Tyrannosauroidea

Skeletal mount of Tyrannosaurus rex, copyright Scott Robert Anselmo.

Belongs within: Coelurosauria.

The Tyrannosauroidea are a clade of theropod dinosaurs known from the Late Jurassic and Cretaceous of Asia and North America including some of the largest of all carnivorous dinosaurs, as well as a number of smaller basal forms. Yutyrannus huali from the lower Cretaceous of China is the largest known dinosaur to preserve direct evidence of a covering of feathers (Xu et al. 2012).

<==Tyrannosauroidea XN04
    |--Kileskus XW12
    |--Dilong Xu, Norell et al. 2004 XW12, XN04
    |    `--*D. paradoxus Xu, Norell et al. 2004 XN04
    |--Stokesosaurus Madsen 1974 XW12, H04
    |    |--S. clevelandi Madsen 1974 H04
    |    `--S. langhami XC09
    |--+--Sinotyrannus XW12
    |  `--+--Guanlong wucaii XW12
    |     `--Proceratosaurus Huene 1926 XW12, HMC04
    |          `--P. bradleyi (Woodward 1910) [=Megalosaurus bradleyi] HMC04
    `--+--Yutyrannus Xu, Wang et al. 2012 XW12
       |    `--*Y. huali Xu, Wang et al. 2012 XW12
       `--+--Eotyrannus Hutt, Naish et al. 2001 XW12, H04
          |    `--E. lengi Hutt, Naish et al. 2001 H04
          `--+--Xiongguanglong baimoensis XW12
             `--+--Dryptosaurus Marsh 1877 XW12, H04
                |    `--D. aquilunguis (Cope 1866) [=Laelaps aquilunguis] H04
                `--+--Raptorex kriegsteini XW12
                   `--+--Appalachiosaurus Carr, Williamson & Schwimmer 2005 XW12, D07
                      |    `--A. montgomeriensis D07
                      `--+--Bistahieversor XW12
                         `--Tyrannosauridae [Deinodontidae, Dinodontidae] XN04
                              |  i. s.: Aublysodon Leidy 1868 [Aublysodontinae] P88
                              |           |--A. amplus Marsh 1892 (n. d.) H04
                              |           |--A. cristatus Marsh 1892 (n. d.) H04
                              |           |--A. lateralis Cope 1876 (n. d.) H04
                              |           `--A. mirandus Leidy 1868 (n. d.) H04
                              |         Labocania Molnar 1974 N85, H04
                              |           `--L. anomala Molnar 1974 H04
                              |         Prodeinodon N85
                              |           |--P. kwangshiensis Hou, Yeh & Zhao 1975 (n. d.) HMC04
                              |           `--P. mongoliensis Osborn 1924 (n. d.) HMC04
                              |         Alectrosaurus Gilmore 1933 H04
                              |           `--A. olseni Gilmore 1933 H04 [=Albertosaurus (Alectrosaurus) olseni P88]
                              |--Albertosaurinae H04
                              |    |--Albertosaurus Osborn 1905 XW12, H04
                              |    |    `--*A. sarcophagus Osborn 1905 N85 [incl. A. arctunguis Parks 1928 H04]
                              |    `--Gorgosaurus Lambe 1914 [incl. Deinodon Leidy 1856] H04
                              |         `--G. libratus Lambe 1914 H04 (see below for synonymy)
                              `--Tyrannosaurinae H04
                                   |--Alioramus Kurzanov 1976 XW12, H04
                                   |    |--A. altai Brusatte, Carr et al. 2009 BC09
                                   |    `--A. remotus Kurzanov 1976 H04
                                   `--+--Teratophoneus XW12
                                      `--+--Daspletosaurus Russell 1970 H04
                                         |    `--D. torosus Russell 1970 H04 (see below for synonymy)
                                         `--+--Tarbosaurus Maleev 1955 (see below for synonymy) H04
                                            |    `--T. bataar (Maleev 1955) H04 (see below for synonymy)
                                            `--+--Nanotyrannus Bakker, Williams & Currie 1988 XN04, H04
                                               |    `--N. lancensis (Gilmore 1946) H04 (see below for synonymy)
                                               `--Tyrannosaurus Osborn 1905 XW12, H04 (see below for synonymy)
                                                    |--T. lanpingensis Yeh 1975 (n. d.) H04
                                                    `--T. rex Osborn 1905 H04 (see below for synonymy)

Tyrannosauroidea incertae sedis:
  Mirischia asymmetrica N10
  Bagaraatan Osmólska 1996 H04
    `--B. ostromi Osmólska 1996 H04
  ‘Ornithomimus’ grandis Marsh 1890 H04
  ‘Chilantaisaurus’ maortuensis Hu 1964 H04
  Santanaraptor Kellner 1999 H04
    `--S. placidus Kellner 1999 H04
  Laelaps macropus Cope 1868 (n. d.) H04
  Diplotomodon Leidy 1868 (n. d.) H04
    `--*D. horrificus (Leidy 1868) (n. d.) [=Tomodon horrificus] H04
  Itemirus Kurzanov 1976 [Itemiridae] H04
    `--I. medullaris Kurzanov 1976 H04
  Iliosuchus Huene 1932 D07, HMC04 [Iliosuchidae]
    `--I. incognitus Huene 1932 HMC04

Daspletosaurus torosus Russell 1970 H04 [=Tyrannosaurus (Daspletosaurus) torosus P88]

Gorgosaurus libratus Lambe 1914 H04 [=Albertosaurus libratus N85; incl. Laelaps falculus Cope 1876 H04, Dromaeosaurus falculus O69, L. hazenianus Cope 1876 H04, Deinodon horridus Leidy 1856 (n. d.) H04, L. incrassatus Cope 1876 H04, Dryptosaurus incrassatus P88, D. kenabekides Hay 1899 H04, Gorgosaurus sternbergi Matthew & Brown 1923 H04, Albertosaurus sternbergi P88]

Nanotyrannus lancensis (Gilmore 1946) H04 [=Gorgosaurus lancensis H04, Albertosaurus (Nanotyrannus) lancensis P88]

Tarbosaurus Maleev 1955 XW12, H04 [incl. Chingkankosaurus Young 1958 H04, Maleevosaurus Carpenter 1992 H04, Shanshanosaurus Dong 1977 H04]

Tarbosaurus bataar (Maleev 1955) H04 [=Tyrannosaurus bataar H04; incl. Ta. efremovi Maleev 1955 H04, Chingkankousaurus fragilis Young 1958 H04, Shanshanosaurus huoyanshanensis Dong 1977 H04, Aublysodon huoyanshanensis P88, Gorgosaurus lancinator Maleev 1955 H04, Tyrannnosaurus luanchuanensis Dong 1979 H04, G. novojilovi Maleev 1955 H04, Albertosaurus novojilovi H04, Aublysodon novojilovi H04, Deinodon novojilovi H04, Maleevosaurus novojilovi H04, Albertosaurus periculosus Riabinin 1930 H04, Tyrannosaurus turpanensis Zhai, Zheng & Tong 1978 H04]

Tyrannosaurus Osborn 1905 XW12, H04 [incl. Dynamosaurus Osborn 1905 H04, Manospondylus Cope 1892 H04]

Tyrannosaurus rex Osborn 1905 H04 [incl. Manospondylus gigas Cope 1892 H04, Dynamosaurus imperiosus Osborn 1905 H04, Albertosaurus megagracilis Paul 1988 H04, Aublysodon molnaris Paul 1988 H04, P88]

*Type species of generic name indicated

REFERENCES

[BC09] Brusatte, S. L., T. D. Carr, G. M. Erickson, G. S. Bever & M. A. Norell. 2009. A long-snouted, multihorned tyrannosaurid from the Late Cretaceous of Mongolia. Proceedings of the National Academy of Sciences of the USA 106 (41): 17261–17266.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[H04] Holtz, T. R. 2004. Tyrannosauroidea. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 111–136. University of California Press: Berkeley.

