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.

Iguanodontia

Reconstruction of Iguanodon atherfieldensis, copyright Steveoc 86.

Belongs within: Ornithopoda.
Contains: Hadrosauroidea.

The Iguanodontia are a diverse group of bipedal and facultatively quadrupedal herbivorous dinosaurs known from the Late Jurassic and Cretaceous, recognised by Norman (2004) as the clade of all taxa closer to Edmontosaurus than Thescelosaurus.

Characters (from Norman 2004): Premaxilla with transversely expanded and edentulous margin; predentary with smoothly convex rostral margin in dorsal aspect; dentary ramus deep, with parallel dorsal and ventral borders; sternal rib ossification absent; one phalanx lost on manus digit III; prepubic process compressed, blade-shaped.

<==Iguanodontia [Iguanodontidae]
    |  i. s.: Albisaurus scutifer Fritsch 1905 (n. d.) N04
    |         Loncosaurus argentinus Ameghino 1898 (n. d.) N04
    |         Tichosteus (n. d.) N04
    |           |--T. aequifacies Cope 1877 (n. d.) N04
    |           `--T. lucasanus Cope 1877 (n. d.) N04
    |--Tenontosaurus Ostrom 1970 N04
    |    |--T. dossi Winkler, Murry & Jacobs 1997 N04
    |    `--T. tilletti Ostrom 1970 N04
    `--+--Muttaburrasaurus Bartholomai & Molnar 1981 N04
       |    `--M. langdoni Bartholomai & Molnar 1981 N04
       `--Dryomorpha N04
            |  i. s.: Callovosaurus Galton 1980 (n. d.) N85, N04
            |           `--*C. leedsi (Lydekker 1889) [=Camptosaurus leedsi] N04
            |         Kangnasaurus coetzeei Haughton 1915 (n. d.) N85, N04
            |--Dryosauridae N04
            |    |--Planicoxa DiCroce & Carpenter 2001 N04
            |    |    `--P. venenica DiCroce & Carpenter 2001 N04
            |    |--Dryosaurus Marsh 1894 [incl. Dysalotosaurus Virchow 1919] N04
            |    |    |--D. altus (Marsh 1878) [=Laosaurus altus] N04
            |    |    `--D. lettowvorbecki (Virchow 1919) [=Dysalotosaurus lettowvorbecki] N04
            |    `--Valdosaurus Galton 1977 N04
            |         |--V. canaliculatus (Galton 1975) (see below for synonymy) N04
            |         `--V. nigeriensis Galton & Taquet 1982 N04
            `--Ankylopollexia N04
                 |  i. s.: Draconyx Mateus & Antunes 2001 N04
                 |           `--D. loureiroi Mateus & Antunes 2001 N04
                 |--Camptosaurus Marsh 1885 (see below for synonymy) N04
                 |    |--C. amplus (Marsh 1879) [=Camptonotus amplus] N04
                 |    |--C. depressus Gilmore 1909 N04
                 |    |--C. dispar (Marsh 1879) (see below for synonymy) N04
                 |    |--C. hoggi (Owen 1874) [=Iguanodon hoggi] N04
                 |    `--C. prestwichii (Hulke 1880) [=Iguanodon prestwichii] N04
                 `--Styracosterna N04
                      |--Lurdusaurus Taquet & Russell 1999 [incl. Gravisaurus Chabli 1988] N04
                      |    `--L. arenatus Taquet & Russell 1999 [incl. Gravisaurus tenerensis Chabli 1988] N04
                      |--Equijubus You, Luo et al. 2003 N04
                      |    `--E. normani You, Luo et al. 2003 N04
                      `--Iguanodontoidea [Hadrosauriformes] N04
                           |  i. s.: Craspedodon Dollo 1883 N04
                           |           `--C. lonzeensis Dollo 1883 N04
                           |         Fukuisaurus Kobayashi & Azuma 2003 N04
                           |           `--F. tetoriensius Kobayashi & Azuma 2003 N04
                           |         Shuangmiaosaurus You, Ji et al. 2003 N04
                           |           `--S. gilmorei You, Ji et al. 2003 N04
                           |--Hadrosauroidea N04
                           `--Iguanodon Mantell 1825 [incl. Heterosaurus Cornuel 1850, Iguanosaurus Ritgen 1828] N04
                                |--*I. anglicus Holl 1829 WU03, N04 (see below for synonymy)
                                |--I. atherfieldensis Hooley 1924 (see below for synonymy) N04
                                |--I. bernissartensis Boulenger 1881 [incl. I. seelyi Hulke 1882] N04
                                |--I. dawsoni Lydekker 1888 N04
                                |--I. exogirarus Fritsch 1878 (n. d.) N04
                                |--I. fittoni Lydekker 1889 [incl. I. hollingtoniensis Lydekker 1889] N04
                                |--I. lakotaensis Weishampel & Bjork 1989 N04
                                `--I. ottingeri Galton & Jensen 1979 (n. d.) N04

