Novialoidea

Cast of Rhamphorhynchus muensteri in the Musée d'Histoire Naturelle, Brussels, copyright M0tty.

Belongs within: Pterosauria.
Contains: Pterodactyloidea.

The Novialoidea are a clade of pterosaurs defined by Kellner (2003) as the most exclusive clade containing Campylognathoides and Quetzalcoatlus. Members of this clade have the third phalanx of the wing finger shorter than the first and second phalanges (Kellner 2003).

<==Novialoidea (see below for synonymy) K03
    |--Campylognathoides Strand 1928 RK13, U03
    |    |--C. indicus Jain 1974 W91
    |    |--C. liasicus (Quenstedt 1858) K03 [=Pterodactylus liasicus E13]
    |    `--C. zitteli [=Campylognathus zitteli] W13
    `--+--Cacibupteryx Gasparini et al. 2004 RK13, A08
       |    `--C. caribensis W13
       |--Dorygnathus Wagner 1860 RK13, U03 [incl. Parapsicephalus Arthaber 1919 W13]
       |    |--D. banthensis (Theodori 1830) K03 (see below for synonymy)
       |    `--D. purdoni Newton 1888 [=Scaphognathus purdoni, Parapsicephalus purdoni] W13
       |--+--Angustinaripterus He, Yan & Su 1983 RK13, U03
       |  |    `--A. longicephalus W13
       |  `--Harpactognathus Carpenter, Unwin et al. 2003 RK13, CU03
       |       `--*H. gentryii Carpenter, Unwin et al. 2003 CU03
       `--Breviquartossa U03
            |--Pterodactyloidea RK13
            `--Rhamphorhynchus von Meyer 1847 RK13, A08
                 |--R. jessoni Lydekker 1890 W91
                 |--R. longicaudus (Münster 1839) WK08, K03 [=Pterodactylus longicaudus W91]
                 |--R. longiceps U03 [incl. R. kokeni W91]
                 |--R. muensteri (Goldfuss 1831) W13 (see below for synonymy)
                 |--R. phyllurus Marsh 1882 W91
                 `--R. tendagurensis Reck 1931 W91

Novialoidea incertae sedis:
  Carniadactylus Dalla Vecchia 2009 W13
    `--*C. rosenfeldi (Dalla Vecchia 1995) W13, DV03 [=Eudimorphodon rosenfeldi W13]
  Fenghuangopterus lii W13
  Jianchangopterus zhaoianus W13
  Pterorhynchus Czerkas & Ji 2002 RK13, A08
    `--P. wellnhoferi W13
  Sericipterus wucaiwanensis W13
  Qinglongopterus guoi W13
  Bellubrunnus rothgaengeri W13
  Nesodactylus Colbert 1969 U03
    `--N. hesperius W13
  Rhamphocephalus Seeley 1880 [incl. Dolichorhamphus Seeley 1885, Pteromonodactylus Teryaev 1967] A08
    |--R. bucklandi (von Meyer 1832) [=Pterodactylus bucklandi] W91
    |--R. depressirostris W13
    `--R. prestwichi W91

Dorygnathus banthensis (Theodori 1830) K03 [=Ornithocephalus banthensis W91, Pterodactylus banthensis W91; incl. Dorygnathus mistelgauensis W13, Parapsicephalus mistelgauensis D07]

Novialoidea [Angustinaripterinae, Campylognathoididae, Monofenestrata, Parapsicephalinae, Pteromonodactylidae, Rhamphorhynchae, Rhamphorhynchinae] K03

Rhamphorhynchus muensteri (Goldfuss 1831) W13 [=Ornithocephalus muensteri W13, Pterodactylus muensteri W13; incl. R. gemmingi U03, R. intermedius U03]

Inorganic: Rhamphorhynchus gemmingi minilorientalus Okamura 1987 O87

*Type species of generic name indicated

REFERENCES

[A08] Averianov, A. O. 2008. Superorder Pterosauria. In: Ivakhnenko, M. F., & E. N. Kurochkin (eds) Fossil Vertebrates of Russia and Adjacent Territories. Fossil Reptiles and Birds. Part 1 pp. 319–342. GEOS: Moscow.

[CU03] Carpenter, K., D. Unwin, K. Cloward, C. Miles & C. Miles. 2003. A new scaphognathine pterosaur from the Upper Jurassic Morrison Formation of Wyoming, USA. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 45–54. The Geological Society: London.

[DV03] Dalla Vecchia, F. M. 2003. New morphological observations on Triassic pterosaurs. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 23–44. The Geological Society: London.

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

[K03] Kellner, A. W. A. 2003. Pterosaur phylogeny and comments on the evolutionary history of the group. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 105–137. The Geological Society: London.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[RK13] Rodrigues, T., & A. W. A. Kellner. 2013. Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England. ZooKeys 308: 1–112.

[U03] Unwin, D. M. 2003. On the phylogeny and evolutionary history of pterosaurs. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 139–190. The Geological Society: London.

[WK08] Wang, X., A. W. A. Kellner, Z. Zhou & D. de A. Campos. 2008. Discovery of a rare arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloidea) from China. Proceedings of the National Academy of Sciences of the USA 105 (6): 1983–1987.

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

[W13] Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press: Princeton (New Jersey).

Sauropodomorpha

Skeletal reconstruction of Massospondylus carinatus, copyright Ballista.

Belongs within: Dinosauria.
Contains: Plateosauridae, Anchisauria, Sauropoda.

The Sauropodomorpha are a clade of often large herbivorous dinosaurs, the best-known members of which are the gigantic sauropods. Non-sauropod sauropodomorphs are commonly referred to as 'prosauropods' though the majority of analyses have not recognised them as a monophyletic clade. The earliest sauropodomorphs, such as Saturnalia tupiniquim and Panphagia protos, are known from the Late Triassic (Carnian) of South America. These were small forms, close to 1.5 metres in length. Guaibasaurus candelariensis is a slightly larger form represented by a partial postcranial skeleton from the Norian of Brazil that has been placed by various analyses as either a basal sauropodomorph or basal theropod.

