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.

Genasauria

Skeletal reconstruction of Agilisaurus louderbacki, copyright Jaime A. Headden.

Belongs within: Dinosauria.
Contains: Stegosauria, Ankylosauria, Ornithopoda, Pachycephalosauria, Ceratopsia.

The Genasauria are a major clade of herbivorous dinosaurs that have been defined as uniting the Thyreophora (armoured dinosaurs), Ornithopoda and Marginocephalia (pachycephalosaurs and ceratopsians). All Ornithischia surviving after the early Jurassic are members of this clade. Basal genasaurs include Lesothosaurus diagnosticus, a lightly built bipedal species about two metres in length from the early Jurassic of southern Africa.

Synapomorphies (from Butler et al. 2007): Metacarpals with proximal ends not block-like; manual phalanges/metacarpals with at least some extensor pits on dorsal surface of distal end absent or poorly developed; manual unguals not strongly recurved, without prominent flexor tubercle; brevis shelf and fossa with fossa facing ventrolaterally, visible in lateral view along entire length, creating deep postacetabular portion.

<==Genasauria BSN07
    |--+--Lesothosaurus Galton 1978 ZY09, NWW04a
    |  |    `--L. diagnosticus Galton 1978 NWW04a
    |  `--Thyreophora NWW04b
    |       |  i. s.: Tatisaurus Simmons 1965 NWW04b
    |       |           `--T. oehleri Simmons 1965 [=Scelidosaurus oehleri] NWW04b
    |       |         Bienosaurus Dong 2001 NWW04b
    |       |           `--B. lufengensis Dong 2001 NWW04b
    |       |         Lusitanosaurus de Lappaarent & Zbyszewski 1957 NWA04b, D07
    |       |           `--L. liasicus Lapparent & Zbyszewski 1957 (n. d.) NWW04b
    |       |--Laquintasaura venezuelae BNB17
    |       |--Scutellosaurus Colbert 1981 NWW04b
    |       |    `--S. lawleri Colbert 1981 NWW04b
    |       `--+--Emausaurus Haubold 1991 NWW04b
    |          |    `--E. ernsti Haubold 1991 NWW04b
    |          `--+--Scelidosaurus Owen 1860 NWW04b
    |             |    `--S. harrisonii Owen 1861 NWW04b
    |             `--Eurypoda [Thyreophoroidea] NWW04b
    |                  |--Stegosauria NWW04b
    |                  `--Ankylosauria NWW04b
    `--Neornithischia [Euornithopoda] BSN07
         |  i. s.: Atlascopcosaurus Rich & Rich 1989 NS04
         |           `--A. loadsi Rich & Rich 1989 NS04
         |         Gongbusaurus NS04
         |           |--G. shiyii Dong, Zhou & Zhang 1983 (n. d.) NWW04a
         |           `--G. wucaiwanensis Dong 1989 NS04
         |         Notohypsilophodon Martínez 1998 NS04
         |           `--N. comodorensis Martínez 1998 NS04
         |         Qantassaurus Rich & Vickers-Rich 1999 NS04
         |           `--Q. intrepidus Rich & Vickers-Rich 1999 NS04
         |         Yandusaurus He 1979 NS04
         |           `--Y. hongheensis He 1979 NS04
         |--Stormbergia dangershoeki BSN07
         `--+--Agilisaurus Peng 1990 BSN07, NS04
            |    |--A. louderbacki Peng 1990 NS04
            |    `--A. multidens (He & Cai 1983) [=Yandusaurus multidens] NS04
            `--+--Hexinlusaurus multidens BSN07
               `--+--Othnielosaurus celer BSN07
                  `--Cerapoda BSN07
                       |--Ornithopoda BSN07
                       `--Marginocephalia BSN07
                            |--Pachycephalosauria BSN07
                            `--Ceratopsia BSN07

*Type species of generic name indicated

REFERENCES

[BNB17] Baron, M. G., D. B. Norman & P. M. Barrett. 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543: 501–506.