[HMC04] Holtz, T. R., Jr, R. E. Molnar & P. J. Currie. 2004. Basal Tetanurae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 71–110. University of California Press: Berkeley.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[O69] Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History 30: 1–165.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[XC09] Xu, X., J. M. Clark, J. Mo, J. Choiniere, C. A. Forster, G. M. Erickson, D. W. E. Hone, C. Sullivan, D. A. Eberth, S. Nesbitt, Q. Zhao, R. Hernandez, C.-K. Jia, F.-L. Han & Y. Guo. 2009. A Jurassic ceratosaur from China helps clarify avian digital homologies. Nature 459: 940–944.

[XN04] Xu, X., M. A. Norell, X. Kuang, X. Wang, Q. Zhao & C. Jia. 2004. Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids. Nature 431: 680–684.

[XW12] Xu, X., K. Wang, K. Zhang, Q. Ma, L. Xing, C. Sullivan, D. Hu, S. Cheng & S. Wang. 2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature 484: 92–95.

Last updated: 11 July 2018.

Ornithurae

Reconstruction of Enaliornis barretti, copyright Nobu Tamura.

Belongs within: Ornithothoraces.
Contains: Aves.

The Ornithurae are a clade of birds including all modern species together with related fossil forms possessing a very short pygostyle and completely fused tarsometatarsus (Zhou & Zhang 2005). Basal members of this clade include the famous toothed seabirds Ichthyornis and Hesperornis from the Late Cretaceous of North America.

Synapomorphies (from Zhou & Zhang 2005): Pygostyle very short; coracoid strut-like with well-developed procoracoid process; furcula U-shaped; sternum elongate; major digit with first phalanx significantly laterally expanded, tightly abutted by reduced minor digit; tarsometatarsus completely fused.

<==Ornithurae [Ichthyornithiformes, Odontoholomorphae, Odontornithes, Odontotormae, Plataleae] C04
    |--Hongshanornis Zhou & Zhang 2005 ZZ05
    |    `--*H. longicresta Zhou & Zhang 2005 ZZ05
    `--+--+--Liaoningornis Hou 1997 ZZ05, P04
       |  |    `--L. longidigitus Hou 1997 P04
       |  `--+--Yixianornis grabaui ZZ05
       |     `--Yanornis Zhou & Zhang 2001 ZZ05, D07
       |          |--Y. chaoyangensis ZZ05
       |          `--Y. martini D07
       `--+--Apsaravis Norell & Clarke 2001 ZZ05, NC01 [Apsaravidae, Apsaraviformes]
          |    `--*A. ukhaana Norell & Clarke 2001 NC01
          `--+--+--Ambiortus Kurochkin 1982 P04
             |  |    `--A. dementjevi Kurochkin 1982 [incl. Holbotia ponomarenkoi Kurochkin 1982] P04
             |  `--Carinatae P04
             |       |--Ichthyornis Marsh 1872 C04 (see below for synonymy)
             |       |    `--I. anceps (Marsh 1872) (see below for synonymy) C04
             |       `--+--Aves C04
             |          |--Apatornis Marsh 1873 [Apatornithidae] C04
             |          |    `--*A. celer Marsh 1873 C04 [=Ichthyornis celer P04]
             |          |--Iaceornis Clarke 2004 C04
             |          |    `--*I. marshi Clarke 2004 C04
             |          `--Guildavis Clarke 2004 C04
             |               `--*G. tener (Marsh 1880) [=Ichthyornis tener] C04
             `--Hesperornithes [Hesperornithiformes, Odontoholcae, Odontolcae] C04
                  |  i. s.: Asiahesperornis Nessov & Prizemlin 1991 P04
                  |           `--A. bazhanovi Nessov & Prizemlin 1991 P04
                  |         Judinornis Nessov & Borkin 1983 P04
                  |           `--J. nogontsavensis Nessov & Borkin 1983 P04
                  |         Pasquiaornis Tokaryk, Cumbaa & Storer 1997 P04
                  |           |--P. hardiei Tokaryk, Cumbaa & Storer 1998 P04
                  |           `--P. tankei Tokaryk, Cumbaa & Storer 1997 P04
                  |         Potamornis Elżanowski, Paul & Stidham 1999 P04
                  |           `--P. skutchi Elżanowski, Paul & Stidham 1999 P04
                  |--Enaliornis Seeley 1876 P04 (see below for synonymy)
                  |    |--E. barretti Seeley 1876 [=Palaeocolymbus barretti] P04
                  |    |--E. sedgwicki (Seeley 1876) [=Pelagornis sedgwicki] P04
                  |    `--E. seeleyi D07
                  `--+--Baptornis Marsh 1877 P04 [Baptornithidae FP64]
                     |    `--B. advenus Marsh 1877 C04
                     `--Hesperornithidae C04
                          |--Coniornis Marsh 1893 FP64, P04
                          |    `--C. altus Marsh 1893 [incl. Hesperornis montanus Shufeldt 1915] P04
                          |--Lestornis Marsh 1876 C04, P04
                          |    `--L. crassipes Marsh 1876 C04 [=Hesperornis crassipes P04]
                          |--Parahesperornis Martin 1983 C04, P04
                          |    `--P. alexi Martin 1983 P04
                          `--Hesperornis Marsh 1872 [incl. Hargeria Lucas 1903] P04
                               |--H. gracilis Marsh 1876 C04 [=Hargeria gracilis P04]
                               `--H. regalis Marsh 1872 P04

Ornithurae incertae sedis:
  Chaoyangia Hou & Zhang 1993 P04
    `--C. beishanensis Hou & Zhang 1993 P04
  Gansus Hou & Liu 1984 P04
    `--G. yumenensis Hou & Liu 1984 P04
  Gargantuavis Buffetaut & Le Loeuff 1998 P04
    `--G. philoinos Buffetaut & Le Loeuff 1998 P04
  Horezmavis Nessov & Borkin 1983 P04
    `--H. eocretacea Nessov & Borkin 1983 P04
  Limenavis Clarke & Chiappe 2001 P04
    `--L. patagonica Clarke & Chiappe 2001 P04
  Qinornis paleocenica Xue 1995 M09
  Cimolopteryx retusus C04