Synapomorphies (from Baron et al. 2017): Premaxilla with posterodorsal process thin, bar-like, restricted to ventral border of external naris; external naris extending posteriorly to overlie maxilla; anterior margin of external naris posterior to midlength of premaxillary body; posterior margin of external naris posterior to first maxillary alveolus; anterior profile of maxilla with a strong inflection (notch) at base of ascending ramus, creating an anterior ramus with parallel dorsal and ventral margins; squamosal with ventral process narrower than one-quarter of its length; ventral ramus of squamosal forming more than half of caudal border of lower temporal fenestra; 18 or more dentary teeth present; premaxillary teeth with crown at least moderately expanded above root; maxillary/dentary tooth serrations present as enlarged and coarser (lower density) denticles; majority of maxillary and dentary teeth not recurved; anterior dentary teeth inclined anteriorly (procumbent); ten or more cervical vertebrae present; cervical vertebrae lacking prezygaodiapophyseal lamina; preacetabular process triangular and pointed; ischial shaft distally expanded into distinct ‘foot’ or 'boot'; tibia with posterolateral margin of distal end concave.

<==Sauropodomorpha [Plateosauria, Prosauropoda]
    |--+--Saturnalia Langer, Abdala et al. 1999 BNB17, GU04
    |  |    `--S. tupiniquim Langer, Abdala et al. 1999 GU04
    |  `--Chromogisaurus NS15
    |--Pampadromaeus barberenai BNB17
    `--+--Guaibasaurus Bonaparte, Ferigolo & Ribeiro 1999 BNB17, L04
       |    `--G. candelariensis Bonaparte, Ferigolo & Ribeiro 1999 L04
       |--Panphagia protos BNB17
       |--Pantydraco caducus BNB17
       |--Thecodontosaurus Riley & Stutchbury 1836 BNB17, GU04 [Thecodontosauridae]
       |    |--*T. antiquus Morris 1843 Y03
       |    |--T. caducus Yates 2003 Y03
       |    |--T. dubius Haughton 1924 (n. d.) GU04
       |    `--T. minor Haughton 1918 (n. d.) GU04
       `--+--Plateosauravus Huene 1932 NS15, GU04 [Plateosauravidae]
          |    `--*P. cullngworthi (Haughton 1924) (n. d.) [=Plateosaurus cullingworthi] GU04
          `--+--Plateosauridae BNB17
             `--+--+--Lufengosaurus Young 1941 BNB17, GU04
                |  |    |--L. changhduensis D07
                |  |    `--L. huenei Young 1941 [incl. L. magnus Young 1947] GU04
                |  `--Gyposaurus GU04
                |       |--G. capensis Broom 1911 (n. d.) GU04
                |       `--G. sinensis Young 1941 GU04 [=Anchisaurus capensis N85]
                `--+--Coloradisaurus Lambert 1983 BNB17, GU04 [incl. Coloradia Bonaparte 1978 GU04]
                   |    `--C. brevis (Bonaparte 1978) [=Coloradia brevis] GU04
                   `--+--Yunnanosaurus Young 1942 BNB17, GU04 [Yunnanosauridae]
                      |    `--Y. huangi Young 1942 [incl. Y. robustus Young 1951] GU04
                      `--+--+--Massospondylus Owen 1854 BNB17, GU04 [Massospondylidae]
                         |  |    |--M. browni Seeley 1895 (n. d.) GU04
                         |  |    |--M. carinatus Owen 1854 GU04
                         |  |    |--M. harriesi Broom 1911 (n. d.) GU04
                         |  |    |--M. hislopi Lydekker 1890 (n. d.) GU04
                         |  |    |--M. kaalea BNB17
                         |  |    `--M. schwarzi Haughton 1924 (n. d.) GU04
                         |  `--+--Leyesaurus marayensis NS15, BNB17
                         |     `--Adeopapposaurus NS15
                         `--+--Anchisauria BNB17
                            `--+--Aardonyx celestae BNB17
                               |--Pulanesaura eocollum BNB17
                               `--Sauropoda BNB17

Sauropodomorpha incertae sedis:
  Avalonianus sanfordi (Seeley 1898) (n. d.) N85, H79 [=Avalonia sanfordi H79]
  Clepsysaurus (n. d.) N85
  Dinosaurus (n. d.) N85
  Nyasasaurus parringtoni BNB17
  Orosauravus (n. d.) N85
  Palaeosauriscus (n. d.) N85
  Picrodon herveyi Seeley 1898 (n. d.) N85, H79
  Thotobolosaurus (n. d.) N85
  Gresslyosaurus GU04
    |--G. ingens Rütimeyer 1856 (n. d.) GU04
    |--G. plieningeri APS03
    |--G. robustus Huene 1907-1908 (n. d.) GU04
    `--G. torgeri Jaekel 1911 (n. d.) GU04
  Aetonyx palustris Broom 1911 (n. d.) GU04
  Dromicosaurus gracilis Hoepen 1920 (n. d.) GU04
  Hortalotarsus skirtopodus Seeley 1894 (n. d.) GU04
  Leptospondylus capensis Owen 1854 (n. d.) GU04
  Pachysauriscus Kuhn 1959 (n. d.) [incl. Pachysaurops Huene 1959] GU04
    `--*P. ajax (Huene 1907-1908) (n. d.) [=Pachysaurus ajax, *Pachysaurops ajax] GU04
  Pachyspondylus orpenii Owen 1854 (n. d.) GU04
  Yimenosaurus Bai, Yang & Wang 1990 GU04
    `--Y. youngi Bai, Yang & Wang 1990 GU04
  Fulengia Carroll & Galton 1977 GU04, E88
    `--F. youngi Carroll & Galton 1977 (n. d.) GU04
  Pachysaurus GU04
    |--P. giganteus Huene 1932 (n. d.) GU04
    |--P. magnus Huene 1907-1908 (n. d.) GU04
    |--P. reiningeri Huene 1907-1908 (n. d.) GU04
    `--P. wetzelianus Huene 1932 (n. d.) GU04
  Palaeosaurus fraserianus Cope 1878 (n. d.) GU04
  Strenusaurus procerus Bonaparte 1969 (n. d.) GU04
  Tawasaurus minor Young 1982 (n. d.) GU04
  ‘Teratosaurus’ minor Huene 1907-1908 (n. d.) GU04
  ‘Teratosaurus’ trossingensis Huene 1907-1908 (n. d.) GU04
  Ruehleia Galton 2001 NS15, GU04
    `--R. bedheimensis Galton 2001 GU04
  Jingshanosaurus Zhang & Yang 1995 GU04
    `--J. xinwaensis Zhang & Yang 1995 GU04
  Agrosaurus mcgillivrayi Seeley 1891 GU04
  Basutodon ferox Huene 1932 (n. d.) N85, H79