Camptosaurus Marsh 1885 [incl. Brachyrophus Cope 1878, Cumnoria Seeley 1999, Symphyrophus Cope 1878; Camptosauridae] N04

Camptosaurus dispar (Marsh 1879) [=Camptonotus dispar; incl. Brachyrophus altarkansanus Cope 1878, Camptosaurus browni Gilmore 1909, Camptosaurus medius Marsh 1894, Symphyrophus musculosus Cope 1878, Camptosaurus nanus Marsh 1894] N04

*Iguanodon anglicus Holl 1829 WU03, N04 [incl. Cetiosaurus brevis Owen 1842 N04, Streptospondylus major Owen 1842 N04]

Iguanodon atherfieldensis Hooley 1924 [incl. Sphenospondylus gracilis Lydekker 1888, I. mantelli Mayer 1832, Heterosaurus neocomiensis Cornuel 1850, Vectisaurus valdensis Hulke 1879] N04

Valdosaurus canaliculatus (Galton 1975) [=Dryosaurus canaliculatus; incl. Camptosaurus valdensis Lydekker 1889] N04

*Type species of generic name indicated

REFERENCES

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

[WU03] Wilson, J. A., & P. Upchurch. 2003. A revision of Titanosaurus Lydekker (Dinosauria—Sauropoda), the first dinosaur genus with a ‘Gondwanan’ distribution. Journal of Systematic Palaeontology 1 (3): 125–160.

Hadrosaurinae

Skeletal mount of Edmontosaurus annectens, copyright Andytang20.

Belongs within: Hadrosauridae.

The Hadrosaurinae are a lineage of hadrosaurid dinosaurs that were mostly crestless, or possessed a solid crest formed from an extension of the nasal bones. The clade as given below is supported by the presence of a caudal margin on the circumnarial fossa (Horner et al. 2004). Members of the genus Edmontosaurus include some of the largest and best known hadrosaurids, reaching lengths of around twelve metres or more.

<==Hadrosaurinae
    |--Lophorhothon Langston 1960 HWF04
    |    `--L. atopus Langston 1960 HWF04
    `--+--+--Naashoibitosaurus Hunt & Lucas 1993 HWF04
       |  |    `--N. ostromi Hunt & Lucas 1993 HWF04
       |  |--Kritosaurus Brown 1910 HWF04, D07
       |  |    |--K. australis Bonaparte, Franchi et al. 1984 HWF04
       |  |    `--K. navajovius Brown 1910 (n. d.) HWF04
       |  `--Saurolophus Brown 1912 HWF04
       |       |--S. angustirostris Rozhdestvensky 1952 HWF04
       |       |--S. kryschtofovici Riabinin 1930 (n. d.) HWF04
       |       `--S. osborni Brown 1912 HWF04
       `--+--Prosaurolophus Brown 1916 HWF04
          |    |--P. blackfeetensis Horner 1992 HWF04
          |    `--P. maximus Brown 1916 HWF04
          `--+--Gryposaurus Lambe 1914 HWF04
             |    |--G. incurvimanus (Parks 1920) [=Kritosaurus incurvimanus] HWF04
             |    |--G. latidens Horner 1992 HWF04
             |    `--G. notabilis Lambe 1914 HWF04 [=Kritosaurus notabilis D07]
             `--+--Edmontosaurus Lambe 1920 (see below for synonymy) HWF04
                |    |--E. annectens (Marsh 1892) HWF04 (see below for synonymy)
                |    |--E. regalis Lambe 1917 [incl. Trachodon atavus Cope 1871, Agathaumas milo Cope 1874] HWF04
                |    `--E. saskatchewanensis (Sternberg 1926) [=Thespesius saskatchewanensis] HWF04
                `--+--Brachylophosaurus Sternberg 1953 HWF04
                   |    `--B. canadensis Sternberg 1953 [incl. B. goodwini Horner 1988] HWF04
                   `--Maiasaura Horner & Makela 1979 HWF04
                        `--M. peeblesorum Horner & Makela 1979 HWF04

