Pseudosuchia

Mounted skeleton of Postosuchus at Museum of Texas Tech University, copyrigh Dallas Krentzel.

Belongs within: Archosauriformes.
Contains: Stagonolepidae, Crocodylomorpha.

The Pseudosuchia are a clade of reptiles including the crocodilians and a number of related fossil taxa once classified as 'thecodontians' such as the aetosaurs, ornithosuchids and 'rauisuchians'. Higher relationships within the clade have differed between analyses though most recent authors have agreed that the 'rauisuchians' (Rauisuchidae, Prestosuchidae and Poposauridae) do not form a single clade. Indeed, an analysis of the archosauriforms by Nesbitt et al. (2017) did not even recover monophyly for the individual rauisuchian families.

Synapomorphies (from Benton & Clark 1988): Septomaxilla absent; lower temporal fenestra reduced in size, triangular with a dorsal point, quadratojugal sloping forward towards postorbital; axial diapophysis reduced or absent; pubo-ischiadic plate absent, pubis and ischium long and narrow, pubis expanding distally into broad forward-facing plate; long distal process of pubis bent down into subvertical orientation; pubis longer than ischium; proximal head of femur turned inwards at about 45°; fully developed crocodile-normal tarsus, capable of rotation of 60–70°; digit V in foot reduced in length, shorter than digit I.

<==Pseudosuchia (see below for synonymy) NS09
    |--Nundasuchus songeaensis NB17
    `--+--Ornithosuchidae B04
       |    |--Venaticosuchus B04
       |    |--Parringtonia gracilis R72, NB17
       |    |--Dyoplax R72
       |    |--Riojasuchus Bonaparte 1969 IN07, R72
       |    |    `--R. tenuisceps IN07
       |    `--Ornithosuchus Newton 1894 NB17, GQ01 [incl. Dasygnathoides R72]
       |         |--*O. longidens (Huxley 1877) [=Dasygnathus longidens; incl. O. woodwardi Newton 1894] GQ01
       |         `--O. rusconii P88
       `--+--+--Stagonolepidae NB17
          |  `--Revueltosaurus Hunt 1989 NS09, PI05
          |       |--R. callenderi Hunt 1989 PI05
          |       `--R. hunti Heckert 2002 PI05
          `--+--Kollamasuchus gonzalezdiazi NB17
             `--+--+--Turfanosuchus dabanensis NB17
                |  `--+--Yonghesuchus sangbiensis NB17
                |     `--Gracilisuchus Romer 1972 NB17, D07
                |          `--G. stipanicicorum GPS06
                `--+--Ticinosuchus Krebs 1965 NB17, D07 (see below for synonymy)
                   |    `--T. ferox Krebs 1965 G00
                   `--+--+--Qianosuchus mixtus NB17
                      |  `--+--+--Arizonasaurus Welles 1947 NB17, G00
                      |     |  |    `--A. babbitti NS09
                      |     |  `--Xilousuchus sapingensis NB17
                      |     `--+--Poposaurus gracilis Mehl 1915 NB17, G00
                      |        `--+--Lotosaurus Zhang 1975 NB17, D07
                      |           |    `--L. adentus Zhang 1975 G00
                      |           `--+--Sillosuchus longicervix Alcober & Parrish 1997 NB17, G00
                      |              `--+--Shuvosaurus Chatterjee 1993 NB17, D07
                      |                 |    `--S. inexpectatus G00
                      |                 `--Effigia okeeffeae NB17
                      `--+--Prestosuchus chiniquensis Huene 1942 NB17, G00 [incl. P. loricatus Huene 1942 G00]
                         `--+--Saurosuchus Reig 1959 NB17, D07
                            |    `--S. galilei Reig 1959 G00
                            `--+--Batrachotomus kupferzellensis Gower 1999 NB17, G00
                               `--+--Fasolasuchus tenax Bonaparte 1978 NB17, G00
                                  `--+--+--Rauisuchus tiradentes Huene 1942 NB17, G00
                                     |  `--+--Postosuchus NB17
                                     |     |    |--P. alisonae NB17
                                     |     |    `--P. kirkpatricki Chatterjee 1985 G00
                                     |     `--Polonosuchus silesiacus NB17
                                     `--+--Trialestes BC88
                                        |--Erpetosuchus Newton 1894 CX04, D07 [=Herpetosuchus R72]
                                        |    `--E. granti D07
                                        `--Crocodylomorpha NB17