*Type species of generic name indicated

REFERENCES

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

[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.

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

[E88] Evans, S. E. 1988. The early history and relationships of the Diapsida. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 221–260. Clarendon Press: Oxford.

[GU04] Galton, P. M., & P. Upchurch. 2004. Prosauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 232–258. University of California Press: Berkeley.

[H79] Heerden, J. van. 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; late Triassic) from South Africa. Navorsinge van die Nasionale Museum 4: 21–84.

[L04] Langer, M. C. 2004. Basal Saurischia. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 25–46. University of California Press: Berkeley.

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

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

[Y03] Yates, A. M. 2003. A new species of the primitive dinosaur Thecodontosaurus (Saurischia: Sauropodomorpha) and its implications for the systematics of early dinosaurs. Journal of Systematic Palaeontology 1: 1–42.

Last updated: 11 July 2018.

Eusauropoda

Reconstruction of Mamenchisaurus, copyright DiBgd.

Belongs within: Sauropoda.
Contains: Neosauropoda.

The Eusauropoda is a clade of sauropods that has been defined as the smallest clade containing Shunosaurus and Saltasaurus; it contains almost all sauropods known from the Late Jurassic and Cretaceous (Upchurch et al. 2004). Basal members of the clade include Cetiosaurus from the Middle Jurassic of Europe, which was the first sauropod to be named (albeit under the inital mistaken belief that it was some form of marine reptile).

Characters (from Upchurch et al. 2004): Snout broadly rounded in dorsal view; caudal margin of external naris lying behind caudal margin of antorbital fenestra; lateral plate present on premaxillae, maxillae and dentaries; contact between dorsal ends of maxilla and lacrimal lying at caudodorsal margin of antorbital fenestra; platelike flange of bone present at base of maxillary ascending process; rostral process of quadratojugal at least twice as long as dorsal process; quadratojugal dorsal process expanded dorsoventrally at its tip; prefrontal rostral process absent; transverse width of frontals greater than rostrocaudal length; rostral tip of maxillary process of palatine expanded transversely; ectopterygoid process of pterygoid lying below rostral rim of orbit; diameter of external mandibular fenestra less than 10% of mandible length; tooth crown enamel wrinkled; caudalmost tooth lying beneath antorbital fenestra; at least twelve cervical vertebrae present; height of midcervical neural arches greater than diameter of centrum; caudal margin of caudal cervical neural spines sloping strongly forward in lateral view; dorsal surfaces of sacral plates level with dorsal margin of ilium; first caudal rib linked to neural arch by lamina; carpals blocklike; metacarpals arranged in U-shaped colonnade; manual phalanges (apart from unguals) wider transversely than proximodistally; manual digits II–IV having two or fewer phalanges; dorsal margin of ilium strongly convex in lateral view; ischial peduncle of ilium reduced, long axis of iliac blade sloping craniodorsally in lateral view; middle and distal portions of pubis lying in same plane as proximal part; femoral cranial trochanter absent; medial malleolus of tibia reduced, exposing caudal fossa of astragalus in caudal view; lateral muscle scar present at midlength on fibula; metatarsals II–IV diverging distally; ratio of length to proximal transverse width for metatarsal I 1.5 or less; pedal digit IV with three phalanges; pedal phalanges lacking collateral ligament pits; nonungual pedal phalanges wider transversely than proximodistally; pedal digits II–IV with rudimentary penultimate phalanges; ratio of ungual length to metatarsal length for pedal digit I greater than 1.0; unguals for pedal digits II and III with transversely compressed proximal articular ends.

