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Maniraptorans Temporal range: Middle Jurassic–Present, 165–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Possible Early Jurassic record
Collage of six maniraptorans. From top left to right: the Zamyn Khondt oviraptorid, Sinornithosaurus, the Prince Creek troodontid, Patagonykus, a short-tailed albatross, and Nothronychus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Clade:	Saurischia
Clade:	Theropoda
Clade:	Maniraptoriformes
Clade:	Maniraptora Gauthier, 1986
Subgroups
†Fukuivenator †Juravenator? †Migmanychion †Ornitholestes? †Unquillosaurus †Yaverlandia †Alvarezsauroidea? †Therizinosauria Pennaraptora Foth, Tischlinger & Rauhut, 2014 †Oviraptorosauria †Scansoriopterygidae Paraves
Synonyms
Metornithes Perle et al., 1993

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 (see Eshanosaurus), and survive today as living birds.

Description

Maniraptorans are characterized by long arms and three-fingered hands (though reduced or fused in some lineages), as well as a "half-moon shaped" (semi-lunate) bone in the wrist (carpus). In 2004, Tom Holtz and Halszka Osmólska pointed out six other maniraptoran characters relating to specific details of the skeleton. Unlike most other saurischian dinosaurs, which have pubic bones that point forward, several groups of maniraptorans have an ornithischian-like backwards-pointing hip bone. A backward-pointing hip characterizes the therizinosaurs, dromaeosaurids, avialans, and some primitive troodontids. The fact that the backward-pointing hip is present in so many diverse maniraptoran groups has led most scientists to conclude that the "primitive" forward-pointing hip seen in advanced troodontids and oviraptorosaurs is an evolutionary reversal, and that these groups evolved from ancestors with backward-pointing hips.

Technical diagnosis

Holtz and Osmólska (2004) diagnosed the clade Maniraptora based on the following characters: reduced or absent olecranon process of the ulna, greater trochanter and cranial trochanter of the femur fused into a trochanteric crest. An elongated, backwards-pointing pubic bone is present in therizinosauroids, dromaeosaurids, avialans, and the basal troodontid Sinovenator, which suggests that the propubic condition in advanced troodontids and oviraptorosaurs is a reversal. Turner et al. (2007) named seven synapomorphies that diagnose Maniraptora.

Feathers and flight

Modern pennaceous feathers and remiges are known in the advanced maniraptoran group Aviremigia. More primitive maniraptorans, such as therizinosaurs (specifically Beipiaosaurus), preserve a combination of simple downy filaments and unique elongated quills. Simple feathers are known from more primitive coelurosaurs such as Sinosauropteryx prima, and possibly from even more distantly related species such as the ornithischian Tianyulong confuciusi and the flying pterosaurs. Thus it appears as if some form of feathers or down-like integument would have been present in all maniraptorans, at least when they were young.

Maniraptora is the only dinosaur group known to include flying members, though how far back in this lineage flight extends is controversial. Powered and/or gliding flight is believed to have been present in some types of non-avialan paravians, including dromaeosaurids, such as Rahonavis and Microraptor. Zhenyuanlong suni, a dromaeosaurid, was too heavy to fly but still had wings with feathers required for flying, which suggests its ancestors had the ability for aerial locomotion. Other groups, like the Oviraptorosauria who had a tail with a tail fan of feathers with caudal anatomy resembling a pygostyle, are not known to have been capable of flight, but some scientists, such as Gregory S. Paul, have suggested that they could be descended from ancestors which flew. Paul has gone as far as to propose that Therizinosauria, Alvarezsauroidea, and the non-maniraptoran group Ornithomimosauria also descended from flying ancestors.

Classification

The Maniraptora was originally named by Jacques Gauthier in 1986, for a branch-based clade defined as all dinosaurs closer to modern birds than to the ornithomimids. Gauthier noted that this group could be easily characterized by their long forelimbs and hands, which he interpreted as adaptations for grasping (hence the name Maniraptora, which means "hand snatchers" in relation to their 'seizing hands'). In 1994, Thomas R. Holtz attempted to define the group based on the characteristics of the hand and wrist alone (an apomorphy-based definition), and included the long, thin fingers, bowed, wing-like forearm bones, and half-moon shaped wrist bone as key characters. Most subsequent studies have not followed this definition, however, preferring the earlier branch-based definition.