Enaliornis Seeley 1876 P04 [incl. Palaeocolymbus Seeley 1876 P04, Pelagornis Seeley 1876 non Lartet 1857 P04; Enaliornithidae FP64]

Ichthyornis Marsh 1872 C04 [incl. Angelinornis Kashin 1972 P04, Colonosaurus Marsh 1872 P04; Ichthyornidae, Ichthyornithes, Ichthyornithidae]

Ichthyornis anceps (Marsh 1872) [=Graculavus anceps; incl. G. agilis Marsh 1873, Ichthyornis agilis, Plegadornis antecessor Wetmore 1962, Angelinornis antecessor, Ichthyornis antecessor, *I. dispar Marsh 1872, Colonosaurus mudgei Marsh 1872, I. validus Marsh 1880, I. victor Marsh 1876] C04

*Type species of generic name indicated

REFERENCES

[C04] Clarke, J. A. 2004. Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae). Bulletin of the American Museum of Natural History 286: 1–179.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[NC01] Norell, M. A., & J. A. Clarke. 2001. Fossil that fills a critical gap in avian evolution. Nature 409: 181–184.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[ZZ05] Zhou, Z., & F. Zhang. 2005. Discovery of an ornithurine bird and its implications for Early Cretaceous avian radiation. Proceedings of the National Academy of Sciences of the USA 102: 18998–19002.

Last updated: 6 August 2017.

Troodontidae

Skeletal reconstruction of Saurornithoides junior, copyright Jaime A. Headden.

Belongs within: Maniraptora.

The Troodontidae are a group of small, cursorial theropods known from the Cretaceous of Asia and North America with a single possible representative, Koparion douglassi, also known from the Upper Jurassic (Makovicky & Norell 2004). Certain fragmentary taxa from the late Cretaceous of Europe, such as Bradycneme draculae and Elopteryx nopcsai, have also been suggested to be troodontids but are not universally accepted as such.

Characters (from Makovicky & Norell 2004, possibly excluding Jinfengopteryx): Dentary bearing groove on at least part of labial face containing exits for neurovascular foramina; quadrate with pneumatic foramen present caudally; basicranium with basisphenoid recess absent; internarial bar dorsoventrally flattened; large number of teeth present; mesial dentary teeth closely packed; laterosphenoid with depression present on ventral surface of postorbital process; reduced basal tubera usually lying directly ventral to occipital condyle; transverse processes elongate; distal caudals with sulcus on dorsal midline in place of neural spine; metatarsus asymmetrical with slender metatarsal II markedly shorter than metatarsals III and IV, metatarsal IV robust.

<==Troodontidae [Saurornithoididae]
    |--Jinfengopteryx GC13
    `--+--Mei Xu & Norell 2004 GC13, XN04
       |    `--*M. long Xu & Norell 2004 XN04
       `--+--Sinovenator Xu, Norell et al. 2002 MN04
          |    `--*S. changii Xu, Norell et al. 2002 XN02
          `--+--Sinusonasus Xu & Wang 2004 GC13, XW04
             |    `--*S. magnodens Xu & Wang 2004 XW04
             `--+--Byronosaurus Norell, Makovicky & Clark 2000 GC13, MN04
                |    `--B. jaffei Norell, Makovicky & Clark 2000 MN04
                `--+--Sinornithoides Russell & Dong 1993 GC13, MN04
                   |    `--S. youngi Russell & Dong 1993 MN04
                   `--+--Zanabazar GC13
                      |--Troodon Leidy 1856 (see below for synonymy) MN04
                      |    |--*T. formosus Leidy 1856 N85 (see below for synonymy)
                      |    |--‘Pectinodon’ asiamericanus Nessov 1995 (n. d.) MN04
                      |    `--T. isfarensis Nessov 1995 (n. d.) MN04
                      `--Saurornithoides Osborn 1924 MN04
                           |--S. asiamericanus D07
                           |--S. isfarensis D07
                           |--S. junior Barsbold 1974 MN04
                           `--S. mongoliensis Osborn 1924 MN04 [=Troodon mongoliensis P88]

Troodontidae incertae sedis:
  Borogovia Osmólska 1987 MN04
    `--B. gracilicrus Osmólska 1987 MN04
  Bradycneme draculae Harrison & Walker 1975 (n. d.) MN04
  Heptasteornis Harrison & Walker 1975 (n. d.) MN04
    `--H. andrewsi Harrison & Walker 1975 [=Troodon andrewsi] P88
  Koparion Chure 1994 H98
    `--K. douglassi Chure 1994 MN04
  Tochisaurus Kurzanov & Osmólska 1991 MN04
    `--T. nemegtensis Kurzanov & Osmólska 1991 MN04
  Elopteryx [Elopterygidae] MN04
    `--E. nopcsai Andrews 1913 (n. d.) MN04
  ‘Laelaps’ cristatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus cristatus O69]
  ‘Laelaps’ explanatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus explanatus O69]
  ‘Laelaps’ laevifrons Cope 1876 (n. d.) MN04 [=Dromaeosaurus laevifrons O69]
  Paronychodon lacustris Cope 1876 (n. d.) MN04
  Zapsalis abradens Cope 1876 (n. d.) MN04

Troodon Leidy 1856 [incl. Pectinodon Carpenter 1982, Polyodontosaurus Gilmore 1932, Stenonychosaurus Sternberg 1932] MN04

*Troodon formosus Leidy 1856 N85 [incl. Pectinodon bakkeri Carpenter 1982 MN04, Polyodontosaurus grandis Gilmore 1932 MN04, Stenonychosaurus inequalis Sternberg 1932 MN04]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[H98] Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5–61.

[MN04] Makovicky, P. J., & M. A. Norell. 2004. Troodontidae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 184–195. University of California Press: Berkeley.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[O69] Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History 30: 1–165.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[XN04] Xu, X., & M. A. Norell. 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature 431: 838–841.

[XN02] Xu, X., M. A. Norell, X.-L. Wang, P. J. Makovicky & X.-C. Wu. 2002. A basal troodontid from the Early Cretaceous of China. Nature 415: 780–784.

[XW04] Xu X. & Wang X. 2004. A new troodontid (Theropoda: Troodontidae) from the Lower Cretaceous Yixian Formation of western Liaoning, China. Acta Geologica Sinica (English Edition) 78: 22–26.

Last updated: 3 July 2018.

Oviraptorosauria

Reconstruction of Caudipteryx zoui, by Matt Martyniuk.

Belongs within: Maniraptora.

The Oviraptorosauria are a group of cursorial theropods known from the Cretaceous of Eurasia and North America. Mostly less than two metres in length, they are characterised by deep, short skulls with the dentition reduced or absent. The ecology of oviraptorosaurs has been subject to much debate; they may have included herbivorous or omnivorous forms.