<==Eusauropoda [Bothrosauropodidae, Cetiosauridae, Ornithopsidae, Somphospondylii]
    |  i. s.: Amygdalodon Cabrera 1947 UBD04
    |           `--A. patagonicus Cabrera 1947 UBD04
    |         Aragosaurus Sanz, Buscalioni et al. 1987 UBD04
    |           `--A. ischiaticus Sanz, Buscalioni et al. 1987 UBD04
    |--Shunosaurus Dong, Zhou & Zhang 1983 R-TCA06, UBD04 [Shunosaurinae]
    |    |--S. lii Dong, Zhou & Zhang 1983 UBD04
    |    `--S. ziliujingensis D07
    `--+--Jobaria Sereno, Beck et al. 1999 R-TCA06, UBD04
       |    `--J. tiguidensis Sereno, Beck et al. 1999 UBD04
       `--+--Atlasaurus Monbaron, Russell & Taquet 1999 R-TCA06, UBD04
          |    `--A. imelakei Monbaron, Russell & Taquet 1999 UBD04
          `--+--+--Neosauropoda R-TCA06
             |  `--Turiasauria R-TCA06
             |       |--Galveosaurus R-TCA06
             |       `--+--Losillasaurus Casanovas, Santafé & Sanz 2001 R-TCA06, UBD04
             |          |    `--L. giganteus Casanovas, Santafé & Sanz 2001 UBD04
             |          `--Turiasaurus Royo-Torres, Cobos & Alcalá 2006 R-TCA06
             |               `--*T. riodevensis Royo-Torres, Cobos & Alcalá 2006 R-TCA06
             `--+--Cetiosaurus Owen 1841 R-TCA06, UM02 [Cetiosaurinae]
                |    |--*C. oxoniensis Phillips 1871 UM02 (see below for synonymy)
                |    |--C. brachyurus (n. d.) UM02
                |    |--C. medius Owen 1842 (n. d.) UM02
                |    `--C. mogrebiensis Lapparent 1955 (n. d.) UBD04
                `--+--Patagosaurus Bonaparte 1979 R-TCA06, UBD04
                   |    `--P. fariasi Bonaparte 1979 UBD04
                   `--+--Barapasaurus Jain, Kutty et al. 1975 R-TCA06, UBD04
                      |    `--B. tagorei Jain, Kutty et al. 1975 UBD04
                      `--Euhelopodidae WS98
                           |  i. s.: Hudiesaurus Dong 1997 Y03, UBD04
                           |           `--H. sinojapanorum Dong 1997 UBD04
                           |--+--Tehuelchesaurus Rich, Vickers-Rich et al. 1999 UBD04
                           |  |    `--T. benitezii Rich, Vickers-Rich et al. 1999 UBD04
                           |  `--+--Cetiosauriscus Huene 1927 RR05, UBD04
                           |     |    |--C. glymptonensis (Phillips 1871) [=Cetiosaurus glymptonensis] UBD04
                           |     |    |--C. greppini D07
                           |     |    `--C. stewarti Charig 1980 [incl. Cetiosaurus leedsi Woodward 1905] UBD04
                           |     `--Omeisaurus Young 1939 R-TCA06, UBD04
                           |          |--*O. junghsiensis Young 1939 WS98, UBD04
                           |          |--O. changshouensis Young 1958 (n. d.) UBD04
                           |          |--O. fuxiensis Dong, Zhou & Zhang 1983 (n. d.) UBD04
                           |          |--O. luoquanensis Li 1988 (n. d.) UBD04
                           |          `--O. tianfuensis He, Li et al. 1984 UBD04
                           `--+--Euhelopus Romer 1956 R-TCA06, UBD04 [=Helopus Wiman 1929 (preoc.) UBD04; Euhelopodinae]
                              |    `--E. zdanskyi (Wiman 1929) [=Helopus zdanskyi] UBD04
                              `--Mamenchisaurus Young 1954 R-TCA06, UBD04 (see below for synonymy)
                                   |--M. anyuensis He, Yang et al. 1996 UBD04
                                   |--M. constructus Young 1954 UBD04
                                   |--M. fuxiensis (Hou, Chao & Chu 1976) (see below for synonymy) UBD04
                                   |--M. hochuanensis Young & Chao 1972 UBD04
                                   |--M. jingyanensis Zhang & Li in Zhang & Chen 1996 UBD04
                                   |--M. sinocanadorum Russell & Zheng 1993 UBD04
                                   `--M. youngi Pi, Ouyang & Ye 1996 UBD04

Nomina nuda: Cetiosaurus epioolithicus UM02
             Cetiosaurus hypoolithicus UM02

*Cetiosaurus oxoniensis Phillips 1871 UM02 [=Cardiodon oxoniensis UM02, Ornithopsis oxoniensis UM02; incl. Cetiosaurus giganteus Owen in Huxley 1870 (n. n.) UBD04, Cetiosaurus longus Owen 1875 UM02, Cetiosauriscus longus UBD04]

Mamenchisaurus Young 1954 R-TCA06, UBD04 [incl. Zigongosaurus Hou, Chao & Chu 1976 UBD04; Mamenchisaurinae]

Mamenchisaurus fuxiensis (Hou, Chao & Chu 1976) [=Zigongosaurus fuxiensis, Zizhongosaurus fuxiensis] UBD04

*Type species of generic name indicated

REFERENCES

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

[RR05] Rauhut, O. W. M., K. Remes, R. Fechner, G. Cladera & P. Puerta. 2005. Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435: 670–672.

[R-TCA06] Royo-Torres, R., A. Cobos & L. Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314: 1925–1927.

[UBD04] Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 259–322. University of California Press: Berkeley.

[UM02] Upchurch, P., & J. Martin. 2002. The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur. Palaeontology 45 (6): 1049–1074.

[WS98] Wilson, J. A., & P. C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Journal of Vertebrate Paleontology 18 (2 Suppl.): 1–68.

[Y03] Yates, A. M. 2003. A new species of the primitive dinosaur Thecodontosaurus (Saurischia: Sauropodomorpha) and its implications for the systematics of early dinosaurs. Journal of Systematic Palaeontology 1: 1–42.

Last updated: 6 August 2017.

Camarasauromorpha

Camarasaurus lentus, from here.

Belongs within: Neosauropoda.
Contains: Titanosauria.

The Camarasauromorpha are a clade of sauropod dinosaurs that have been defined as the smallest clade containing Camarasaurus and Saltasaurus (Upchurch et al. 2004). Camarasaurus is the most common sauropod in the Late Jurassic Morrison Formation of North America; it possessed an arched skull and blunt snout.

Characters (from Upchurch et al. 2004): Premaxillary ascending process directed dorsally over most of its length; rostral end of splenial reaching mandibular symphysis; pleurocoels in cranial dorsal centra having acute caudal ends; metacarpal I longer than metacarpal IV; length of longest metacarpal at least 40% radius length.