[BSN07] Butler, R. J., R. M. H. Smith & D. B. Norman. 2007. A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia. Proceedings of the Royal Society of London Series B—Biological Sciences 274 (1621): 2041–2046.

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

[NS04] Norman, D. B., H.-D. Sues, L. M. Witmer & R. A. Coria. 2004. Basal Ornithopoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 393–412. University of California Press: Berkeley.

[NWW04a] Norman, D. B., L. M. Witmer & D. B. Weishampel. 2004a. Basal Ornithischia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 325–334. University of California Press: Berkeley.

[NWW04b] Norman, D. B., L. M. Witmer & D. B. Weishampel. 2004b. Basal Thyreophora. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 335–342. University of California Press: Berkeley.

[ZY09] Zheng, X.-T., H.-L. You, X. Xu & Z.-M. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333–336.

Last updated: 6 August 2017.

Ankylosauria

Reconstruction of Edmontonia rugosidens by E. M. Fulda.

Belongs within: Genasauria.
Contains: Ankylosaurinae.

The Ankylosauria is a clade of more or less heavily armoured, quadrupedal, herbivorous dinosaurs that has been defined as including all taxa closer to Ankylosaurus than Stegosaurus. Most classifications of ankylosaurs have recognised the families Nodosauridae, with relatively slender snouts, and Ankylosauridae, with broader snouts and often with a large bony club on the end of the tail.

Synapomorphies (from Vickaryous et al. 2004): Skull with cranial ornamentation present across rostral region and lateral surface of angular; antorbital, supratemporal and external mandibular fenestrae absent; maxillary tooth row inset from lateral edge of maxilla; multiple parasagittal rows of osteoderms present across dorsal surface of neck and postcervical region of body; acetabulum imperforate; synsacrum composed of co-ossified dorsal, sacral and caudal vertebrae.