Hadrosaurinae incertae sedis:
  Anasazisaurus Hunt & Lucas 1993 HWF04
    `--A. horneri Hunt & Lucas 1993 HWF04
  Aralosaurus Rozhdestvensky 1968 HWF04
    `--A. tuberiferus Rozhdestvensky 1968 HWF04
  Shantungosaurus Hu 1973 HWF04
    `--S. giganteus Hu 1973 HWF04
  Kerberosaurus Bolotsky & Godefroit 2004 D07
    `--K. manakini D07
  Hadrosaurus Leidy 1858 HWF04
    |--H. breviceps Marsh 1889 (n. d.) HWF04
    |--H. cavatus Cope 1871 (n. d.) HWF04
    |--H. foulkii Leidy 1858 [incl. Ornithotarsus immanis Cope 1869] HWF04
    |--H. minor Marsh 1870 (n. d.) HWF04
    `--H. paucidens Marsh 1889 (n. d.) HWF04

Edmontosaurus Lambe 1920 [incl. Anatosaurus Lull & Wright 1942, Anatotitan Brett-Surman in Chapman & Brett-Surman 1990] HWF04

Edmontosaurus annectens (Marsh 1892) HWF04 [=Claosaurus annectens HWF04; incl. Anatosaurus copei Lull & Wright 1942 HWF04, Anatotitan copei D07, Thespesius edmontonensis Gilmore 1924 HWF04, Trachodon longiceps Marsh 1890 HWF04, Anatotitan longiceps D07]

*Type species of generic name indicated

REFERENCES

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

[HWF04] Horner, J. R., D. B. Weishampel & C. A. Forster. 2004. Hadrosauridae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 438–463. University of California Press: Berkeley.

Last updated: 3 July 2018.

Hadrosauridae

Skeletal mount of Hypacrosaurus altispinus, copyright Roland Tanglao.

Belongs within: Hadrosauroidea.
Contains: Hadrosaurinae.

The Hadrosauridae are a group of large herbivorous dinosaurs known from the Late Cretaceous of Eurasia and North and South America, recognised by Horner et al. (2004) as the least inclusive clade containing Telmatosaurus and Parasaurolophus. Most hadrosaurids can be divided between the sublineages Hadrosaurinae and Lambeosaurinae; members of the latter group often possessed large crests formed from posterior expansions of the nasal bones.

Characters (from Horner et al. 2004): Three or more replacement teeth present per tooth family; dentary tooth row extending distally to terminate caudal to apex of coronoid process, caudalmost termination of dentary well behind coronoid process; surangular foramen absent; ectopterygoid and jugal not in contact; supraorbital absent or fused to orbit rim; neural arches with cervical zygapophyseal peduncles elevated to extend well above level of neural canal; postzygapophyses long, dorsally arched; coracoid bearing long cranioventral process extending well below glenoid; proximal scapula dorsoventrally narrow, with acromion process projecting horizontally, notched cranioventral corner, reduced area for coracoid articulation; femur bearing deep intercondylar extensor groove, with edges meeting or nearly meeting cranially to enclose extensor tunnel.