The branch-based definition usually includes the major groups Dromaeosauridae, Troodontidae, Oviraptorosauria, Therizinosauria, and Avialae. Other taxa often found to be maniraptorans include the alvarezsaurs and Ornitholestes. Several taxa have been assigned to the Maniraptora more definitively, though their exact placement within the group remains uncertain. These forms include the scansoriopterygids, Pedopenna, and Yixianosaurus.

In 1993, Perle and colleagues coined the name Metornithes to include alvarezsaurids and modern birds, which the researchers believed were members of the Avialae. This group was defined as a clade by Luis Chiappe in 1995 as the last common ancestor of Mononykus and modern birds, and all its descendants.

Pennaraptora (Latin penna "bird feather" + raptor "thief", from rapere "snatch"; a feathered bird-like predator) is a clade within Maniraptora, 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 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". Ancestral morphology relating to pennaceous feathers suggests that basal species of Pennaraptora were capable of scansorial locomotion and gliding, and further evolution of said adaptation within the clade would eventually give rise to the origin of flight in avian species.

The following cladogram follows the results of a phylogenetic study by Cau (2020).

Alternative interpretations

In 2002, Czerkas and Yuan reported that some maniraptoran traits, such as a long, backwards-pointed pubis and short ischia were present in Scansoriopteryx, a scansoriopterygid. The authors considered it to be more primitive than true theropods, and hypothesized that maniraptorans may have branched off from theropods at a very early point, or may even have descended from pre-theropod dinosaurs. Zhang et al., in describing the closely related or conspecific specimen Epidendrosaurus (now considered a synonym of Scansoriopteryx), did not report any of the primitive traits mentioned by Czerkas and Yuan, but did find that the shoulder blade of Epidendrosaurus appeared primitive. Despite this, they placed Epidendrosaurus firmly within Maniraptora due to a number of synapomorphies.

Paleobiology

Diet

Scientists traditionally assumed that maniraptorans were ancestrally hypercarnivorous, that is, that most non-avialan species primarily ate and hunted only other vertebrates. However, a number of discoveries made during the first decade of the 21st century, as well as re-evaluation of older evidence, began to suggest that maniraptorans were a primarily omnivorous group, including a number of sub-groups that ate mainly plants, insects, or other food sources besides meat. Additionally, phylogenetic studies of maniraptoran relationships began to more consistently show that herbivorous or omnivorous groups were spread throughout the Maniraptora, rather than representing a single side-branch as previously thought. This led scientists such as Lindsay Zanno to conclude that the ancestral maniraptoran must have been omnivorous, giving rise to several purely herbivorous groups (such as the therizinosaurs, primitive oviraptorosaurs, and some avialans) and that, among non-avians, only one group reverted to pure carnivores (the dromaeosaurids). Most other groups fell somewhere in between the two extremes, with alvarezsaurids and some avialans being insectivorous, and with advanced oviraptorosaurs and troodontids being omnivorous.

Reproduction

A 2023 study analyzing fossil eggshells assigned to Troodon with clumped isotope thermometry found that Troodon, and likely other non-avian maniraptorans, had a slowed calcification of eggs akin to that of most reptiles. This contrasts with the rapid calcification of eggs found in modern birds, indicating that most maniraptorans aside from birds retained this basal trait. This would also indicate that most non-avian maniraptorans possessed two functional ovaries, contrasting with the one functional ovary in birds, and were thus limited in the numbers of eggs each individual could produce.