<==Ankylosauria
    |--Polacanthidae D07
    |    |--Mymoorapelta Kirkland & Carpenter 1994 D07
    |    |    `--M. maysi Kirkland & Carpenter 1994 VMW04
    |    `--Polacanthus Huxley 1867 VMW04
    |         |--P. foxii Hulke 1881 [=Hylaeosaurus foxii; incl. P. becklesi Hennig 1924] VMW04
    |         `--P. rudgwickensis Blows 1996 VMW04
    |--Ankylosauridae VMW04
    |    |  i. s.: Gastonia Kirklan 1998 VMW04
    |    |           `--G. burgei Kirkland 1998 VMW04
    |    |         Tianchisaurus Dong 1993 D07
    |    |           `--T. nedegoaperferima Dong 1993 (n. d.) VMW04
    |    |         Aletopelta Ford & Kirkland 2001 D07
    |    |           `--A. coombsi Ford & Kirkland 2001 (n. d.) VMW04
    |    |         Crichtonsaurus Dong 2002 D07
    |    |           `--C. bohlini Dong 2002 VMW04
    |    |         Shanxia Barrett, You et al. 1998 D07
    |    |           `--S. tianzhensis Barrett, You et al. 1998 (n. d.) VMW04
    |    |         Maleevus Tumanova 1987 (n. d.) D07, VMW04
    |    |           `--*M. disparoserratus (Maleev 1952) [=Syrmosaurus disparoserratus] VMW04
    |    |--Gargoyleosaurus Carpenter, Miles & Cloward 1998 VMW04
    |    |    `--G. parkpinorum Carpenter, Miles & Cloward 1998 VMW04
    |    `--+--Minmi Molnar 1980 VMW04
    |       |    `--M. paravertebra Molnar 1980 VMW04
    |       `--+--Ankylosaurinae VMW04
    |          `--+--Gobisaurus Vickaryous, Russell et al. 2001 VMW04
    |             |    `--G. domoculus Vickaryous, Russell et al. 2001 VMW04
    |             `--Shamosaurus Tumanova 1983 VMW04
    |                  `--S. scutatus Tumanova 1983 VMW04
    `--Nodosauridae VMW04
         |  i. s.: Struthiosaurus Bunzel 1871 VMW04 (see below for synonymy)
         |           |--S. austriacus Bunzel 1871 (see below for synonymy) VMW04
         |           |--S. languedocensis Garcia & Pereda-Suberbiola 2003 VMW04
         |           |--S. lugdunensis D07
         |           `--S. transylvanicus Nopcsa 1915 VMW04
         |         Rhodanosaurus lugdunensis Nopcsa 1929 (n. d.) N85, VMW04
         |         Dracopelta Galton 1980 N85
         |           `--D. zbyszewskii Galton 1980 VMW04
         |         Sarcolestes Lydekker 1893 N85, VMW04
         |           `--S. leedsi Lydekker 1893 VMW04
         |         Priodontognathus Seeley 1875 (n. d.) VMW04
         |           `--*P. phillipsii (Seeley 1869) [=Iguanodon phillipsii] VMW04
         |         Cryptodraco Lydekker 1889 (n. d.) N85, VMW04
         |           `--*C. eumerus (Seeley 1869) [=Cryptosaurus eumerus] VMW04
         |         Hoplitosaurus Lucas 1902 N85, VMW04
         |           `--H. marshi (Lucas 1901) [=Stegosaurus marshi, Polacanthus marshi] VMW04
         |         Polacanthoides ponderosus Nopcsa 1929 (n. d.) N85, VMW04
         |         Priconodon crassus Marsh 1888 (n. d.) N85, VMW04
         |         Animantarx Carpenter, Kirkland et al. 1999 VMW04
         |           `--A. ramaljonesi Carpenter, Kirkland et al. 1999 VMW04
         |         Liaoningosaurus Xu, Wang & You 2001 XWY01
         |           `--*L. paradoxus Xu, Wang & You 2001 XWY01
         |         Hylaeosaurus Mantell 1833 XWY01, VMW04 [incl. Vectensia Delair 1982 VMW04]
         |           `--H. armatus Mantell 1833 (see below for synonymy) VMW04
         |         Nodosaurus Marsh 1889 VMW04
         |           `--N. textilis Marsh 1889 VMW04
         |         Acanthopholis Huxley 1867 D07
         |           |--A. eucercus Seeley 1869 (n. d.) VMW04
         |           |--A. horridus Huxley 1867 (n. d.) VMW04
         |           |--A. hughesii D07
         |           |--A. keepingi D07
         |           |--A. macrocercus Seeley 1869 (n. d.) VMW04
         |           |--A. platypus Seeley 1869 (n. d.) VMW04
         |           `--A. stereocercus Seeley 1869 (n. d.) VMW04
         |         Niobrarasaurus Carpenter, Dilkes & Weishampel 1995 D07, VMW04
         |           `--N. coleii (Mehl 1936) [=Hierosaurus coleii, Nodosaurus coleii] VMW04
         |         Anoplosaurus Seeley 1879 D07, VMW04
         |           |--A. curtonotus Seeley 1879 VMW04
         |           |--A. major Seeley 1879 (n. d.) VMW04
         |           `--A. tanyspondulus D07
         |--Cedarpelta Carpenter, Kirkland et al. 2001 VMW04
         |    `--C. bilbeyhallorum Carpenter, Kirkland et al. 2001 VMW04
         `--+--Pawpawsaurus Lee 1996 VMW04
            |    `--P. campbelli Lee 1996 VMW04
            |--Sauropelta Ostrom 1970 N85
            |    `--S. edwardsorum Ostrom 1970 VMW04
            |--Silvisaurus Eaton 1960 VMW04
            |    `--S. condrayi Eaton 1960 VMW04
            `--+--Panoplosaurus Lambe 1919 VMW04
               |    `--P. mirus Lambe 1919 VMW04
               `--Edmontonia Sternberg 1928 [incl. Chassternbergia Bakker 1988, Denversaurus Bakker 1988] VMW04
                    |--E. australis Ford 2000 (n. d.) VMW04
                    |--E. longiceps Sternberg 1928 VMW04 (see below for synonymy)
                    `--E. rugosidens (Gilmore 1930) (see below for synonymy) VMW04