The majority of oviraptorosaurs belong to a clade Caenagnathoidea, which has been defined as the least inclusive clade including Oviraptor philoceratops and Caenagnathus collinsi. This clade is supported by several synapomorphies, including the presence of two longitudinal ridges and a toothlike ventral process on the palatal shelf of the maxilla, and a jaw joint close to the midline of the skull (Osmólska et al. 2004). Non-caenagnathoid oviraptorosaurs include the early Cretaceous Caudipteryx, the type species of which, C. zoui, is known from fossils in which a fan of feathers is preserved at the end of the tail.

Synapomorphies (from Osmólska et al. 2004): Premaxilla with crenulated ventral margin; parietal at least as long as frontal; ascending process of quadratojugal bordering more than three-quarters of infratemporal fenestra; U-shaped mandibular symphysis present; dentary edentulous; pubic foot with cranial process longer than caudal process.

<==Oviraptorosauria
    |  i. s.: Thecocoelurus Huene 1923 OCB04
    |           `--*T. daviesi (Seeley 1888) OCB04 [=Thecospondylus daviesi OCB04, Coelurus daviesi N02]
    |--+--Protarchaeopteryx Ji & Ji 1997 GC13, P04
    |  |    `--P. robusta Ji & Ji 1997 P04
    |  `--Incisivosaurus Xu, Cheng et al. 2002 XT07, OCB04
    |       `--*I. gauthieri Xu, Cheng et al. 2002 OCB04
    `--+--Similicaudipteryx GC13
       `--+--Avimimus Kurzanov 1981 [Avimimidae] OCB04
          |    `--A. portentosus Kurzanov 1981 OCB04
          `--+--Caudipteryx Ji, Currie et al. 1998 XT07, OCB04 [Caudipterygidae]
             |    |--C. dongi Zhou & Wang 2000 OCB04
             |    `--C. zoui Ji, Currie et al. 1998 OCB04
             |--Microvenator Ostrom 1970 GC13, OCB04
             |    `--M. celer Ostrom 1970 OCB04
             `--Caenagnathoidea [Caenagnathinae, Oviraptoroidea] OCB04
                  |  i. s.: Shixinggia oblita N10
                  |--Caenagnathidae [Elmisauridae] OCB04
                  |    |--Hagryphus giganteus N10
                  |    |--Elmisaurus Osmólska 1981 MOW02, OCB04
                  |    |    `--E. rarus Osmólska 1981 OCB04 [=Chirostenotes rarus P88]
                  |    |--Caenagnathasia Currie, Godfrey & Nessov 1993 OCB04
                  |    |    `--C. martinsoni Currie, Godfrey & Nessov 1993 1994 OCB04
                  |    |--Nomingia Barsbold, Osmólska et al. 2000 OCB04
                  |    |    `--N. gobiensis Barsbold, Osmólska et al. 2000 OCB04
                  |    `--Chirostenotes Gilmore 1924 OCB04 (see below for synonymy)
                  |         |--C. elegans (Parks 1933) OCB04 (see below for synonymy)
                  |         `--C. pergracilis Gilmore 1924 (see below for synonymy) OCB04
                  `--Oviraptoridae XT07
                       |  i. s.: Nemegtomaia barsboldi (Lü et al. 2004) [=Nemegtia barsboldi] N10
                       |--Gigantoraptor Xu, Tan et al. 2007 XT07
                       |    `--*G. erlianensis Xu, Tan et al. 2007 XT07
                       `--+--Oviraptorinae OCB04
                          |    |--Oviraptor Osborn 1924 OCB04
                          |    |    `--O. philoceratops Osborn 1924 OCB04
                          |    |--Rinchenia Barsbold 1997 OCB04
                          |    |    `--R. mongoliensis (Barsbold 1986) [=Oviraptor mongoliensis] OCB04
                          |    `--Citipati Clark, Norell & Barsbold 2001 OCB04
                          |         `--C. osmolskae Clark, Norell & Barsbold 2001 OCB04
                          `--Ingeniinae OCB04
                               |--+--Conchoraptor Barsbold 1986 OCB04
                               |  |    `--C. gracilis Barsbold 1986 OCB04
                               |  `--Khaan Clark, Norell & Barsbold 2001 OCB04
                               |       `--K. mckennai Clark, Norell & Barsbold 2001 OCB04
                               `--+--‘Ingenia’ Barsbold 1981 non Gerlach 1957 OCB04
                                  |    `--I. yanshini Barsbold 1981 OCB04 [=Oviraptor (Ingenia) yanshini P88]
                                  `--Heyuannia Lü 2003 ZZ08, OCB04
                                       `--H. huangi Lü 2003 OCB04

Chirostenotes Gilmore 1924 OCB04 [incl. Caenagnathus Sternberg 1940 OCB04, Macrophalangia Sternberg 1932 N85]

Chirostenotes elegans (Parks 1933) OCB04 [=Ornithomimus elegans OCB04; incl. Caenagnathus sternbergi Cracraft 1971 OCB04, Chirostenotes sternbergi D07]

Chirostenotes pergracilis Gilmore 1924 [incl. Caenagnathus collinsi Sternberg 1940, Macrophalangia canadensis Sternberg 1932] OCB04

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[MOW02] Maryańska, T., H. Osmólska & M. Wolsan. 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica 47 (1): 97–116.

[N02] Naish, D. 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria) Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings of the Geologists' Association 113: 153–163.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[OCB04] Osmólska, H., P. J. Currie & R. Barsbold. 2004. Oviraptorosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 165–183. University of California Press: Berkeley.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[XT07] Xu, X., Q. Tan, J. Wang, X. Zhao & L. Tan. 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature 447: 844–847.

[ZZ08] Zhang, F., Z. Zhou, X. Xu, X. Wang & C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105–1108.

Last updated: 11 July 2018.

Dromaeosauridae

Reconstruction of the roughly human-sized Achillobator giganticus, from here.

Belongs within: Maniraptora.

The Dromaeosauridae included some of the most hypercarnivorous of all maniraptoran coelurosaurs, including such famed taxa as Deinonychus antirrhopus and Velociraptor mongoliensis. They varied in size from members of the genus Utahraptor, estimated to be well over five metres in length, to species of Microraptor with lengths less than one metre (among the smallest known non-avian dinosaurs). The dromaeosaurs are most famous for the development of a large hyperextendable sickle claw on digit II of the foot, most commonly presumed to have played some part in the subduing and killing of prey animals. Well-preserved specimens of dromaeosaurids (particularly of Microraptor species) have demonstrated the presence of extensive feather covering; in Microraptor, elongate feathers were present on the legs as well as on the arms, and may have served an aerodynamic function.

The majority of dromaeosaurid species are known from the Cretaceous of Eurasia and North America. However, the family also includes a basal clade of Unenlagia and related taxa from South America.

Characters (from Norell & Makovicky 2004): Frontals short, T-shaped, with sinusoidal edge demarcating rostral boundary of supratemporal fenestra; squamosal with caudolateral overhanging shelf; quadrate bearing lateral process contacting quadratojugal dorsally above enlarged quadrate foramen; dorsal vertebrae bearing raised, stalklike parapophyses; digit II modified, raptorial; caudal vertebrae with chevrons and prezygapophyses elongate, spanning several vertebrae; coracoid bearing subglenoid fossa.