<==Camarasauromorpha
    |  i. s.: Chondrosteosaurus gigas Owen 1876 (n. d.) N10, UBD04
    |         Oplosaurus Gervais 1852 N10, UBD04
    |           `--O. armatus Gervais 1852 [=Hoplosaurus armatus] UBD04
    |--Lourinhasaurus Dantas, Sanz et al. 1998 R-TCA06, UBD04
    |    `--L. alenquerensis (Lapparent & Zbyszewski 1957) (see below for synonymy) UBD04
    |--Klamelisaurinae D07
    |    |--Bellusaurus Dong 1990 R-TCA06, UBD04
    |    |    `--B. sui Dong 1990 UBD04
    |    `--Klamelisaurus Zhao 1993 D07
    |         `--K. gobiensis Zhao 1993 UBD04
    |--Camarasaurus Cope 1877 (see below for synonymy) UBD04
    |    |--‘Morosaurus’ agilis Marsh 1889 UBD04
    |    |--C. grandis (Marsh 1877) (see below for synonymy) UBD04
    |    |--C. lentus (Marsh 1889) UBD04 (see below for synonymy)
    |    |--C. lewisi (Jensen 1988) [=Cathetosaurus lewisi] UBD04
    |    |--C. supremus Cope 1877 (see below for synonymy) UBD04
    |    `--‘Apatosaurus’ yahnahpin [=Eobrontosaurus yahnahpin] UBD04
    `--Titanosauriformes SL01
         |  i. s.: Lapparentosaurus Bonaparte 1986 UBD04
         |           `--L. madagascariensis Bonaparte 1986 UBD04
         |         Amazonsaurus RR05
         |         Tehuelchesaurus Rich, Vickers-Rich et al. 1999 RR05, UBD04
         |           `--T. benitezii Rich, Vickers-Rich et al. 1999 UBD04
         |--Titanosauria SL01
         `--Brachiosauridae UBD04
              |  i. s.: Pleurocoelus Marsh 1888 N10, UBD04 [Pleurocoelidae]
              |           |--P. altus Marsh 1888 UBD04
              |           |--P. nanus Marsh 1888 UBD04
              |           `--P. valdensis Ludekker 1889 (n. d.) UBD04
              |         Pelorosaurus Mantell 1850 N10, UBD04
              |           |--P. becklesii Mantell 1852 UBD04
              |           `--P. conybearei Mantell 1850 [=Cetiosaurus conybeari] UBD04
              |         Astrodon (n. d.) [Astrodontidae] N85
              |           `--A. johnstoni Leidy 1865 (n. d.) UBD04
              |         Eucamerotus foxi Blows 1995 N10, UBD04
              |         Sauroposeidon Wedel, Cifelli & Sanders 2000 UBD04
              |           `--S. proteles Wedel, Cifelli & Sanders 2000 UBD04
              |         Dystylosaurus Jensen 1985 UBD04
              |           `--D. edwini Jensen 1985 UBD04
              |         ‘Cetiosaurus’ humerocristatus Hulke 1874 UBD04
              |         Cedarosaurus Tidwell, Carpenter & Brooks 1999 UBD04
              |           `--C. weiskopfae Tidwell, Carpenter & Brooks 1999 UBD04
              |--Bothriospondylus Owen 1875 WS98, D07
              |    |--B. elongatus Owen 1875 (n. d.) UBD04
              |    |--B. madagascariensis Lydekker 1895 (n. d.) UBD04
              |    |--B. magnus Owen 1875 (n. d.) UBD04
              |    |--B. robustus Owen 1875 (n. d.) UBD04
              |    `--B. suffosus Owen 1875 (n. d.) UBD04
              `--+--Ornithopsis Seeley 1870 WS98, UBD04
                 |    |--O. eucamerotus Hulke 1882 (n. d.) UBD04
                 |    |--O. greppini Huene 1922 (n. d.) UBD04
                 |    |--O. hulkei Seeley 1870 UBD04
                 |    `--O. leedsi Hulke 1887 (n. d.) UBD04
                 `--Brachiosaurus Riggs 1903 WS98, UBD04 (see below for synonymy)
                      |--B. altithorax Riggs 1903 [=Giraffatitan altithorax] UBD04
                      |--B. atalaiensis Lapparent & Zbyszewski 1957 UBD04 [=Lusotitan atalaiensis D07]
                      |--B. brancai Janensch 1914 UBD04 [=Giraffatitan brancai N10; incl. B. fraasi Janensch 1914 UBD04]
                      `--B. nougaredi Lapparent 1960 UBD04

Brachiosaurus Riggs 1903 WS98, UBD04 [incl. Giraffatitan Paul 1988 UBD04, Lusotitan Teles, Antunes & Mateus 2003 D07; Brachiosaurinae]

Camarasaurus Cope 1877 [incl. Cathetosaurus Jensen 1988, Caulodon Cope 1877, Eobrontosaurus Bakker 1998, Morosaurus Marsh 1878, Uintasaurus Holland 1919; Camarasaurinae, Morosauridae] UBD04

Camarasaurus grandis (Marsh 1877) [=Apatosaurus grandis; incl. Morosaurus impar Marsh 1878, Pleurocoelus montanus Marsh 1896, M. robustus Marsh 1878] UBD04

Camarasaurus lentus (Marsh 1889) UBD04 [=Morosaurus lentus UBD04; incl. C. annae Ellinger 1950 UBD04, *Uintasaurus douglassi Holland 1919 N85, UBD04]

Camarasaurus supremus Cope 1877 [incl. Caulodon diversidens Cope 1877, Amphicoelias latus Cope 1877, Camarasaurus leptodirus Cope 1879, Caulodon leptoganus Cope 1878] UBD04

Lourinhasaurus alenquerensis (Lapparent & Zbyszewski 1957) [=Apatosaurus alenquerensis, Atlantosaurus alequerensis, Camarasaurus alequerensis] UBD04

*Type species of generic name indicated

REFERENCES

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

[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.