Ankylosauria incertae sedis:
  Sauroplites scutiger Bohlin 1953 (n. d.) VMW04
  Amtosaurus Kurzanov & Tumanova 1978 VMW04
    |--A. archibaldi Averianov 2002 VMW04
    `--A. magnus Kurzanov & Tumanova 1978 VMW04
  Stegosaurides [=Stegosauroides] N85
    `--S. excavatus Bohlin 1953 (n. d.) VMW04
  Stegopeltinae D07
    |--Stegopelta Williston 1905 VMW04
    |    `--S. landerensis Williston 1905 VMW04
    `--Glyptodontopelta Ford 2000 D07
         `--G. mimus Ford 2000 (n. d.) VMW04
  Texasetes Coombs 1995 VMW04
    `--T. pleurohalio Coombs 1995 VMW04
  Brachypodosaurus gravis Chakravarti 1934 (n. d.) VMW04
  Eucercosaurus tanyspondylus Seeley 1879 (n. d.) VMW04
  Hierosaurus sternbergi Wieland 1909 (n. d.) VMW04
  Lametasaurus indicus Matley 1923 (n. d.) VMW04
  Palaeoscincus Leidy 1856 D07
    |--P. asper Lambe 1902 (n. d.) VMW04
    |--P. costatus Leidy 1856 (n. d.) VMW04
    `--P. latus Marsh 1892 (n. d.) VMW04
  Peishansaurus philemys Bohlin 1953 (n. d.) VMW04
  Rhadinosaurus alcinus Seeley 1881 (n. d.) VMW04
  Sygonosaurus macrocercus Seeley 1879 (n. d.) VMW04

Edmontonia longiceps Sternberg 1928 VMW04 [incl. Denversaurus schlessmani Bakker 1988 VMW04, E. schlessmani D07]

Edmontonia rugosidens (Gilmore 1930) [=Palaeoscincus rugosidens, Chassternbergia rugosidens, Panoplosaurus rugosidens] VMW04

Hylaeosaurus armatus Mantell 1833 [incl. Hylosaurus mantelli Fitzinger 1843, Hyla. oweni Mantell 1844] VMW04

Struthiosaurus Bunzel 1871 VMW04 [incl. Cretaeomus Seeley 1881 VMW04, Danubiosaurus Bunzel 1871 VMW04, Danubriosaurus N85, Leipsanosaurus Nopcsa 1918 VMW04, Lepanosaurus N85, Pleuropeltis Seeley 1881 VMW04, Pleuropeltus N85]

Struthiosaurus austriacus Bunzel 1871 [incl. Danubiosaurus anceps Bunzel 1871, Hoplosaurus ischyrus Seeley 1881, Crataeomus lepidophorus Seeley 1881, Leipsanosaurus noricus Nopcsa 1918, C. pawlowitschii Seeley 1881, Pleuropeltis suessi Seeley 1881] VMW04

*Type species of generic name indicated

REFERENCES

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

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

[VMW04] Vickaryous, M. K., T. Maryańska & D. B. Weishampel. 2004. Ankylosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 363–392. University of California Press: Berkeley.

[XWY01] Xu, X., X.-L. Wang & H.-L. You. 2001. A juvenile ankylosaur from China. Naturwissenschaften 88: 297–300.

Last updated: 6 August 2017.

Stegosauria

Composite skeletal mount of Stegosaurus armatus, copyright Perry Quan.

Belongs within: Genasauria.