<==Dromaeosauridae [Dromaeosaurinae, Velociraptorinae]
    |--Mahakala GC13
    `--+--+--Austroraptor GC13
       |  |--Neuquenraptor argentinus GC13, XC09
       |  |--Buitreraptor Makovicky 2005 GC13, D07
       |  |    `--B. gonzalezorum HH09
       |  `--Unenlagia Novas & Puerta 1997 GC13, NM04
       |       |--U. comahuensis Novas & Puerta 1997 NM04
       |       `--U. paynemili CG13
       `--+--Shanag GC13
          `--+--+--Graciliraptor Xu & Wang 2004 GC13, D07
             |  |    `--G. lujiatunensis D07
             |  `--+--Sinornithosaurus Xu, Wang & Wu 1999 GC13, NM04
             |     |    |--S. haoiana D07
             |     |    `--S. millenii Xu, Wang & Wu 1999 NM04
             |     `--Microraptor Xu, Zhou & Wang 2000 GC13, NM04
             |          |--M. gui Xu, Zhou et al. 2003 NM04
             |          `--M. zhaoianus Xu, Zhou & Wang 2000 NM04
             `--+--Tsaagan mangas GC13, XT07
                `--+--+--Tianyuraptor GC13
                   |  `--Bambiraptor Burnham, Derstler et al. 2000 GC13, NM04
                   |       `--B. feinbergorum Burnham, Derstler et al. 2000 NM04
                   `--+--Adasaurus Barsbold 1983 HH09, NM04
                      |    `--A. mongoliensis Barsbold 1983 NM04
                      `--+--Velociraptor Osborn 1924 GC13, NM04
                         |    `--V. mongoliensis Osborn 1924 NM04
                         `--+--Saurornitholestes Sues 1978 GC13, NM04
                            |    `--S. langstoni Sues 1978 NM04 [=Velociraptor langstoni P88]
                            `--+--Deinonychus Ostrom 1969 HH09, NM04
                               |    `-*D. antirrhopus Ostrom 1969 O69 [=Velociraptor antirrhopus P88]
                               `--+--Atrociraptor marshalli HH09
                                  `--+--Achillobator Perle, Norell & Clark 1999 HH09, NM04
                                     |    `--A. giganticus Perle, Norell & Clark 1999 NM04
                                     `--+--Dromaeosaurus Matthew & Brown 1922 GC13, NM04
                                        |    |--D. albertensis Matthew & Brown 1922 NM04
                                        |    `--D. gracilis (Marsh 1888) (n. d.) [=Coelurus gracilis] O69
                                        `--Utahraptor Kirkland, Burge & Gaston 1993 GC13, NM04
                                             |--U. ostrommaysi Kirkland, Burge & Gaston 1993 NM04
                                             `--U. spielbergi D07

Dromaeosauridae incertae sedis:
  Pyroraptor Allain & Taquet 2000 NM04
    `--P. olympius Allain & Taquet 2000 NM04
  Variraptor Le Loeuff & Buffetaut 1998 NM04
    `--V. mechinorum Le Loeuff & Buffetaut 1998 NM04
  Hulsanpes Osmólska 1982 NM04
    `--H. perlei Osmólska 1982 NM04
  Phaedrolosaurus ilikensis Dong 1973 (n. d.) P88
  Euronychodon Antunes & Sigogneau-Russell 1991 NM04
    |--E. asiaticus Nessov 1995 NM04
    `--E. portucalensis Antunes & Sigogneau-Russell 1991 NM04
  Ornithodesmus Seeley 1887 NM04, W91
    `--*O. cluniculus Seeley 1887 W91
  Unquillosaurus Powell 1979 NM04
    `--U. ceibalii Powell 1979 NM04
  Palaeopteryx thomsoni Jensen 1981 (n. d.) NM04

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[HMC04] Holtz, T. R., Jr, R. E. Molnar & P. J. Currie. 2004. Basal Tetanurae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 71–110. University of California Press: Berkeley.

[HH09] Hu, D., L. Hou, L. Zhang & X. Xu. 2009. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature 461: 640–643.

[NM04] Norell, M. A., & P. J. Makovicky. 2004. Dromaeosauridae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 196–209. University of California Press: Berkeley.

[O69] Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History 30: 1–165.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[W91] Wellnhofer, P. 1991. The Illustrated Encyclopedia of Pterosaurs.  Salamander Books: London (reprinted 2000. In: Norman, D., & P. Wellnhofer. The Illustrated Encyclopedia of Dinosaurs. Salamander Books).

[XC09] Xu, X., J. M. Clark, J. Mo, J. Choiniere, C. A. Forster, G. M. Erickson, D. W. E. Hone, C. Sullivan, D. A. Eberth, S. Nesbitt, Q. Zhao, R. Hernandez, C.-K. Jia, F.-L. Han & Y. Guo. 2009. A Jurassic ceratosaur from China helps clarify avian digital homologies. Nature 459: 940–944.

[XT07] Xu, X., Q. Tan, J. Wang, X. Zhao & L. Tan. 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature 447: 844–847.

Last updated: 3 July 2018.

Avialae

Reconstruction of the Early Cretaceous Jeholornis prima by Matt Martyniuk.

Belongs within: Maniraptora.
Contains: Ornithothoraces.

Though the specific definition has varied from author to author, the clade Avialae has always effectively referred to the total group of all 'birds', i. e. the union of living birds with other flying fossil dinosaurs. Basal members of the clade such as Archaeopteryx and Jeholornis retained a long skeletal tail. Eleven specimens of Archaeopterygidae are known from the Late Jurassic of Europe; some authors have assigned all these specimens to a single species Archaeopteryx lithographica whereas others have recognised multiple species.

More derived avialans in the clade Pygostylia had the tail reduced as in modern birds. This clade includes the Confuciusornithidae from the lower Cretaceous Yixian Formation of China, which are characterised by toothless jaws with a hooked rostrum (Padian 2004).

Synapomorphies (from Holtz 1998): Vertebral foramen to cranial articular facet ratio of dorsals 0.4 or greater; less than 25 caudal vertebrae present; glenoid oriented laterally; proximal ulnar shaft arched; preacetabular portion of ilium significantly longer than postacetabular portion; ischial proximodorsal process just distal to iliac process; ischiadic terminal processes separate; proximal region of fibular medial face flat; metatarsal IV subequal in length to metatarsal II; metatarsal I completely reverted.