[RR05] Rauhut, O. W. M., K. Remes, R. Fechner, G. Cladera & P. Puerta. 2005. Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435: 670–672.

[R-TCA06] Royo-Torres, R., A. Cobos & L. Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314: 1925–1927.

[SL01] Smith, J. B., M. C. Lamanna, K. J. Lacovara, P. Dodson, J. R. Smith, J. C. Poole, R. Giegengack & Y. Attia. 2001. A giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt. Science 292: 1704–1706.

[UBD04] Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 259–322. University of California Press: Berkeley.

[WS98] Wilson, J. A. & P. C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Journal of Vertebrate Paleontology 18 (2 Suppl.): 1–68.

Last updated: 6 August 2017.

Eutitanosauria

Reconstruction of Saltasaurus, from here.

Belongs within: Lithostrotia.

The Eutitanosauria are a clade of titanosaurian sauropods defined by the inclusion of Argyrosaurus, Lirainosaurus and Saltasaurus, together with the "Peirópolis titanosaur" (Wilson & Upchurch 2003).

Eutitanosauria
    |  i. s.: Argyrosaurus Lydekker 1893 WU03
    |           `--A. superbus Lydekker 1893 UBD04
    |--Lirainosaurus Sanz, Powell et al. 1999 UBD04
    |    `--L. astibiae Sanz, Powell et al. 1999 UBD04
    `--+--+--Alamosaurus Gilmore 1922 UBD04
       |  |    `--A. sanjuanensis Gilmore 1922 UBD04
       |  `--Pellegrinisaurus Salgado 1996 UBD04
       |       `--P. powelli Salgado 1996 UBD04
       `--Saltasauridae R-TCA06
            |--Opisthocoelicaudia Borsuk-Bialynicka 1977 UBD04 [Opisthocoelicaudiinae WU03]
            |    `--O. skarzynskii Borsuk-Bialynicka 1977 UBD04
            `--Saltasaurinae WU03
                 |--Neuquensaurus Powell 1992 WU03
                 |    `--N. australis (Lydekker 1893) (see below for synonymy) UBD04
                 `--Saltasaurus Bonaparte & Powell 1980 [incl. Loricosaurus Huene 1929, Microcoelus Ludekker 1893] UBD04
                      |--S. loricatus Bonaparte & Powell 1980 UBD04
                      `--S. robustus (Huene 1929) [=Titanosaurus robustus, Neuquensaurus robustus] UBD04

Neuquensaurus australis (Lydekker 1893) [=Titanosaurus australis, Saltasaurus australis; incl. T. nanus Lydekker 1893, Microcoelus patagonicus Lydekker 1893, Loricosaurus scutatus Huene 1929] UBD04

*Type species of generic name indicated

REFERENCES

[R-TCA06] Royo-Torres, R., A. Cobos & L. Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314: 1925–1927.

[UBD04] Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 259–322. 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.

Titanosauria

Femora of Antarctosaurus wichmannianus, copyright Beatrice Murch.

Belongs within: Camarasauromorpha.
Contains: Eutitanosauria.

The Titanosauria are a clade of sauropods that has been defined as all taxa closer to Saltasaurus than Brachiosaurus. Members of this clade are characterised by a prominent caudolateral expansion of the caudal end of the sternal plate, and an extremely robust radius and ulna with the radius proximal width at least 33% of its length (Upchurch et al. 2004). Many titanosaurs, placed in the clade Lithostrotia, possessed a dorsal armour of scattered dermal osteoderms.

<==Titanosauria [Titanosauridae, Titanosaurinae, Titanosauroidea]
    |--Andesaurus Calvo & Bonaparte 1991 WU03, UBD04 [Andesauridae]
    |    `--A. delgadoi Calvo & Bonaparte 1991 UBD04
    |--Phuwiangosaurus Martin, Buffetaut & Suteethorn 1994 UBD04
    |    `--P. sirindhornae Martin, Buffetaut & Suteethorn 1994 UBD04
    `--+--Austrosaurus Longman 1933 UBD04
       |    `--A. mckillopi Longman 1933 UBD04
       `--Lithostrotia WU03
            |  i. s.: Epachthosaurus Powell 1990 WU03, UBD04
            |           `--E. sciuttoi Powell 1990 UBD04
            |         Aeolosaurus Poell 1987 UBD04
            |           `--A. rionegrinus Powell 1987 UBD04
            |         Ampelosaurus Le Loeuff 1995 UBD04
            |           `--A. atacis Le Loeuff 1995 UBD04
            |         Jainosaurus Hunt, Lockley et al. 1994 UBD04
            |           `--J. septentrionalis (Huene & Matley 1933) [=Antarctosaurus septentrionalis] UBD04
            |         Jiangshanosaurus Tang, Kang et al. 2001 UBD04
            |           `--J. lixianensis Tang, Kang et al. 2001 UBD04
            |         Laplatasaurus Huene 1929 UBD04
            |           `--L. araukanicus Huene 1929 [=Titanosaurus araukanicus] WU03
            |         Magyarosaurus Huene 1932 UBD04
            |           `--M. dacus (Nopcsa 1915) (see below for synonymy) UBD04
            |         Paralititan Smith, Lamanna et al. 2001 SL01
            |           `--*P. stromeri Smith, Lamanna et al. 2001 SL01
            |         Rocasaurus Salgado & Azpilicueta 2000 UBD04
            |           `--R. muniozi Salgado & Azpilicueta 2000 UBD04
            |         Titanosaurus Lydekker 1877 WU03
            |           |--*T. indicus Lydekker 1877 (n. d.) WU03
            |           |--T. blanfordi Lydekker 1879 (n. d.) WU03
            |           |--T. falloti Hoffet 1942 (n. d.) [=Tangvayosaurus falloti] WU03
            |           |--T. madagascariensis Depéret 1896 (n. d.) [=Laplatasaurus madagascariensis] WU03
            |           `--T. rahioliensis Mathur & Srivastava 1987 (n. d.) WU03
            |--+--Gondwanatitan Kellner & de Azevedo 1999 UBD04
            |  |    `--G. faustoi Kellner & de Azevedo 1999 UBD04
            |  `--+--Eutitanosauria WU03
            |     `--Isisaurus Wilson & Upchurchu 2003 WU03
            |          `--*I. colberti (Jain & Bandyopadhyay 1997) [=Titanosaurus colberti] WU03
            `--+--Antarctosaurus Huene 1929 CRF01, WU03
               |    |--*A. wichmannianus Huene 1929 WU03, UBD04
               |    |--A. brasiliensis Arid & Vizotto 1971 (n. d.) UBD04
               |    |--A. giganteus Huene 1929 (n. d.) UBD04
               |    `--A. jaxartensis Riabinin 1939 (n. d.) UBD04
               `--+--Malawisaurus Jacobs, Winkler et al. 1993 UBD04
                  |    `--M. dixeyi (Haughton 1928) [=Gigantosaurus dixeyi] UBD04
                  `--Nemegtosauridae UBD04
                       |--Quaesitosaurus Kurzanov & Bannikov 1983 CRF01, UBD04
                       |    `--Q. orientalis Kurzanov & Bannikov 1983 UBD04
                       `--+--Rapetosaurus Curry Rogers & Forster 2001 CRF01
                          |    `--*R. krausei Curry Rogers & Forster 2001 CRF01
                          `--Nemegtosaurus Nowinski 1971 CRF01, UBD04
                               |--N. mongoliensis Nowinski 1971 UBD04
                               `--N. pachi Dong 1977 (n. d.) UBD04