The Stegosauria are a group of quadrupedal herbivorous dinosaurs known from the Middle Jurassic to the Lower Cretaceous, members of which bore a characteristic armature of dorsal vertical plates (Galton & Upchurch 2004). The largest and best known stegosaur is Stegosaurus from the Late Jurassic of North America, some examples of which were over seven metres in length.

Characters (from Galton & Upchurch 2004): Parietals with flattened dorsal surface; quadrate with transversely compressed proximal head; neural canals of cranial dorsal vertebrae with diameter greater than half that of centra; dorsal neural arches at least 1.5 times as high as dorsal centra; mid-dorsal transverse processes oriented at 50° to horizontal or more steeply; proximal caudals with dorsal portions of neural spines transversely widened and axially compressed; length:width ratio of distal caudal central 1.2 or lower; proximal scapular plate with larger surface area than coracoid; humerus with prominent triceps tubercle and descending ridge located caudolateral to deltopectoral crest; dorsoventral thickness of carpal elements exceeding 10% of humeral length; ulnare and intermedium fused in adults; proximal carpals block-shaped in cranial view; distal carpals absent or failing to ossify; ilium with preacetabular process directed outward at 35° to sagittal plane; pubis with acetabular portion cup-shaped, facing laterally; prepubis:pubic shaft length ratio higher than 0.4; pubic shaft length:acetabular diameter ratio at least 2.0; pedal digit I absent; pedal digit III with no more than three phalanges; pedial digit IV with no more than three phalanges; parasagittal rows of plates or spines present; ossified epaxial tendons absent.

<==Stegosauria
    |  i. s.: Regnosaurus northamptoni (n. d.) GU04
    |--Huayangosaurus Dong, Tang & Zhou 1982 [Huayangosauridae] GU04
    |    `--H. taibaii Dong, Tang & Zhou 1982 GU04
    `--Stegosauridae GU04
         |  i. s.: Craterosaurus Seeley 1874 GU04
         |           `--C. pottonensis Seeley 1874 GU04
         |         Dravidosaurus Yadagiri & Ayyasami 1979 GU04
         |           `--D. blanfordi Yadagiri & Ayyasami 1979 GU04
         |         Monkonosaurus Zhou in Dong 1990 GU04
         |           `--M. lawulacus Zhou in Dong 1990 GU04
         |         Paranthodon Nopcsa 1929 GU04
         |           `--P. africanus (Broom 1912) (see below for synonymy) GU04
         |         Yingshanosaurus Zhou 1984 D07
         |           `--Y. jichuanensis D07
         |--Dacentrurus Lucas 1902 [incl. Omosaurus Owen 1875] GU04
         |    |--D. armatus (Owen 1875) GU04 (see below for synonymy)
         |    |--D. phillipsi (Seeley 1893) (n. d.) D07, GU04 [=Omosaurus’ phillipsi GU04]
         |    `--‘Omosaurus’ vetustus Huene 1910 (n. d.) GU04
         `--Stegosaurinae GU04
              |--Chungkingosaurus Dong, Zhou & Zhang 1983 GU04
              |    `--C. jiangbeiensis Dong, Zhou & Zhang 1983 GU04
              `--+--Wuerhosaurus Dong 1973 GU04
                 |    |--W. homheni Dong 1973 GU04
                 |    `--W. ordosensis Dong 1993 GU04
                 `--+--Hesperosaurus Carpenter, Miles & Cloward 2001 GU04
                    |    `--H. mjosi Carpenter, Miles & Cloward 2001 GU04
                    |--Lexovisaurus Hoffstetter 1957 GU04
                    |    |--L. durobrivensis (Hulke 1887) (see below for synonymy) GU04
                    |    `--L. vetustus D07
                    `--+--Stegosaurus Marsh 1877 [incl. Diracodon Marsh 1881, Hypsirhophus Cope 1879] GU04
                       |    |--S. affinis Marsh 1881 (n. d.) GU04
                       |    |--S. armatus Marsh 1877 (see below for synonymy) GU04
                       |    |--S. longispinus Gilmore 1914 GU04
                       |    |--S. madagascariensis Piveteau 1926 (n. d.) NWW04
                       |    `--S. stenops Marsh 1887 [incl. Diracodon laticeps Marsh 1881] GU04
                       `--+--Kentrosaurus Hennig 1915 (see below for synonymy) GU04
                          |    |--K. aethiopicus Hennig 1915 GU04
                          |    `--K. longispinus D07
                          `--+--Tuojiangosaurus Dong, Li et al. 1977 GU04
                             |    `--T. multispinus Dong, Li et al. 1977 GU04
                             `--Chialingosaurus Young 1959 GU04
                                  |--C. guangyuanensis D07
                                  `--C. kuani Young 1959 GU04