<==Avialae [Archaeornithes, Palaeoaves, Sauriurae]
    |--Aurornis Godefroit, Cau et al. 2013 GC13
    |    `--*A. xui Godefroit, Cau et al. 2013 GC13
    `--+--Anchiornis huxleyi GC13
       `--+--Archaeopterygidae [Archaeopterygiformes, Archaeopteryginae] P04
          |    |--Wellnhoferia Elżanowski 2001 P04
          |    |    `--W. grandis Elżanowski 2001 P04
          |    `--Archaeopteryx von Meyer 1861 GC13, P04 (see below for synonymy)
          |         |--A. bavarica Wellnhofer 1993 P04
          |         `--A. lithographica von Meyer 1861 (see below for synonymy) P04
          `--+--Xiaotingia zhengi GC13
             `--+--+--Rahonavis Forster, Sampson et al. 1998 GC13, P04 (see below for synonymy)
                |  |    `--R. ostromi (Forster, Sampson et al. 1998) [=Rahona ostromi Forster, Sampson et al. 1998] P04
                |  `--Shenzhouraptor Zhou & Zhang 2002 GC13, D07
                |       `--S. sinensis D07
                `--+--Balaur GC13
                   |--Jeholornis Zhou & Zhang 2002 ZZ02
                   |    `--*J. prima Zhou & Zhang 2002 ZZ02
                   `--+--Sapeornis Zhou & Zhang 2002 GC13, P04
                      |    `--S. chaoyangensis Zhou & Zhang 2002 P04
                      `--Pygostylia P04
                           |--Ornithothoraces GC13
                           `--Confuciusornithidae [Confuciusornithiformes] P04
                                |--Changchengornis Ji, Chiappe & Ji 1999 P04
                                |    `--C. hengdaoziensis Hi, Chiappe & Ji 1999 P04
                                `--Confuciusornis Hou, Zhou et al. 1995 GC13, P04
                                     |--C. dui Hou, Martin et al. 1999 P04
                                     `--C. sanctus Hou, Zhou et al. 1995 (see below for synonymy) P04

Avialae incertae sedis:
  Songlingornis linghensis ZBH03
  Jibeinia Hou 2000 P04
    `--J. luahera Hou 2000 P04
  Wileyia valdensis Harrison & Walker 1973 (n. d.) P04
  Ilerdopteryx viai Lacasa-Ruiz 1991 (n. d.) P04
  ‘Ornithomimus’ minutus Marsh 1892 P04
  Platannavis nana Nessov 1992 (n. d.) P04
  Protoavis texensis Chatterjee 1991 (n. d.) P04
  Cuspirostrisornis Hou 1997 D07
    `--C. houi D07
  Yandangornis Cai & Zhao 1999 D07
    `--Y. longicaudatus D07

Archaeopteryx von Meyer 1861 GC13, P04 [incl. Archaeornis Petronievics in Petronievics & Woodward 1917 P04, Griphosaurus Wagner 1861 P04, Jurapteryx Howgate 1985 P04]

Archaeopteryx lithographica von Meyer 1861 [incl. Pterodactylus crassipes von Meyer 1857, Archaeopteryx crassipes, Rhamphorhynchus crassipes, Scaphognathus crassipes, Griphornis longicaudatus Woodward 1862, Griphosaurus longicaudatus, Archaeopteryx macrura Owen 1862, Archaeopteryx oweni Petronievics 1921, Griphosaurus problematicus Woodward 1862, Archaeopteryx recurva Howgate 1984, Jurapteryx recurva, Archaeopteryx siemensii Dames 1897, Archaeornis siemensi] P04

Confuciusornis sanctus Hou, Zhou et al. 1995 [incl. C. chuonzhous Hou 1997, C. suniae Hou 1997] P04

Rahonavis Forster, Sampson et al. 1998 GC13, P04 [incl. Rahona Forster, Sampson et al. 1998 P04; Rahonavidae, Rahonaviformes]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5-61.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

[ZZ02] Zhou, Z., & F. Zhang. 2002. A long-tailed, seed-eating bird from the Early Cretaceous of China. Nature 418: 405–409.

Last updated: 6 August 2017.

Maniraptora

Mounted skeleton of Falcarius utahensis with first author James Kirkland, via the New York Times.

Belongs within: Coelurosauria.
Contains: Oviraptorosauria, Dromaeosauridae, Troodontidae, Avialae.

The clade Maniraptora contains all theropods more closely related to the birds than to ornithomimosaurs. The name refers to the large, grasping hands used in capturing prey by some members of the clade, such as the dromaeosaurs, though members of the clade were morphologically and ecologically diverse. These include the Alvarezsauroidea, a group of small, cursorial forms in which the forelimbs had the first digit enlarged relative to the others, the latter often being reduced and non-functional. The Therizinosauria, in contrast, were mostly large, heavy-bodied forms with small leaf-shaped teeth and edentulous beaks that were probably herbivorous (Kirkland et al. 2005). The Scansoriopterygidae were small arboreal theropods capable of gliding flight.

See also: A new stem-bird and publication in the digital age.