<==Hadrosauridae
    |--Telmatosaurus Nopcsa 1903 [incl. Hecatasaurus Brown 1910, Limnosaurus Nopcsa 1900] HWF04
    |    |--T. cantabrigiensis D07
    |    `--T. transsylvanicus Nopcsa 1900 HWF04 [=Limnosaurus transsylvanicus HWF04, Orthomerus transsylvanicus PST99]
    `--Euhadrosauria HWF04
         |--Hadrosaurinae HWF04
         `--Lambeosaurinae HWF04
              |  i. s.: Pararhabdodon isonesis WJ03
              |         Barsboldia Maryańska & Osmólska 1981 HWF04
              |           `--B. sicinskii Maryańska & Osmólska 1981 HWF04
              |         Nipponosaurus Nagao 1936 HWF04
              |           `--N. sachalinensis Nagao 1936 HWF04
              |         Bactrosaurus Gilmore 1933 D07
              |           |--B. johnsoni Gilmore 1933 HWF04
              |           |--B. kyzylkumensis D07
              |           `--B. prynadai Riabinin 1939 (n. d.) HWF04
              |--Tsintaosaurus Young 1958 HWF04
              |    `--T. spinorhinus Young 1958 [incl. Tanius laiyangensis Zhen 1976] HWF04
              `--+--Jaxartosaurus Riabinin 1939 GBA03, HWF04
                 |    |--J. aralensis Riabinin 1939 [incl. Procheneosaurus convincens Rozhdestvensky 1968] HWF04
                 |    `--J. fuyunensis Wu 1984 (n. d.) HWF04
                 `--+--Amurosaurus Bolotsky & Kurzanov 1991 GBA03, HWF04
                    |    `--A. riabinini Bolotsky & Kurzanov 1991 HWF04
                    `--+--+--Charonosaurus Godefroit, Zan & Jin 2000 GBA03, HWF04
                       |  |    `--C. jiayinensis Godefroit, Zan & Jin 2000 HWF04
                       |  `--Parasaurolophus Parks 1922 GBA03, HWF04
                       |       |--P. cyrtocristatus Ostrom 1961 HWF04
                       |       |--P. tubicen Wiman 1931 HWF04
                       |       `--P. walkeri Parks 1922 HWF04
                       `--+--Lambeosaurus Parks 1923 GBA03, HWF04 [incl. Didanodon Osborn 1902 HWF04]
                          |    |--L. lambei Parks 1923 HWF04 (see below for synonymy)
                          |    |--L. laticaudus Morris 1981 HWF04
                          |    `--L. magnicristatus Sternberg 1935 HWF04
                          `--+--Corythosaurus Brown 1914 GBA03, HWF04 [incl. Procheneosaurus Matthew 1920 HWF04]
                             |    `--C. casuarius Brown 1914 HWF04 (see below for synonymy)
                             |--Olorotitan Godefroit, Boltsky & Alifanov 2003 GBA03
                             |    `--*O. arharensis Godefroit, Bolotsky & Alifanov 2003 GBA03
                             `--Hypacrosaurus Brown 1913 GBA03, HWF04
                                  |--H. altispinus Brown 1913 [incl. Cheneosaurus tolmanensis Lambe 1917] HWF04
                                  |--H. casuarius P88
                                  `--H. stebingeri Horner & Currie 1994 HWF04

Hadrosauridae incertae sedis:
  Gilmoreosaurus Brett-Surman 1979 HWF04
    |--G. arkhangelskyi D07
    |--G. atavus D07
    `--G. mongoliensis (Gilmore 1933) [=Mandschurosaurus mongoliensis] HWF04
  Tanius Wiman 1929 HWF04
    `--T. sinensis Wiman 1929 [incl. T. chingkankouensis Young 1958] HWF04
  Claosaurus Marsh 1890 HWF04
    |--C. affinis Wieland 1903 (n. d.) HWF04
    `--C. agilis (Marsh 1872) [=Hadrosaurus agilis] HWF04
  Secernosaurus Brett-Surman 1979 HWF04
    `--S. koerneri Brett-Surman 1979 HWF04
  Arstanosaurus akkurganensis Suslov & Shilin 1982 (n. d.) HWF04
  Cionodon HWF04
    |--C. arctatus Cope 1874 (n. d.) HWF04
    |--C. kysylkumensis Riabinin 1931 (n. d.) HWF04
    `--C. stenopsis Cope 1875 (n. d.) HWF04
  Claorhynchus trihedrus Cope 1892 (n. d.) HWF04
  Diclonius HWF04
    |--D. calamarius Cope 1876 (n. d.) HWF04
    |--D. pentagonus Cope 1876 (n. d.) HWF04
    `--D. perangulatus Cope 1876 (n. d.) HWF04
  Hypsibema crassicauda Cope 1869 (n. d.) HWF04
  ‘Iguanodon’ hilli Newton 1892 (n. d.) HWF04
  Mandschurosaurus Riabinin 1930 (n. d.) HWF04
    |--M. amurensis (Riabinin 1925) [=Trachodon amurensis] HWF04
    `--M. laosensis Hoffet 1943 (n. d.) HWF04
  Microhadrosaurus nanshiungensis Dong 1979 (n. d.) HWF04
  Orthomerus HWF04
    |--O. dolloi Seeley 1883 (n. d.) HWF04
    `--O. weberi Riabinin 1945 (n. d.) HWF04
  Parrosaurus Gilmore 1945 (n. d.) HWF04
    `--*P. missouriensis (Gilmore & Stewart 1945) [=Neosaurus missouriensis] HWF04
  Pteropelyx grallipes Cope 1889 (n. d.) HWF04
  Thespesius occidentalis Leidy 1856 (n. d.) HWF04
  Trachodon HWF04
    |--T. altidens Lambe 1902 (n. d.) HWF04
    |--T. cantabrigiensis Lydekker 1888 (n. d.) HWF04
    |--T. marginatus Lambe 1902 (n. d.) HWF04
    |--T. mirabilis Leidy 1856 (n. d.) HWF04
    `--T. selwyni Lambe 1902 (n. d.) HWF04