Dacentrurus armatus (Owen 1875) GU04 [=Omosaurus armatus GU04; incl. O. hastiger Owen 1877 GU04, O. lennieri Nopcsa 1911 GU04, Dacentrurus lennieri APS03, Astrodon pusillus Lapparent & Zbyszewski 1957 GU04]

Kentrosaurus Hennig 1915 [incl. Doryphorosaurus Nopcsa 1916, Kentrurosaurus Hennig 1916] GU04

Lexovisaurus durobrivensis (Hulke 1887) [=Omosaurus durobrivensis; incl. O. leedsi Seeley 1901, Stegosaurus priscus Nopcsa 1911] GU04

Paranthodon africanus (Broom 1912) [=Palaeoscincus africanus; incl. Par. owenii Nopcsa 1929, Anthodon serrarius Owen 1876] GU04

Stegosaurus armatus Marsh 1877 [incl. S. duplex Marsh 1887, Hypsirhophus seeleyanus Cope 1879, S. sulcatus Marsh 1887, S. ungulatus Marsh 1879] GU04

*Type species of generic name indicated

REFERENCES

[APS03] Allain, R., & X. Pereda Suberbiola. 2003. Dinosaurs of France. Comptes Rendus Palevol 2 (1): 27–44.

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

[GU04] Galton, P. M., & P. Upchurch. 2004. Stegosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 343–362. University of California Press: Berkeley.

[NWW04] Norman, D. B., L. M. Witmer & D. B. Weishampel. 2004. Basal Ornithischia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 325–334. University of California Press: Berkeley.

Last updated: 6 August 2017.

Ornithopoda

Reconstruction of Hypsilophodon, copyright Arthur Weasley.

Belongs within: Genasauria.
Contains: Iguanodontia.

The Ornithopoda is a clade of bipedal and facultatively quadrupedal herbivorous dinosaurs that has been defined as including all taxa closer to Edmontosaurus than to Triceratops (Norman et al. 2004). The composition of this clade is currently subject to revision; recent phylogenetic analyses have found that several taxa previously regarded as basal Ornithopoda should be placed outside the Ornithopoda + Marginocephalia clade. One of the best known basal ornithopods is Hypsilophodon foxii from the Early Cretaceous of England, a lightly built bipedal form less than two metres in length.

<==Ornithopoda [Hypsilophodontidae]
    |--Orodromeus Horner & Weishampel 1988 NS04
    |    `--O. makelai Horner & Weishampel 1988 NS04
    `--+--+--Zephyrosaurus Sues 1980 NS04
       |  |    `--Z. schaffi Sues 1980 NS04
       |  `--Hypsilophodon Huxley 1869 NS04
       |       |--H. foxii Huxley 1869 NS04 [=Iguanodon foxii N85]
       |       `--H. wielandi Galton & Jensen 1979 (n. d.) NS04
       `--+--Gasparinisaura Coria & Salgado 1996 NS04
          |    `--G. cincosaltensis Coria & Salgado 1996 NS04
          `--+--Iguanodontia WJ03
             `--+--Parksosaurus Sternberg 1937 NS04
                |    `--P. warreni (Parks 1926) [=Thescelosaurus warreni] NS04
                |--Thescelosaurus Gilmore 1913 NS04
                |    `--T. neglectus Gilmore 1913 [incl. T. edmontonensis Sternberg 1940] NS04
                `--Bugenasaura Galton 1995 NS04
                     `--B. infernalis Galton 1995 NS04 (see below for synonymy)