References

1. Zhang, H.; Wang, M.; Liu, X. (2008). "Constraints on the upper boundary age of the Tiaojishan Formation volcanic rocks in West Liaoning-North Hebei by LA-ICP-MS dating". Chinese Science Bulletin. 53 (22): 3574–3584. Bibcode:2008SciBu..53.3574Z. doi:10.1007/s11434-008-0287-4.
2. 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. Bibcode:2009Palgy..52..681B. doi:10.1111/j.1475-4983.2009.00887.x.
3. Meso, Jorge Gustavo; Pol, Diego; Chiappe, Luis; Qin, Zichuan; Díaz-Martínez, Ignacio; Gianechini, Federico; Apesteguía, Sebastián; Makovicky, Peter J.; Pittman, Michael. "Body size and evolutionary rate analyses reveal complex evolutionary history of Alvarezsauria". Cladistics. n/a (n/a). doi:10.1111/cla.12600. ISSN 1096-0031.
4. ^ Holtz, T.R. and Osmólska, H. (2004). "Saurischia." In Weishampel, Dodson and Osmólska (eds.), The Dinosauria, second edition. Berkeley: University of California Press.
5. ^ Turner, A.H.; Pol, D.; Clarke, J.A.; Erickson, G.M.; Norell, M. (2007). "A basal dromaeosaurid and size evolution preceding avian flight" (PDF). Science. 317 (5843): 1378–1381. Bibcode:2007Sci...317.1378T. doi:10.1126/science.1144066. PMID 17823350. S2CID 2519726.
6. Xu, X.; Tang, Z-L.; Wang, X-L. (1999). "A therizinosauroid dinosaur with integumentary structures from China". Nature. 399 (6734): 350–354. Bibcode:1999Natur.399..350X. doi:10.1038/20670. S2CID 204993327.
7. Xu X., Zheng X.-t. and You, H.-l. (2009). "A new feather type in a nonavian theropod and the early evolution of feathers." Proceedings of the National Academy of Sciences (Philadelphia), doi:10.1073/pnas.0810055106
8. Chiappe, L.M. (2007). Glorified Dinosaurs: The Origin and Early Evolution of Birds. Sydney: UNSW Press.
9. Scientists find a new dinosaur with well preserved, bird-like wings — but not for flight
10. Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
11. ^ Zanno, L.E., Gillette, D.D., Albright, L.B., and Titus, A.L. (2009). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution." Proceedings of the Royal Society B, Published online before print July 15, 2009, doi:10.1098/rspb.2009.1029.
12. Chiappe, L.M. (1995). "The first 85 million years of avian evolution". Nature. 378 (6555): 349–355. Bibcode:1995Natur.378..349C. doi:10.1038/378349a0. S2CID 4245171.
13. Foth, Christian; Tischlinger, Helmut; Rauhut, Oliver (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.
14. Gauthier, J. and de Queiroz, K. (2001). "Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name 'Aves'". Pp. 7-41 in Gauthier, J. and L.F. Gall (eds.), New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. New Haven: Peabody Museum of Natural History, Yale University. ISBN 0-912532-57-2.
15. Sorkin, Boris (2021-12-02). "Scansorial and aerial ability in Scansoriopterygidae and basal Oviraptorosauria". Historical Biology. 33 (12): 3202–3214. Bibcode:2021HBio...33.3202S. doi:10.1080/08912963.2020.1855158. ISSN 0891-2963.
16. Cau, Andrea. “The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory.” PeerJ vol. 8 e8672. 25 Feb. 2020, doi:10.7717/peerj.8672
17. Czerkas, S.A., and Yuan, C. (2002). "An arboreal maniraptoran from northeast China." Pp. 63-95 in Czerkas, S.J. (Ed.), Feathered Dinosaurs and the Origin of Flight. The Dinosaur Museum Journal 1. The Dinosaur Museum, Blanding, U.S.A. PDF abridged version
18. Zhang, F., Zhou, Z., Xu, X. & Wang, X. (2002). "A juvenile coelurosaurian theropod from China indicates arboreal habits." Naturwissenschaften, 89(9): 394-398. doi:10.1007 /s00114-002-0353-8.
19. Longrich, Nicholas R.; Currie, Philip J. (2009). "Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae". Cretaceous Research. 30 (1): 239–252. Bibcode:2009CrRes..30..239L. doi:10.1016/j.cretres.2008.07.005.
20. Holtz, T.R. Jr.; Brinkman, D.L.; Chandler, C.L. (1998). "Dental morphometrics and a possibly omnivorous feeding habit for the theropod dinosaur Troodon". GAIA. 15: 159–166.
21. Tagliavento, Mattia; Davies, Amelia J.; Bernecker, Miguel; Fiebig, Jens (April 3, 2023). "Evidence for heterothermic endothermy and reptile-like eggshell mineralization in Troodon, a non-avian maniraptoran theropod". PNAS. 120 (15): e2213987120. Bibcode:2023PNAS..12013987T. doi:10.1073/pnas.2213987120. PMC 10104568. PMID 37011196.