<==Maniraptora [Metornithes]
    |--Alvarezsauroidea CX10
    |    |--Haplocheirus Choiniere, Xu et al. 2010 GC13, CX10
    |    |    `--*H. sollers Choiniere, Xu et al. 2010 CX10
    |    `--Alvarezsauridae [Alvarezsauria] CX10
    |         |--Alvarezsaurus Bonaparte 1991 GC13, P04
    |         |    `--A. calvoi Bonaparte 1991 P04
    |         |--Achillesaurus manazzonei XS11
    |         `--+--+--Bonapartenykus GC13
    |            |  `--Patagonykus Novas 1996 GC13, P04
    |            |       `--P. puertai Novas 1996 P04
    |            `--Parvicursorinae XS11
    |                 |  i. s.: Ceratonykus osculans CX10
    |                 |--Linhenykus Xu, Sullivan et al. XS11
    |                 |    `--*L. monodactylus Xu, Sullivan et al. XS11
    |                 `--+--Albertonykus boreale XS11
    |                    `--+--Xixianykus zhangi XS11
    |                       `--+--Parvicursor Karhu & Rautian 1996 XS11, P04
    |                          |    `--P. remotus Karhu & Rautian 1996 P04
    |                          `--+--Mononykus Perle, Norell et al. 1993 XS11, P04
    |                             |    `--M. olecranus Perle, Norell et al. 1993 P04
    |                             `--Shuuvuia Chiappe, Norell & Clark 1998 XS11, P04
    |                                  `--S. deserti Chiappe, Norell & Clark 1998 P04
    `--+--+--Oviraptorosauria GC13
       |  `--Paraves [Eumaniraptora] GC13
       |       |--Scansoriopterygidae GC13
       |       |    |--Epidexipteryx Zhang, Zhou et al. 2008 GC13, ZZ08
       |       |    |    `--*E. hui Zhang, Zhou et al. 2008 ZZ08
       |       |    `--Epidendrosaurus Zhang, Zhao et al. 2002 GC13, P04 [incl. Scansoriopteryx Czerkas & Yuan 2000 P04]
       |       |         `--E. ningchengensis Zhang, Zhou et al. 2002 GC13 (see below for synonymy)
       |       `--+--Eosinopteryx brevipinna GC13
       |          `--Eumaniraptora [Deinonychosauria] GC13
       |               |  i. s.: Nuthetes Owen 1854 D07
       |               |           `--N. destructor Owen 1854 (n. d.) HMC04
       |               |         Cryptovolans Cxerkas, Zhang et al. 2002 D07
       |               |           `--C. pauli D07
       |               |         Yixianosaurus Xu & Wang 2003 D07
       |               |           `--Y. longimanus D07
       |               |--Dromaeosauridae GC13
       |               `--+--Troodontidae GC13
       |                  `--Avialae GC13
       `--Therizinosauria [Segnosauria, Segnosaurischia, Therizinosauroidea] ZZ08
            |  i. s.: ‘Chilantaisaurus’ zheziangensis Dong 1979 CMB04
            |         Enigmosaurus Barsbold & Perle 1983 CMB04
            |           `--E. mongoliensis Barsbold & Perle 1983 CMB04
            |         Jianchangosaurus NS15
            |--Falcarius Kirkland, Zanno et al. 2005 GC13, KZ05
            |    `--*F. utahensis Kirkland, Zanno et al. 2005 KZ05
            `--+--Beipiaosaurus Xu, Tang & Wang 1999 GC13, CMB04
               |    `--B. inexpectus Xu, Tang & Wang 1999 CMB04
               `--+--Alxasaurus Russell & Dong 1993 GC13, CMB04
                  |    `--A. elesitaiensis Russell & Dong 1993 CMB04
                  `--Therizinosauridae CMB04
                       |  i. s.: Eshanosaurus Xu, Zhao & Clark 2001 CMB04
                       |           `--E. deguchiianus Xu, Zhao & Clark 2001 CMB04
                       |--Erlianosaurus Xu, Zhang et al. 2002 ZZ08, CMB04
                       |    `--E. bellamanus Xu, Zhang et al. 2002 CMB04
                       |--Nothronychus Kirkland & Wolfe 2001 GC13, CMB04
                       |    |--N. graffami GC13
                       |    `--N. mckinleyi Kirkland & Wolfe 2001 CMB04
                       |--Nanshiungosaurus Dong 1979 ZZ08, CMB04
                       |    |--N. bohlini Dong & Yu 1997 CMB04
                       |    `--N. brevispinus Dong 1979 CMB04
                       `--+--Neimongosaurus Zhang, Xu et al. 2001 ZZ08, CMB04
                          |    `--N. yangi Zhang, Xu et al. 2001 CMB04
                          `--+--Segnosaurus Perle 1979 ZZ08, CMB04
                             |    `--S. galbinensis Perle 1979 CMB04
                             `--+--Therizinosaurus Maleev 1954 ZZ08, CMB04
                                |    `--T. cheloniformis Maleev 1954 CMB04
                                `--Erlikosaurus Perle 1981 GC13, CMB04
                                     `--E. andrewsi Perle 1981 CMB04

Epidendrosaurus ningchengensis Zhang, Zhou et al. 2002 GC13 [incl. Scansoriopteryx heilmanni Czerkas & Yuan 2000 P04

*Type species of generic name indicated

REFERENCES

[CX10] Choiniere, J. N., X. Xu, J. M. Clark, C. A. Forster, Y. Guo & F. Han. 2010. A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China. Science 327: 571–574.

[CMB04] Clark, J. M., T. Maryańska & R. Barsbold. 2004. Therizinosauroidea. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 151–164. University of California Press: Berkeley.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[HMC04] Holtz, T. R., Jr, R. E. Molnar & P. J. Currie. 2004. Basal Tetanurae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 71–110. University of California Press: Berkeley.

[KZ05] Kirkland, J. I., L. E. Zanno, S. D. Sampson, J. M. Clark & D. D. DeBlieux. 2005. A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature 435: 84–87.

[NS15] Novas, F. E., L. Salgado, M. Suárez, F. L. Agnolin, M. D. Ezcurra, N. R. Chimento, R. de la Cruz, M. P. Isasi, A. O. Vargas & D. Rubilar-Rogers. 2015. An enigmatic plant-eating theropod from the Late Jurassic period of Chile. Nature 522: 331–334.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[XS11] Xu, X., C. Sullivan, M. Pittman, J. N. Choiniere, D. Hone, P. Upchurch, Q. Tan, D. Xiao, L. Tan & F. Han. 2011. A monodactyl nonavian dinosaur and the complex evolution of the alvarezsauroid hand. Proceedings of the National Academy of Sciences of the USA 108 (6): 2338–2342.

[ZZ08] Zhang, F., Z. Zhou, X. Xu, X. Wang & C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105–1108.

Last updated: 11 July 2018.

Coelurosauria

Reconstruction of Coelurus fragilis by FunkMonk.

Belongs within: Tetanurae.
Contains: Tyrannosauroidea, Maniraptora.

The Coelurosauria is the clade of theropod dinosaurs including all species more closely related to modern birds than to Allosaurus. This is perhaps the most ecologically and morphologically diverse group of dinosaurs. Even without considering the birds themselves, it includes the tyrannosaurs, the giant graviportal therizinosaurs and the small derived alvarezsaurs. Within the coelurosaurs we have definitive evidence for the presence of protofeathers in basal forms such as Tyrannosauroidea and Compsognathidae. These protofeathers were filamentous, hair-like structures; true aerodynamic feathers, with a knitted planar surface formed from branching filaments, probably developed within the Maniraptora.

Notable subgroups of the Coelurosauria include the Ornithomimosauria, slender cursorial theropods from the Cretaceous of Eurasia and North America. Most ornithomimosaurs lacked teeth, and they may have included omnivorous or herbivorous forms. The Compsognathidae are small coelurosaurs known from the Late Jurassic and Early Cretaceous of Eurasia; until recently, Compsognathus longipes was regarded as the smallest known non-avian dinosaur (smaller forms are now known among the Maniraptora). Coelurus fragilis is a small theropod (about two metres in length) known from the late Jurassic of Wyoming.

Synapomorphies (from Holtz 1998): Cervical central surfaces amphiplatyan; pubic boot boat-shaped (pointed cranially and caudally) in ventral view, angle between shaft and caudal portion of boot acute; proximal end of fibula 75% or more proximal width of tibia.