Ornithopoda incertae sedis:
  Othnielia Galton 1977 NS04
    `--O. rex (Marsh 1877) (see below for synonymy) NS04
  Drinker Bakker, Galton et al. 1990 NS04
    `--D. nisti Bakker, Galton et al. 1990 NS04
  Leaellynasaura Rich & Rich 1989 NS04
    `--L. amicagraphica Rich & Rich 1989 NS04
  Fulgurotherium Huene 1932 NS04
    `--F. australe Huene 1932 NS04
  Oryctodromeus Varricchio, Martin & Katsura 2007 D07
    `--O. cubicularis D07
  Anabisetia Coria & Calvo 1998 ZY09, NS04
    `--A. saldiviai Coria & Calvo 2002 NS04
  Talenkauen ZY09
  Jeholosaurus Xu, Wang & You 2000 NS04
    `--J. shangyuanensis Xu, Wang & You 2000 NS04

Bugenasaura infernalis Galton 1995 NS04 [incl. Thescelosaurus garbanii Morris 1976 NS04, Stygimoloch garbanii S03]

Othnielia rex (Marsh 1877) [=Nanosaurus rex; incl. Laosaurus consors Marsh 1894, L. gracilis Marsh 1878] NS04

*Type species of generic name indicated

REFERENCES

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

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

[NS04] Norman, D. B., H.-D. Sues, L. M. Witmer & R. A. Coria. 2004. Basal Ornithopoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 393–412. University of California Press: Berkeley.

[S03] Sullivan, R. M. 2003. Revision of the dinosaur Stegoceras Lambe (Ornithischia, Pachycephalosauridae). Journal of Vertebrate Paleontology 23 (1): 181–207.

[WJ03] Weishampel, D. B., C.-M. Jianu, Z. Csiki & D. B. Norman. 2003. Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. Journal of Systematic Palaeontology 1: 65–123.

[ZY09] Zheng, X.-T., H.-L. You, X. Xu & Z.-M. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333–336.

Last updated: 6 August 2017.

Ankylosaurinae

Skeletal reconstruction of Pinacosaurus, copyright Kabacchi.

Belongs within: Ankylosauria.

The Ankylosaurinae is a clade of ankylosaurs from the Late Cretaceous of Asia and North America that has been defined as including all taxa closer to Ankylosaurus than Shamosaurus (Vickaryous et al. 2004). Well known members of the group include the North American Euoplocephalus tutus, one of the the largest ankylosaurs reaching a length of around six metres.

Synapomorphies (from Vickaryous et al. 2004): Nuchal shelf obscuring occiput in dorsal view; quadrate condyle obscured laterally by quadratojugal boss; buccal emargination strongly concave.

<==Ankylosaurinae VMW04
    |  i. s.: Nodocephalosaurus Sullivan 1999 VMW04
    |           `--N. kirtlandensis Sullivan 1999 VMW04
    |--Tsagantegia Tumanova 1993 VMW04
    |    `--T. longicranialis Tumanova 1993 VMW04
    `--+--Tarchia Maryańska 1977 VMW04
       |    `--T. gigantea (Maleev 1956) [=Dyoplosaurus giganteus; incl. T. kielanae Maryańska 1977] VMW04
       `--+--+--Euoplocephalus Lambe 1910 (see below for synonymy) VMW04
          |  |    `--E. tutus (Lambe 1902) VMW04 (see below for synonymy)
          |  `--Ankylosaurus Brown 1908 VMW04
          |       `--A. magniventris Brown 1908 VMW04
          `--+--Saichania Maryańska 1977 VMW04
             |    `--S. chulsanensis Maryańska 1977 VMW04
             |--Talarurus Maleev 1952 VMW04
             |    `--T. plicatospineus Maleev 1952 VMW04
             `--Pinacosaurus Gilmore 1933 VMW04 [incl. Ninghsiasaurus N85, Virminicaudus N85]
                  |--P. grangeri Gilmore 1933 (see below for synonymy) VMW04
                  `--+--P. mephistocephalus Godefroit, Pereda-Suberbiola et al. 1999 VMW04
                     `--Tianzhenosaurus Pang & Cheng 1998 VMW04
                          `--T. youngi Pang & Cheng 1998 VMW04