v t e Theropoda
Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Avemetatarsalia Clade: Dinosauria Avemetatarsalia see Avemetatarsalia Theropoda see below↓
Theropoda Theropoda Chindesaurus? Daemonosaurus? Eodromaeus? Erythrovenator Guaibasaurus? Nhandumirim? Tawa? Herrerasauria? Neotheropoda Dilophosaurus Dolichosuchus? Dracovenator Gojirasaurus Lepidus Liliensternus Lophostropheus Notatesseraeraptor Sarcosaurus Shuangbaisaurus Sinosaurus Tachiraptor Velocipes? Zupaysaurus Coelophysoidea Dracoraptor? Panguraptor Pendraig Podokesaurus Powellvenator Coelophysidae Camposaurus Coelophysis Lucianovenator Megapnosaurus? Panguraptor? Procompsognathus Pterospondylus? Segisaurus Averostra see below↓ Dubious neotheropods Teinurosaurus Halticosaurus
Averostra Averostra Dornraptor Lophostropheus? Ceratosauria Berberosaurus Fosterovenator? Jubbulpuria Lukousaurus? Saltriovenator Ceratosauridae Ceratosaurus Genyodectes Abelisauroidea Betasuchus? Lametasaurus? Ornithomimoides? Ozraptor? Noasauridae Afromimus? Austrocheirus? Berthasaura Compsosuchus? Dahalokely? Deltadromeus? Genusaurus? Kiyacursor Ligabueino Spinostropheus Elaphrosaurinae Elaphrosaurus Huinculsaurus Limusaurus Noasaurinae Laevisuchus Masiakasaurus Noasaurus Velocisaurus Vespersaurus Abelisauridae Chenanisaurus Dryptosauroides? Eoabelisaurus Indosuchus Kryptops Kurupi Rugops Spectrovenator Tarascosaurus Tralkasaurus Xenotarsosaurus Majungasaurinae Arcovenator Dahalokely? Genusaurus Indosaurus Majungasaurus Rahiolisaurus? Rajasaurus Carnotaurinae Majungasaurinae? Brachyrostra Dahalokely? Ekrixinatosaurus Elemgasem Guemesia Ilokelesia Llukalkan Niebla Rahiolisaurus? Skorpiovenator Thanos Furileusauria Abelisaurus Aucasaurus Caletodraco Carnotaurus Elemgasem? Koleken Pycnonemosaurus Quilmesaurus Viavenator Tetanurae see below↓
Tetanurae Tetanurae Calamospondylus? Chienkosaurus? Chilesaurus? Chuandongocoelurus Cruxicheiros? Cryolophosaurus Kaijiangosaurus Kayentavenator Monolophosaurus Pandoravenator Sinosaurus? Szechuanosaurus Vectaerovenator Orionides Megalosauroidea Yunyangosaurus Piatnitzkysauridae Condorraptor Marshosaurus Piatnitzkysaurus Xuanhanosaurus? Megalosauria Streptospondylus? Megalosauridae Eustreptospondylus Megalosaurinae Duriavenator Megalosaurus Torvosaurus Wiehenvenator Afrovenatorinae Afrovenator Dubreuillosaurus Leshansaurus Magnosaurus Piveteausaurus Poekilopleuron? Spinosauridae Camarillasaurus? Cristatusaurus Iberospinus Ostafrikasaurus? Baryonychinae Baryonyx Protathlitis Riojavenatrix Suchosaurus? Vallibonavenatrix? Ceratosuchopsini Ceratosuchops Riparovenator Suchomimus Spinosaurinae Camarillasaurus? Ichthyovenator Irritator Oxalaia Siamosaurus Spinosaurini Sigilmassasaurus Spinosaurus Avetheropoda see below↓