Titanosauria incertae sedis:
  Aegyptosaurus Stromer 1932 UBD04
    `--A. baharijensis Stromer 1932 UBD04
  Aepysaurus elephantinus Gervais 1853 (n. d.) N85, UBD04
  Algoasaurus bauri Broom 1904 (n. d.) N85, UBD04
  Campylodoniscus Kuhn 1961 (n. d.) N85, UBD04
    `--*C. ameghinoi (Huene 1929) [=Campylodon ameghinoi] UBD04
  Macrurosaurus semnus Seeley 1869 (n. d.) N85, UBD04
  Tangvayosaurus Allain, Taquet et al. 1999 UBD04
    `--T. hoffeti Allain, Taquet et al. 1999 UBD04
  Iuticosaurus Le Loeuff 1993 (n. d.) WU03
    |--*I. valdensis (Huene 1929) (n. d.) [=Titanosaurus valdensis] UBD04
    `--I. lydekkeri (Huene 1929) (n. d.) [=Titanosaurus lydekkeri] WU03
  Hypselosaurus Matheron 1869 D07
    `--H. priscus Matheron 1869 (n. d.) APS03
  Argentinosaurus Bonaparte & Coria 1993 UBD04
    `--A. huinculensis Bonaparte & Coria 1993 UBD04
  Chubutisaurus Corro 1974 UBD04
    `--C. insignis Corro 1974 UBD04
  Mongolosaurus haplodon Gilmore 1933 (n. d.) WS98, UBD04
  Agustinia Bonaparte 1999 UBD04
    `--A. ligabuei Bonaparte 1999 UBD04
  Huabeisaurus Pang & Cheng 2000 UBD04
    `--H. allocotus Pang & Cheng 2000 UBD04
  Janenschia Wild 1991 UBD04
    `--J. robusta (Fraas 1908) [=Gigantosaurus robustus, Tornieria robustus] UBD04
  Venenosaurus Tidwell, Carpenter & Meyer 2001 UBD04
    `--V. dicrocei Tidwell, Carpenter & Meyer 2001 UBD04
  Bruhathkayosaurus Yadagiri & Ayyasami 1989 D07
    `--B. matleyi Yadagiri & Ayyasami 1987 (n. d.) UBD04
  Bonitasaura Apesteguia 2004 D07
    `--B. salgadoi D07

Magyarosaurus dacus (Nopcsa 1915) [=Titanosaurus dacus; incl. M. hungaricus Huene 1932, M. transsylvanicus Huene 1932] UBD04

*Type species of generic name indicated

REFERENCES

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

[CRF01] Curry Rogers, K., & C. A. Forster. 2001. The last of the dinosaur titans: A new sauropod from Madagascar. Nature 412: 530–534.

[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.

[SL01] Smith, J. B., M. C. Lamanna, K. J. Lacovara, P. Dodson, J. R. Smith, J. C. Poole, R. Giegengack & Y. Attia. 2001. A giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt. Science 292: 1704–1706.

[UBD04] Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 259–322. University of California Press: Berkeley.

[WS98] Wilson, J. A. & P. C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Journal of Vertebrate Paleontology 18 (2 Suppl.): 1–68.

[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.

Last updated: 6 August 2017.

Neotheropoda

Skeletal mounts of Coelophysis bauri, copyright Firsfron.

Belongs within: Theropoda.
Contains: Ceratosauria, Tetanurae.

The Neotheropoda are a clade containing the greater part of the theropod dinosaurs, first appearing in the Late Triassic. Three main lineages are recognised within the Neotheropoda—the Coelophysoidea, Ceratosauria and Tetanurae—with most recent analyses placing the Coelophysoidea outside the clade formed by the other two. The Triassic Coelophysis bauri is one of the best known of all fossil dinosaurs, represented by a large number of specimens collected from the Ghost Ranch deposit in New Mexico.