<==Coelurosauria [Avofilopluma, Coeluridae, Ornitholestinae]
    |--Sciurumimus GC13
    |--Zuolong GC13
    |--Gasosaurus Dong & Tang 1985 H98, HMC04
    |    `--G. constructus Dong & Tang 1985 HMC04
    |--Tugulusaurus Dong 1973 GC13, HMC04
    |    `--T. faciles Dong 1973 HMC04
    `--+--+--Coelurus Marsh 1879 GC13, HMC04
       |  |    `--C. fragilis Marsh 1879 [incl. C. agilis Marsh 1884] HMC04
       |  `--+--Tanycolagreus topwilsoni GC13, ZZ08
       |     `--Tyrannosauroidea GC13
       `--+--Sinocalliopteryx GC13
          `--+--+--Huaxiagnathus Hwang, Norell et al. 2004 GC13, D07
             |  |    `--H. orientalis ZZ08
             |  `--Compsognathidae [Compsognathia] HMC04
             |       |--Compsognathus Wagner 1861 GC13, HMC04
             |       |    `--C. longipes Wagner 1861 [incl. C. corallestris Bidar, Demay & Thomel 1972] HMC04
             |       |--Sinosauropteryx Ji & Ji 1996 GC13, HMC04
             |       |    `--S. prima Ji & Ji 1996 HMC04
             |       `--Aristosuchus Seeley 1887 N02
             |            `--*A. pusillus (Owen 1876) [=Poekilopleuron pusillus, Poikilopleuron pusillus] N02
             `--+--+--Juravenator GC13
                |  `--Ornitholestes Osborn 1903 GC13, HMC04
                |       `--O. hermanni Osborn 1903 H98 [=Coelurus hermanni P88]
                `--Maniraptoriformes [Bullatosauria, Protoavia] MOW02
                     |--Maniraptora GC13
                     `--+--Nqwebasaurus de Klerk, Forster et al. 2000 GC13, HMC04
                        |    `--N. thwazi de Klerk, Forster et al. 2000 HMC04
                        `--Ornithomimosauria [Arctometatarsalia, Garudimiminae, Ornithomimidae, Ornithomiminae] MKC04
                             |  i. s.: Sinornithomimus Kobayashi & Lü 2003 MKC04
                             |           `--S. dongi Kobayashi & Lü 2003 MKC04
                             |         Coelosaurus (n. d.) MKC04
                             |           |--C. affinis Gilmore 1920 (n. d.) MKC04
                             |           `--C. antiquus Leidy 1865 (n. d.) MKC04
                             |         Timimus hermani Rich & Vickers-Rich 1994 MKC04
                             |--Harpymimus Barsbold & Perle 1984 ZZ08, MKC04
                             |    `--H. okladnikovi Barsbold & Perle 1984 MKC04
                             `--+--Deinocheirus Osmólska & Roniewicz 1970 ZZ08, MKC04 [Deinocheiridae, Deinocheirinae]
                                |    `--D. mirificus Osmólska & Roniewicz 1970 MKC04
                                `--+--Shenzhousaurus Ji, Norell et al. 2003 ZZ08, MKC04
                                   |    `--S. orientalis Ji, Norell et al. 2003 MKC04
                                   `--+--Pelecanimimus Pérez-Moreno, Sanz et al. 1994 ZZ08, MKC04
                                      |    |--P. hoyasi SR05
                                      |    `--P. polyodon Pérez-Moreno, Sanz et al. 1994 MKC04
                                      `--+--Garudimimus Barsbold 1981 GC13, MKC04
                                         |    `--G. brevipes Barsbold 1981 MKC04
                                         `--+--Archaeornithomimus Russell 1972 ZZ08, MKC04
                                            |    |--A. affinis (Gilmore 1920) (see below for synonymy) P88
                                            |    |--A. asiaticus (Gilmore 1933) [=Ornithomimus asiaticus] MKC04
                                            |    `--A. bissektensis Nessov 1995 (n. d.) MKC04
                                            `--+--Gallimimus Osmólska, Roniewicz & Barsbold 1972 GC13, MKC04
                                               |    |--G. bullatus Osmólska, Roniewicz & Barsbold 1972 MKC04 (see below for synonymy)
                                               |    `--G. mongoliensis D07
                                               `--+--Struthiomimus Osborn 1917 ZZ08, MKC04
                                                  |    |--S. altus (Lambe 1902) [=Ornithomimus altus] MKC04
                                                  |    `--S. sedens D07
                                                  `--+--Anserimimus Barsbold 1988 ZZ08, MKC04
                                                     |    `--A. planinychus Barsbold 1988 MKC04
                                                     `--Ornithomimus Marsh 1890 ZZ08, MKC04 (see below for synonymy)
                                                          |--O. antiquus D07
                                                          |--O. edmontonicus Sternberg 1933 MKC04 (see below for synonymy)
                                                          |--O. lonzeensis D07
                                                          |--O. sedens Marsh 1892 (n. d.) MKC04
                                                          |--O. tenuis Marsh 1890 (n. d.) MKC04
                                                          `--O. velox Marsh 1890 MKC04

Coelurosauria incertae sedis:
  Kakuru Molnar & Pledge 1980 HMC04
    `--K. kujani Molnar & Pledge 1980 HMC04
  Scipionyx Dal Sasso & Signore 1998 HMC04
    `--S. samniticus Dal Sasso & Signore 1998 HMC04
  Nedcolbertia Kirkland, Britt et al. 1998 HMC04
    `--N. justinhofmanni Kirkland, Britt et al. 1998 HMC04
  Archaeornithoides Elżanowski & Wellnhofer 1992 HMC04
    `--A. deinosauriscus Elżanowski & Wellnhofer 1992 HMC04
  Ricardoestesia Currie, Rigby & Sloan 1990 HMC04
    |--R. gilmorei Currie, Rigby & Sloan 1990 HMC04
    `--R. isosceles Sankey 2001 HMC04
  Rapator Huene 1932 P88, HMC04
    `--R. ornitholestoides Huene 1932 HMC04
  Pedopenna Xu & Zhang 2005 D07
    `--P. daohugouensis D07

Archaeornithomimus affinis (Gilmore 1920) [=Ornithomimus (Archaeornithomimus) affinis] P88

Gallimimus bullatus Osmólska, Roniewicz & Barsbold 1972 MKC04 [=Ornithomimus bullatus P88]

Ornithomimus Marsh 1890 ZZ08, MKC04 [incl. Dromiceiomimus Russell 1972 MKC04]

Ornithomimus edmontonicus Sternberg 1933 MKC04 [incl. Struthiomimus brevitertius Parks 1926 MKC04, Dromiceiomimus brevitertius N85, Ornithomimus brevitertius P88, Struthiomimus currelli Parks 1933 MKC04, S. ingens Parks 1933 MKC04, Ornithomimus ingens O69, S. samueli Parks 1928 MKC04, Dromiceiomimus samueli P88, Struthiomimus samueli P88]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[H98] Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5–61.

[HMC04] Holtz, T. R., Jr, R. E. Molnar & P. J. Currie. 2004. Basal Tetanurae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 71–110. University of California Press: Berkeley.

[MKC04] Makovicky, P. J., Y. Kobayashi & P. J. Currie. 2004. Ornithomimosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 137–150. University of California Press: Berkeley.

[MOW02] Maryańska, T., H. Osmólska & M. Wolsan. 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica 47 (1): 97–116.

[N02] Naish, D. 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria) Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings of the Geologists' Association 113: 153–163.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[O69] Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History 30: 1–165.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[SR05] Senter, P., & J. H. Robins. 2005. Range of motion in the forelimb of the theropod dinosaur Acrocanthosaurus atokensis, and implications for predatory behaviour. Journal of Zoology 266: 307–318.

[ZZ08] Zhang, F., Z. Zhou, X. Xu, X. Wang & C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105–1108.

Last updated: 11 July 2018.