Avetheropoda Avetheropoda Gasosaurus? Lourinhanosaurus? Carnosauria Altispinax Monolophosaurus? Megalosauridae? Spinosauridae? Allosauroidea Asfaltovenator Erectopus Xuanhanosaurus? Piatnitzkysauridae? Metriacanthosauridae Xuanhanosaurus? Yangchuanosaurus Metriacanthosaurinae Alpkarakush Metriacanthosaurus Shidaisaurus Siamotyrannus Sinraptor Allosauria Allosauridae Allosaurus Antrodemus? Epanterias? Saurophaganax? Carcharodontosauria Datanglong Lusovenator Siamraptor Siats? Ulughbegsaurus? Veterupristisaurus Neovenatoridae Chilantaisaurus? Gualicho? Neovenator Siats? Megaraptora? Carcharodontosauridae Acrocanthosaurus Altispinax? Concavenator Eocarcharia Kelmayisaurus Lajasvenator Sauroniops Shaochilong Ulughbegsaurus? Veterupristisaurus Carcharodontosaurinae Carcharodontosaurus Giganotosaurini Giganotosaurus Mapusaurus Meraxes Taurovenator Tyrannotitan Megaraptora? Aoniraptor Bahariasaurus? Deltadromeus? Fukuiraptor Phuwiangvenator Rapator Siats? Chilantaisaurus? Megaraptoridae Aerosteon Australovenator Maip Megaraptor Murusraptor Orkoraptor Tratayenia Coelurosauria see below↓
Coelurosauria Coelurosauria Aratasaurus Asiamericana Bahariasaurus? Bicentenaria Chilantaisaurus? Gualicho? Richardoestesia Sciurumimus? Vayuraptor Xinjiangovenator Zuolong Tyrannoraptora Tyrannosauroidea Chingkankousaurus Labocania? Coeluridae? Coelurus Tanycolagreus? Proceratosauridae Guanlong Kileskus Proceratosaurus Sinotyrannus Yutyrannus Pantyrannosauria Alectrosaurus Aviatyrannis Bagaraatan? Dilong Eotyrannus Jinbeisaurus Juratyrant Moros Santanaraptor Stokesosaurus Suskityrannus Timimus? Timurlengia Xiongguanlong Megaraptora? Eutyrannosauria Appalachiosaurus Bistahieversor? Dryptosaurus Tyrannosauridae Albertosaurinae Albertosaurus Gorgosaurus Tyrannosaurinae Asiatyrannus Nanuqsaurus Alioramini Alioramus Qianzhousaurus Daspletosaurini Daspletosaurus Thanatotheristes Teratophoneini Bistahieversor? Dynamoterror Labocania? Lythronax Teratophoneus Tyrannosaurini Tarbosaurus Tyrannosaurus Zhuchengtyrannus Maniraptoromorpha see below↓ Dubious coelurosaurs Iliosuchus Kakuru Phaedrolosaurus Shanyangosaurus
Maniraptoromorpha Maniraptoromorpha Coelurus? Juravenator Ornitholestes Neocoelurosauria Aniksosaurus Compsognathidae Aniksosaurus? Aristosuchus Beipiaognathus? Compsognathus Huaxiagnathus Mirischia Scipionyx? Sinocalliopteryx Sinosauropteryx Xunmenglong Maniraptoriformes Compsognathidae? Ornithomimosauria Arkansaurus Aviatyrannis? Calamosaurus? Hexing Kinnareemimus Nedcolbertia Nqwebasaurus Thecocoelurus? Valdoraptor Macrocheiriformes Harpymimus Pelecanimimus Shenzhousaurus Deinocheiridae Beishanlong Deinocheirus Garudimimus Harpymimus? Paraxenisaurus? Tyrannomimus Ornithomimidae Aepyornithomimus Anserimimus Archaeornithomimus "Coelosaurus" Dromiceiomimus Gallimimus Ornithomimus Qiupalong Rativates Sinornithomimus Struthiomimus Tototlmimus Maniraptora see Maniraptora includes birds