Synapomorphies (from Novas et al. 2015): Nasal process of premaxilla with posterior tip that extending posterior to posterior tip of posterolateral premaxillary process; premaxilla-nasal suture on internarial bar W-shaped; maxillary antorbital fossa shallow, margins formed by low ridges, sharp rim may be present only in front of promaxillary foramen; depth of ventral antorbital fossa much greater than depth of maxilla below ventral margin of antorbital fossa; prefrontal ventral process short and subtriangular or absent; squamosal lateral surface of main body separated from ventral process by sub-horizontal shelf; basisphenoid recess present between basispenoid and basioccipital; dentary rostral end dorsally raised over distance of first three to four alveoli; enlarged, fang-like teeth in anterior part of dentary; pleurocoels in dorsal vertebrae present in anterior dorsals (‘pectorals’); more than three sacral vertebrae present; distal end of metacarpal I with strongly asymmetrical condyles, lateral condyle extending further distally than medial condyle; shaft of metacarpal III considerably more slender than metacarpal II (less than 70% of width of metacarpal II); third finger of manus shorter than second finger; ilium without tuberosity contacting preacetabular process with supraacetabular crest; iliac supraacetabular crest hood-like and extensive, extending laterally and also curving ventrally, to cover dorsal portion of acetabulum in lateral view; brevis fossa very strongly expanded posteriorly; dorsolateral trochanter on proximal femur reduced or absent; femur with mediodistal crest in distal end; ridge on lateral side of tibia for connection with fibula; tibial lateral malleolus tabular; tibia with ventral extension of lateral malleolus distinctly ventral to level of ventral margin of medial malleolus, separated by an inflexion; tibia with concave posterolateral surface of distal end; eeep groove on medial side of proximal end of fibula and medial side of fibula bearing distinct fossa; astragalus without proximally facing basin posterior to base of ascending process; metatarsal I not contacting ankle joint; shaft of metatarsal I contacting metatarsal II along its entire extension; metatarsal I length <50% metatarsal II length; distal articular surface of metatarsal IV taller dorsoplantarlly than broad mediolaterally.

<==Neotheropoda [Halticosaurinae, Paleotheropoda]
    |  i. s.: Dracoraptor hanigani BNB17
    |--+--Zupaysaurus rougieri Arcucci & Coria 2003 XC09, R05
    |  `--+--Dilophosaurus Welles 1970 NS15, TR04
    |     |    |--D. breedorum D07
    |     |    `--D. wetherilli (Welles 1954) [=Megalosaurus wetherilli] TR04
    |     `--+--Dracovenator regenti NS15, BNB17
    |        `--+--‘Dilophosaurus’ sinensis Hu 1993 NS15, TR04
    |           `--Averostra [Intertheropoda] XC09
    |               |--Ceratosauria XC09
    |               `--Tetanurae XC09
    `--Coelophysoidea XC09
         |  i. s.: Longosaurus Welles 1984 N85, TR04
         |           `--*L. longicollis (Cope 1887) [=Coelurus longicollis] TR04
         |         Lukousaurus Young 1948 N85, P88
         |         Avipes Huene 1932 N85, P88
         |         Dolichosuchus cristatus Huene 1932 N85, TR04
         |         Velocipes guerichi Huene 1932 (n. d.) N85, TR04
         |         Gojirasaurus Carpenter 1997 TR04
         |           `--G. quayi Carpenter 1997 TR04
         |         Camposaurus Hunt, Lucas et al. 1998 TR04
         |           `--C. arizonensis Hunt, Lucas et al. 1998 TR04
         |         Podokesaurus Talbot 1911 TR04 [Podokesauridae]
         |           `--P. holyokensis Talbot 1911 [=Coelophysis holyokensis] TR04
         |--Panguraptor lufengensis BNB17
         `--+--+--Procompsognathus Fraas 1913 [Procompsognathidae, Procompsognathinae] TR04
            |  |    `--P. triassicus Fraas 1913 TR04
            |  `--Coelophysidae [Coelophysinae] TR04
            |       |--Segisaurus Camp 1936 [Segisauridae] TR04
            |       |    `--S. halli Camp 1936 TR04
            |       |--Syntarsus Raath 1969 [incl. Megapnosaurus Ivie, Slipinsky & Wegrzynowicz 2001] TR04
            |       |    `--S. kayentakatae Rowe 1989 [=Megapnosaurus kayentakatae] TR04
            |       `--Coelophysis Cope 1889 XC09, TR04
            |            |--*C. bauri (Cope 1889) WU03, TR04 (see below for synonymy)
            |            `--C. rhodesiensis (Raath 1969) XC09, TR04 (see below for synonymy)
            `--+--Cryolophosaurus Hammer & Hickerson 1994 BNB17, HMC04
               |    `--C. ellioti Hammer & Hickerson 1994 HMC04
               |--Sinosaurus Young 1948 BNB17, G00
               |    `--S. triassicus BNB17
               `--Lophostropheus airelensis BNB17

*Coelophysis bauri (Cope 1889) WU03, TR04 [=Coelurus bauri TR04, Rioarribasaurus bauri TR04, Tanystrophaeus bauri TR04; incl. R. colberti Hunt & Lucas 1991 TR04, Syntarsus colberti TR04, Coelophysis longicollis N85, Coelophysis willistoni N85]

Coelophysis rhodesiensis (Raath 1969) XC09, TR04 [=Syntarsus rhodesiensis P88, Megapnosaurus rhodesiensis TR04]

*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.

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

[G00] Gower, D. J. 2000. Rauisuchian archosaurs (Reptilia, Diapsida): an overview. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 218 (3): 447–488.

[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.

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

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

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

[R05] Rauhut, O. W. M. 2005. Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia. Palaeontology 48 (1): 87–110.

[TR04] Tykoski, R. S., & T. Rowe. 2004. Ceratosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 47–70. 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.

[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.

Last updated: 11 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.