in 1842 when Sir Richard Owen placed Iguanodon, Megalosaurus, and Hylaeosaurus in "a distinct tribe or suborder of Saurian Reptiles, for which I would propose the name of Dinosauria." In 1887 and 1888 Harry Seeley divided dinosaurs into the two orders Saurischia and Ornithischia, based on their hip structure. These divisions have proved remarkably enduring, even through several seismic changes in the taxonomy of dinosaurs.

The largest change was prompted by entomologist Willi Hennig's work in the 1950s, which evolved into modern cladistics. For specimens known only from fossils, the rigorous analysis of characters to determine evolutionary relationships between different groups of animals (clades) proved incredibly useful. When computer-based analysis using cladistics came into its own in the 1990s, paleontologists became among the first zoologists to almost wholeheartedly adopt the system. Progressive scrutiny and work upon dinosaurian interrelationships, with the aid of new discoveries that have shed light on previously uncertain relationships between taxa, have begun to yield a stabilizing classification since the mid-2000s. While cladistics is the predominant classificatory system among paleontology professionals, the Linnean system is still in use, especially in works intended for popular distribution.


Fukuivenator ("hunter of Fukui Prefecture") is an extinct genus of coelurosaurian theropod dinosaur from the Early Cretaceous of Japan.


Halszkaraptor (; meaning "Halszka's seizer") is a genus of dromaeosaurid dinosaur from Mongolia that lived during the Late Cretaceous period. It contains only one known species, Halszkaraptor escuilliei.Scientists compared the type specimen (holotype) to the bones of extant crocodilians and aquatic birds, and found evidence of a semiaquatic lifestyle. A phylogenetic analysis revealed it was a member of the basal subfamily Halszkaraptorinae along with Mahakala and Hulsanpes.


Juravenator is a genus of small (75 cm long) coelurosaurian theropod dinosaur, which lived in the area which would someday become the top of the Franconian Jura of Germany, about 151 or 152 million years ago. It is known from a single, juvenile specimen.

Ksar Metlili Formation

The Ksar Metlili Formation is a geological formation in eastern High Atlas of Morocco, it is late Tithonian to Berriasian in age. It is approximately 80 metres (260 ft) thick and primarily consists of mudstone and sandstone, with thin calcareous beds. One of these calcareous beds near the middle of the sequence is an important microvertebrate locality.


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, 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 (see Eshanosaurus), and are regarded as surviving today as living birds.


Maniraptoromorpha is a clade of coelurosaurian theropod dinosaurs that includes the taxa Ornitholestes, Coelurus, and Maniraptoriformes. There has been several phylogenetic analyses that have shown support in the grouping of Maniraptoriformes with at least the aforementioned Ornitholestes. This group was named by Andrea Cau, who defined it as the "most inclusive clade containing Vultur gryphus Linnaeus, 1758, and excluding Tyrannosaurus rex Osborn, 1905."This group of coelurosaurs according to Cau (2018) has the following synapomorphies:

Keel or carinae in the postaxial cervical centra, absence of hyposphene-hypantra in caudal vertebrae (reversal to the plesiomorphic theropodan condition), a prominent dorsomedial process on the semilunate carpal, a convex ventral margin of the pubic foot, a subrectangular distal end of tibia and a sulcus along the posterior margin of the proximal end of fibula.


Nqwebasaurus (IPA: [ᵑǃʷɛbaˈsɔɹəs]; anglicized as or ) 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 lightning. Currently it is the only known coelurosaur discovered 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.


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 ago) of Western Laurasia (the area that was to become North America).

To date, Ornitholestes is known only from a single partial skeleton with a badly crushed skull found at the Bone Cabin Quarry near Medicine Bow, Wyoming, in 1900. It was described by Henry Fairfield Osborn in 1903. An incomplete hand was later attributed to Ornitholestes, although it now appears to belong to Tanycolagreus. The type (and only known) species is O. hermanni. The specific name honors the American Museum of Natural History preparator Adam Hermann.


The Ornithomimosauria, ornithomimosaurs ("bird-mimic lizards") or ostrich dinosaurs are theropod dinosaurs which bore a superficial resemblance to modern ostriches. 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. 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).


Pennaraptora (Latin penna "bird feather" + raptor "thief", from rapere "snatch"; a feathered bird-like predator) is a clade defined as the most recent common ancestor of Oviraptor philoceratops, Deinonychus antirrhopus, and Passer domesticus (the house sparrow), and all descendants thereof, by Foth et al., 2014. The earliest known definitive member of this clade is Anchiornis, from the late Jurassic period of China, about 160 million years ago.

The clade "Aviremigia" was conditionally proposed along with several other apomorphy-based clades relating to birds by Jacques Gauthier and Kevin de Queiroz in a 2001 paper. Their proposed definition for the group was "the clade stemming from the first panavian with ... remiges and rectrices, that is, enlarged, stiff-shafted, closed-vaned (= barbules bearing hooked distal pennulae), pennaceous feathers arising from the distal forelimbs and tail".


Sinocalliopteryx (meaning 'Chinese beautiful feather') is a genus of carnivorous compsognathid theropod dinosaurs from the Lower Cretaceous Yixian Formation of China (Jianshangou Beds, dating to 124.6 Ma).

While similar to the related Huaxiagnathus, Sinocalliopteryx were larger. The type specimen, at 2.37 meters (7.78 ft) in length, in 2007 was the largest known compsognathid exemplar. In 2012 an even larger specimen was reported.

Thomas R. Holtz Jr.

Thomas Richard Holtz Jr., Ph.D. 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: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages 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 on Discovery.Holtz is also the director of the Science and Global Change Program within the College Park Scholars living-learning community at the University of Maryland, College Park.


Unquillosaurus (meaning "Unquillo river lizard") is a genus of maniraptoran dinosaur from the Late Cretaceous Period, discovered in Argentina. Known only from a single fossilized pubis (a pelvic bone), its total body length may have reached 2 to 3 metres (6.6 to 9.8 ft).



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