v t e Maniraptora
Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: Theropoda Clade: Maniraptoriformes Avemetatarsalia see Avemetatarsalia Theropoda see Theropoda Maniraptora see below↓
Maniraptora Maniraptora †Elopteryx? †Fukuivenator? †Kakuru? †Migmanychion †Yaverlandia? †Alvarezsauroidea Achillesaurus Aorun? Bannykus Bonapartenykus Haplocheirus Patagonykus Shishugounykus Tugulusaurus? Xiyunykus Alvarezsauridae Achillesaurus? Alnashetri Alvarezsaurus Bonapartenykus? Bradycneme Heptasteornis Patagonykus? Parvicursorinae Dzharaonyx Jaculinykus Khulsanurus Kol? Nemegtonykus Ondogurvel Parvicursor Qiupanykus Trierarchuncus Ceratonykini Albinykus Ceratonykus Xixianykus Mononykini Albertonykus Linhenykus Mononykus Shuvuuia †Therizinosauria Eshanosaurus? Falcarius Fukuivenator? Jianchangosaurus Lingyuanosaurus Therizinosauroidea Alxasaurus Beipiaosaurus Enigmosaurus Martharaptor Suzhousaurus Therizinosauridae Erliansaurus Erlikosaurus Nanshiungosaurus Neimongosaurus Nothronychus Paralitherizinosaurus Segnosaurus Therizinosaurus Pennaraptora †Oviraptorosauria see Oviraptorosauria Paraves see below↓
Paraves Paraves †Palaeopteryx? †Pneumatoraptor †Rahonavis †Scansoriopterygidae? Ambopteryx Epidexipteryx Scansoriopteryx Yi †Anchiornithidae Anchiornis Aurornis Caihong Eosinopteryx Fujianvenator Liaoningvenator? Ostromia Pedopenna Serikornis Xiaotingia Yixianosaurus †Archaeopterygidae Alcmonavis? Archaeopteryx Wellnhoferia Anchiornithidae? †Dromaeosauridae see Dromaeosauridae †Troodontidae see Troodontidae Avialae †Alcmonavis †Balaur? †Cretaaviculus? †Fukuipteryx †Overoraptor †Rahonavis? †Yandangornis †Zhongornis †Anchiornithidae? †Archaeopterygidae? †Scansoriopterygidae? †Jeholornithiformes Dalianraptor? Jeholornis Jixiangornis? Kompsornis Neimengornis Pygostylia †"Proornis" †Omnivoropterygidae? Omnivoropteryx Sapeornis †Confuciusornithidae Changchengornis Confuciusornis Eoconfuciusornis Yangavis †Jinguofortisidae Chongmingia Cratonavis Jinguofortis Ornithothoraces †Enantiornithes see Enantiornithes Euornithes see below↓
Euornithes Euornithes †Archaeorhynchus †Bellulornis †Brevidentavis †Changmaornis †Changzuiornis †Chaoyangia †Dingavis †Eogranivora †Gargantuavis? †Hollanda †Horezmavis †Jianchangornis †Jiuquanornis †Juehuaornis †Kaririavis †Meemannavis †Platanavis †Vorona †Wyleyia? †Xinghaiornis †Yumenornis †Zhongjianornis †Schizoouridae Mengciusornis Schizooura †Patagopterygiformes Alamitornis Kuszholia? Patagopteryx †Ambiortiformes Ambiortus Apsaravis? Palintropus? †Hongshanornithidae Archaeornithura Hongshanornis Longicrusavis Parahongshanornis Tianyuornis †Songlingornithidae Hollanda? Piscivoravis? Songlingornis Yanornis? Yixianornis? †Yanornithidae Abitusavis Similiyanornis Yanornis †Gansuidae? Changzuiornis Gansus Hollanda? Iteravis Khinganornis Shuilingornis Ornithurae †Antarcticavis? †Apatornis †Cerebavis †Gallornis †Guildavis †Iaceornis †Kookne †Limenavis †Tingmiatornis †Ichthyornithes Ichthyornis Janavis †Hesperornithes Baptornis Brodavis Chupkaornis Enaliornis Judinornis Pasquiaornis Potamornis Hesperornithidae Asiahesperornis Canadaga Fumicollis Hesperornis Parahesperornis †Vegaviidae Australornis? Maaqwi? Neogaeornis? Polarornis Vegavis †Cimolopterygidae Ceramornis Cimolopteryx Lamarqueavis? Aves / Neornithes see below↓
Aves / Neornithes Palaeognathae see Palaeognathae Neognathae Neoaves Galloanserae Anserimorphae see Anserimorphae Pangalliformes see Pangalliformes Incertae sedis †Asteriornis †Perplexicervix †Qinornis? †Dromornithidae Barawertornis Dromornis Genyornis Ilbandornis †Gastornithiformes Brontornis? Gastornis Dromornithidae? †Pelagornithidae Aequornis Caspiodontornis Cyphornis Dasornis Gigantornis Laornis? Lutetodontopteris Macrodontopteryx Manu? Odontopteryx Odontoptila Osteodontornis Palaeochenoides Pelagornis Proceriavis? Protodontopteryx Protopelicanus? Pseudodontornis Tympanonesiotes
See also: Archaeornithes Carinatae Deinonychosauria Odontognathae Odontornithes Sauriurae Unenlagiidae Category

v t e Fins, limbs and wings
Fins	Aquatic locomotion Cephalopod fin Fish locomotion Fin and flipper locomotion Caudal fin Dorsal fin Fish fin Flipper Lobe-finned fish Ray-finned fish Pectoral fins Pelvic fin
Limbs	Limb development Limb morphology digitigrade plantigrade unguligrade uniped biped facultative biped triped quadruped Arthropod Cephalopod Tetrapod dactyly Digit Webbed foot
Wings	Flying and gliding animals Bat wing Bird wing keel skeleton feathers Insect wing Pterosaur wing Wingspan
Evolution	Evolution of fish Evolution of tetrapods Evolution of birds Origin of birds Origin of avian flight Evolution of cetaceans Comparative anatomy Convergent evolution Analogous structures Homologous structures
Related	Animal locomotion Gait Robot locomotion Samara Terrestrial locomotion Tradeoffs for locomotion in air and water Rotating locomotion Undulatory locomotion

• Birds
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• Maniraptora
• Coelurosaurs
• Oxfordian first appearances
• Extant Late Jurassic first appearances
• Taxa named by Jacques Gauthier