Corythosaurus casuarius Brown 1914 HWF04 [incl. C. bicristatus Parks 1935 HWF04, C. brevicristatus Parks 1935 HWF04, Tetragonosaurus cranibrevis Sternberg 1935 HWF04, Procheneosaurus cranibrevis D75, T. erectofrons Parks 1931 HWF04, P. erectofrons D75, Corythosaurus excavatus Gilmore 1923 HWF04, C. intermedius Parks 1923 HWF04]

Lambeosaurus lambei Parks 1923 HWF04 [incl. L. clavinitialis Sternberg 1935 HWF04, Corythosaurus frontalis Parks 1935 HWF04, Tetragonosaurus praeceps Parks 1931 HWF04, Procheneosaurus praeceps D75]

*Type species of generic name indicated

REFERENCES

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

[D75] Dodson, P. 1975. Taxonomic implications of relative growth in lambeosaurine hadrosaurs. Systematic Zoology 24 (1): 37–54.

[GBA03] Godefroit, P., Y. Bolotsky & V. Alifanov. 2003. A remarkable hollow-crested hadrosaur from Russia: an Asian origin for lambeosaurines. Comptes Rendus Palevol 2: 143–151.

[HWF04] Horner, J. R., D. B. Weishampel & C. A. Forster. 2004. Hadrosauridae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 438–463. University of California Press: Berkeley.

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

[PST99] Pereda Suberbiola, X., & P. Taquet. 1999. Restes de Rhabdodon (dinosaure ornithopode) de Transylvanie donnés par Nopcsa au Muséum national d'Histoire naturelle de Paris. Geodiversitas 21 (2): 157–166.

[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.

Last updated: 3 July 2018.

Ceratopsia

Skeletal mounts of Psittacosaurus mongoliensis, copyright Ninjatacoshell.

Belongs within: Genasauria.
Contains: Ceratopsidae.

The Ceratopsia are a group of herbivorous dinosaurs known from the Late Jurassic to Cretaceous of Asia and North America (with some less definite records from other parts of the world). Members of this group are characterised by a novel bone, the rostral bone, at the front of the upper jaw (which forms a narrow beak), and a tendency to the develop of a frill on the rear margin of the skull (You & Dodson 2004).

See also: Big horned lizards; Ceratopsids: a Cretaceous flash in the pan.

Synapomorphies (from You & Dodson 2004): External naris high, separated from ventral border of premaxilla by flat area; rostral bone present; premaxilla enlarged; jugal with well-developed lateral flaring; wide dorsoventral length of infraorbital ramus of jugal; palatal extensions of maxillae contacting rostral to choana.