Pseudosuchia incertae sedis:
  Belodon C77
  Strigosuchus R72
  Wangisuchus Young 1974 G00
    `--W. tzeyii G00
  Vytshegdosuchus zheshartensis Sennikov 1988 G00
  Energosuchus garjainovi Ochev 1986 G00
  Dongusia Huene 1940 G00
  Pallisteria R72
  Tsylmosuchus G00
    |--T. donensis Sennikov 1990 G00
    |--T. jakovlevi Sennikov 1990 G00
    `--T. samariensis Sennikov 1990 G00
  Episcoposaurus G00
  Dolichobrachium G00
  Chirotherium G00
  Vjushkovisaurini G00
    |--Vjushkovisaurus berdjanensis Ochev 1982 G00
    |--Jaikosuchus magnus Sennikov 1990 G00
    |--Jushatyria vjushkovi Sennikov in Kalandadze & Sennikov 1985 G00
    `--Youngosuchus Sennikov in Kalandadze & Sennikov 1985 G00
         `--*Y. sinensis (Young 1973) [=Vjushkovia sinensis] G00
  Stagonosuchus nyassicus Huene 1938 G00
  Zanclodon Plieninger 1846 G00
  Tikisuchus romeri Chatterjee & Majumdar 1987 G00
  Luperosuchus fractus Romer 1971 G00
  Hoplitosuchus raui Huene 1942 G00
  Procerosuchus Huene 1944 G00, R72
    `--P. celer Huene 1942 G00
  Heptasuchus clarki Dawley, Zawiskie & Cosgriff 1979 G00
  Lythrosuchus langstoni Long & Murray 1995 G00
  Bromsgroveia walkeri Galton 1985 G00
  Teratosaurus suevicus v. Meyer 1861 G00
  Chatterjeea [Chatterjeeidae] G00
    `--C. elegans Long & Murray 1995 G00

Nomen nudum: Mandasuchus tanyauchen Charig in Appleby et al. 1967 G00

Pseudosuchia [Crocodylotarsi, Crurotarsi, Dromaeosuchia, Poposauridae, Prestosuchidae, Procrocodylomorpha, Rauisuchia, Rauisuchidae, Rauisuchiformes, Rauisuchinae, Rauisuchini, Suchia] NS09

Ticinosuchus Krebs 1965 NB17, D07 [incl. Mandasuchus Charig in Appleby et al. 1967 (n. n.) G00; Ticinosuchinae]

*Type species of generic name indicated

REFERENCES

[B04] Benton, M. J. 2004. Origin and relationships of Dinosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 7–19. University of California Press: Berkeley.

[BC88] Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[CX04] Clark, J. M., X. Xu, C. A. Forster & Y. Wang. 2004. A Middle Jurassic ‘sphenosuchian’ from China and the origin of the crocodylian skull. Nature 430: 1021–1024.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

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

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[GQ01] Gauthier, J., & K. de Queiroz. 2001. Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name "Aves". In: Gauthier, J., & L. F. Gall (eds) New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom pp. 7–41. Peabody Museum of Natural History, Yale University: New Haven.

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

[IN07] Irmis, R. B., S. J. Nesbitt, K. Padian, N. D. Smith, A. H. Turner, D. Woody & A. Downs. 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317: 358–361.

[NB17] Nesbitt, S. J., R. J. Butler, M. D. Ezcurra, P. M. Barrett, M. R. Stocker, K. D. Angielczyk, R. M. H. Smith, C. A. Sidor, G. Niedźwiedzki, A. G. Sennikov & A. J. Charig. 2017. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544: 484–487.

[NS09] Nesbitt, S. J., N. D. Smith, R. B. Irmis, A. H. Turner, A. Downs & M. A. Norell. 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326: 1530–1533.

[PI05] Parker, W. G., R. B. Irmis, S. J. Nesbitt, J. W. Martz & L. S. Browne. 2005. The Late Triassic pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs. Proceedings of the Royal Society of London Series B—Biological Sciences 272: 963–969.

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

[R72] Romer, A. S. 1972. The Chañares (Argentina) Triassic reptile fauna. XVI. Thecodont classification. Breviora 395: 1–24.

Archosauriformes

Reconstruction of Proterosuchus fergusi, copyright Nobu Tamura.

Belongs within: Archosauromorpha.
Contains: Pseudosuchia, Ornithodira, Parasuchia.

The Archosauriformes are a clade of reptiles united by the presence of an antorbital fenestra in the skull (Benton 2004). This clade has been referred to in the past as the Archosauria but that name is used by most current authors to refer to the restricted crown clade including the birds and crocodiles. The Archosauria sensu stricto also include the extinct dinosaurs and pterosaurs. The two basal lineages within the archosaurs, the Pseudosuchia and Avemetatarsalia, have been distinguished in the past by the conformation of the ankle bones; however, the basal avemetatarsalian (stem-bird) Teleocrater rhadinus retains a plesiomorphic 'crocodile-normal' ankle (Nesbitt et al. 2017).

Synapomorphies (from Benton 2004): Antorbital fenestra present; postfrontal reduced to less than half dimensions of postorbital or absent; ossified laterosphenoid present; prootic midline contact on endocranial cavity floor; teeth with serrated margins; lateral mandibular fenestra present.