Euoplocephalus Lambe 1910 [=Stereocephalus Lambe 1902 (preoc.); incl. Anodontosaurus Sternberg 1929, Dyoplosaurus Parks 1924, Scolosaurus Nopcsa 1928] VMW04

Euoplocephalus tutus (Lambe 1902) VMW04 [=Stereocephalus tutus VMW04; incl. Dyoplosaurus acutosquameus Parks 1924 VMW04, Euoplocephalus acutosquameus D07, Scolosaurus cutleri Nopcsa 1929 VMW04, E. cutleri N85, Anodontosaurus lambei Sternberg 1929 VMW04]

Pinacosaurus grangeri Gilmore 1933 [incl. P. ninghsiensis Young 1935, Syrmosaurus viminicaudus Maleev 1952] VMW04

*Type species of generic name indicated

REFERENCES

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

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

[VMW04] Vickaryous, M. K., T. Maryańska & D. B. Weishampel. 2004. Ankylosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 363–392. University of California Press: Berkeley.

Last updated: 6 August 2017.

Hadrosauroidea

Skeletal mount of Probactrosaurus gobiensis, copyright Greg Goebel.

Belongs within: Iguanodontia.
Contains: Hadrosauridae.

The name Hadrosauroidea has been used for a clade uniting the Hadrosauridae and Ouranosaurus; it is here used for the smallest clade uniting these two taxa as found in the phylogenetic analysis by Norman (2004). Supporting characters for this clade were not explicitly listed. Ouranosaurus nigeriensis was a relatively large ornithopod from the early Cretaceous of Niger, most noted from the relatively long neural spines arising from its vertebrae.

<==Hadrosauroidea
    |  i. s.: Jinzhousaurus Wang & Xu 2001 N04
    |           `--J. yangi Wang & Xu 2001 N04
    |--Ouranosaurus Taquet 1976 N04
    |    `--O. nigeriensis Taquet 1976 N04
    `--+--+--Eolambia Kirkland 1998 N04
       |  |    `--E. caroljonesa Kirkland 1998 N04
       |  `--Altirhinus Norman 1998 N04
       |       `--A. kurzanovi Norman 1998 N04 [incl. Iguanodon orientalis D07]
       `--+--Protohadros Head 1998 N04
          |    `--P. byrdi Head 1998 N04
          `--+--+--Hadrosauridae N04
             |  `--Nanyangosaurus Xu, Zhao et al. 2000 XZ00
             |       `--*N. zhugeii Xu, Zhao et al. 2000 XZ00
             `--Probactrosaurus Rozhdestvensky 1966 N04
                  |--P. alashanicus Rozhdestvensky 1966 N04
                  |--P. gobiensis Rozhdestvensky 1966 N04
                  `--P. mazongshanensis Lü 1997 N04

*Type species of generic name indicated

REFERENCES

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

[N04] Norman, D. B. 2004. Basal Iguanodontia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 413–437. University of California Press: Berkeley.

[XZ00] Xu X., Zhao X.-J., Lü J.-C., Huang W.-B., Li Z.-Y. & Dong Z.-M. 2000. A new iguanodontian from Sangping Formation of Neixiang, Henan and its stratigraphical implication. Vertebrata PalAsiatica 38 (3): 176–191.

Last updated: 5 August 2017.