<==Ceratopsia
    |  i. s.: Micropachycephalosaurus Dong 1978 ZY09, MCW04
    |           `--M. hongtuyanensis Dong 1978 MCW04
    |--Psittacosaurus Osborn 1923 [incl. Protiguanodon Osborn 1923] YD04
    |    |--P. guyangensis Cheng 1983 YD04
    |    |--P. mazongshanensis Xu 1997 YD04
    |    |--P. meileyingensis Sereno, Chao et al. 1988 YD04
    |    |--P. mongoliensis Osborn 1923 (see below for synonymy) YD04
    |    |--P. neimongoliensis Russell & Zhao 1996 YD04
    |    |--P. ordosensis Russell & Zhao 1996 YD04
    |    |--P. osborni Young 1931 [incl. P. tingi Young 1931] YD04
    |    |--P. sattayaraki Buffetaut & Suteethorn 1992 (n. d.) YD04
    |    |--P. sinensis Young 1958 YD04
    |    |--P. xinjiangensis Sereno & Chao 1988 YD04
    |    `--P. youngi Chao 1962 YD04
    `--Neoceratopsia [Protoceratopidae, Protoceratopoidea] YD04
         |  i. s.: Microceratops YD04
         |           |--M. gobiensis Bohlin 1953 (n. d.) YD04
         |           `--M. sulcidens Bohlin 1953 (n. d.) YD04
         |         Kulceratops kulensis Nessov 1995 (n. d.) YD04
         |         Yamaceratops dorngobiensis BN08
         |         Bainoceratops efremovi T08
         |--Yinlong downsi BSN07
         `--+--Chaoyangsaurus Zhao, Cheng & Xu 1999 YD04
            |    `--C. youngi Zhao, Cheng & Xu 1999 YD04
            `--+--Liaoceratops Xu, Makovicky et al. 2002 XM02
               |    `--*L. yanzigouensis Xu, Makovicky et al. 2002 XM02
               `--+--Archaeoceratops Dong & Azuma 1997 [Archaeoceratopidae] YD04
                  |    `--A. oshimai Dong & Azuma 1997 YD04
                  `--Coronosauria YD04
                       |--Ceratopsoidea YD04
                       |    |--+--Ceratopsidae YD04
                       |    |  `--Zuniceratops Wolfe & Kirkland 1998 YD04
                       |    |       `--Z. christopheri Wolfe & Kirkland 1998 YD04
                       |    `--+--Asiaceratops salsopaludalis Nessov, Kaznyshkina & Cherepanov 1989 (n. d.) XM02, YD04
                       |       `--Leptoceratopsidae YD04
                       |            |--Montanoceratops Sternberg 1951 YD04
                       |            |    `--M. cerorhynchus (Brown & Schlaikjer 1942) [=Leptoceratops cerorhynchus] YD04
                       |            `--+--Leptoceratops Brown 1914 YD04
                       |               |    `--L. gracilis Brown 1914 YD04
                       |               `--Udanoceratops Kurzanov 1992 XM02, YD04
                       |                    `--U. tschizhovi Kurzanov 1992 YD04
                       `--Protoceratopsidae YD04
                            |--Graciliceratops Sereno 2000 YD04
                            |    `--G. mongoliensis Sereno 2000 YD04
                            |--Protoceratops Granger & Gregory 1923 YD04
                            |    |--P. andrewsi Granger & Gregory 1923 YD04
                            |    `--P. hellenikorhinus Lambert, Godefroit et al. 2001 YD04
                            `--Bagaceratopidae A03
                                 |--Bagaceratops Maryańska & Osmólska 1975 [incl. Breviceratops Kurzanov 1990] YD04
                                 |    `--B. rozhdestvenskyi Maryańska & Osmólska 1975 (see below for synonymy) YD04
                                 |--Lamaceratops Alifanov 2003 A03
                                 |    `--*L. tereschenkoi Alifanov 2003 A03
                                 `--Platyceratops Alifanov 2003 A03
                                      `--*P. tatarinovi Alifanov 2003 A03

Bagaceratops rozhdestvenskyi Maryańska & Osmólska 1975 [incl. Protoceratops kozlowskii Maryańska & Osmólska 1975, Breviceratops kozlowskii] YD04

Psittacosaurus mongoliensis Osborn 1923 [incl. Protiguanodon mongoliensis Osborn 1923, Ps. protiguanodonensis Young 1958] YD04

*Type species of generic name indicated

REFERENCES

[A03] Alifanov, V. R. 2003. Two new dinosaurs of the infraorder Neoceratopsia (Ornithischia) from the Upper Cretaceous of the Nemegt Depression, Mongolian People’s Republic. Paleontologicheskii Zhurnal 2003 (5): 77–88 (transl. Paleontological Journal 37 (5): 524–534).

[BN08] Balanoff, A. M., M. A. Norell, G. Grellet-Tinner & M. R. Lewin. 2008. Digital preparation of a probable neoceratopsian preserved within an egg, with comments on microstructural anatomy of ornithischian eggshells. Naturwissenschaften 95: 493–500.

[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.

[MCW04] Maryańska, T., R. E. Chapman & D. B. Weishampel. 2004. Pachycephalosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 464–477. University of California Press: Berkeley.

[T08] Tereschenko, V. S. 2008. Adaptive features of protoceratopoids (Ornithischia: Neoceratopsia). Paleontologicheskii Zhurnal 2008 (3): 50–64.

[XM02] Xu, X., P. J. Makovicky, X.-L. Wang, M. A. Norell & H.-L. You. 2002. A ceratopsian dinosaur from China and the early evolution of Ceratopsia. Nature 416: 314–317.

[YD04] You H. & P. Dodson. 2004. Basal Ceratopsia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 478–493. 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.