<==Archosauriformes [Erythrosuchia, Euparkeriidae, Proterosuchia, Thecodontia]
    |  i. s.: Seemannia R72
    |         Crenelosaurus R72
    |         Ocoyuntaia Rusconi 1947 R72
    |         Clarencea R72
    |         Ctenosauriscus koeneni R72, FB17
    |         Hypselorhachis mirabilis R72, NB17
    |         Arcticodactylus cromptonellus NB17
    |         Diandongosuchus fuyuanensis NB17
    |--Proterosuchidae B04
    |    |--Proterosuchus [incl. Ankistrodon, Chasmatosaurus Haughton 1924, Epicampodon] R72
    |    |    |--P. fergusi NB17
    |    |    `--+--P. alexanderi NB17
    |    |       `--+--P. goweri NB17
    |    |          `--‘Chasmatosaurus’ yuani NB17
    |    |--Elaphrosuchus R72
    |    |--Chasmatosuchus R72
    |    `--Archosaurus rossicus R72, NB17
    `--Erythrosuchidae J94
         |--Fugusuchus hejipanensis NB17
         `--+--Sarmatosuchus otschevi NB17
            `--+--Cuyosuchus Rieg 1961 NB17, R72
               |    `--C. huenei NB17
               `--+--+--Guchengosuchus shigualensis NB17
                  |  `--+--+--Erythrosuchus Broom 1905 NB17, D07
                  |     |  |    `--E. africanus B04
                  |     |  `--+--Chalishevia cothurnata NB17
                  |     |     `--Shansisuchus NB17
                  |     |          |--*S. shansisuchus [incl. Fenhosuchus cristatus Young 1964] G00
                  |     |          `--S. heiyuekouensis G00
                  |     `--Garjainia Ochev 1958 NB17, G00 [incl. Vjushkovia Huene 1960 G00]
                  |          |--*G. prima Ochev 1958 [incl. Vjushkovia triplicostata Huene 1960] G00
                  |          `--G. madiba NB17
                  `--+--Dorosuchus neoetus NB17
                     |--Asperoris mnyama NB17
                     `--+--Euparkeria Broom 1913 NB17, D07 [incl. Browniella R72]
                        |    `--E. capensis B04
                        `--+--+--+--Vancleavea campi NB17
                           |  |  `--+--+--Doswellia Weems 1980 J94, D07
                           |  |     |  |    `--D. kaltenbachi D07
                           |  |     |  `--Jaxtasuchus salomoni NB17
                           |  |     `--+--Tarjadia ruthae NB17
                           |  |        `--Archeopelta arborensis NB17
                           |  `--Proterochampsidae J94
                           |       |--Proterochampsa NB17
                           |       |    |--P. barrionuevoi B04
                           |       |    `--P. nodosa NB17
                           |       `--+--Cerritosaurus binsfeldi NB17
                           |          `--+--Tropidosuchus romeri NB17
                           |             `--+--Pseudochampsa ischigualastensis NB17
                           |                `--+--Gualosuchus reigi NB17
                           |                   `--+--Chanaresuchus bonapartei NB17
                           |                      `--Rhadinosuchus Huene 1944 NB17, R72
                           |                           `--R. gracilis NB17
                           `--+--Archosauria [Avesuchia, Ornithosuchia] NB17
                              |    |--Pseudosuchia NS10
                              |    `--Avemetatarsalia [Ornithosuchia, Ornithotarsi, Panaves] NB17
                              |         |--Aphanosauria NB17
                              |         |    |--Spondylosoma Huene 1942 NB17, L04
                              |         |    |    `--S. absconditum Huene 1942 L04
                              |         |    `--+--Teleocrater Nesbitt, Butler et al. 2017 [Aphanosauria] NB17
                              |         |       |    `--*T. rhadinus Nesbitt, Butler et al. 2017 NB17
                              |         |       |--Dongusuchus efremovi Sennikov 1988 NB17, G00
                              |         |       `--Yarasuchus deccanensis Sen 2005 NB17
                              |         `--Ornithodira NB17
                              `--Parasuchia NB17

Inorganic: Chasmatosaurus minilorientalus Okamura 1987 O87

*Type species of generic name indicated

REFERENCES

[B04] Benton, M. J. 2004. Origin and relationships of Dinosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 7–19. University of California Press: Berkeley.

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

[FB17] Feng, Y.-J., D. C. Blackburn, D. Liang, D. M. Hillis, D. B. Wake, D. C. Cannatella & P. Zhang. 2017. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 114 (29): E5864–E5870.

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

[J94] Juul, L. 1994. The phylogeny of basal archosaurs. Palaeontologia Africana 31: 1–38.

[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.

[NB17] Nesbitt, S. J., R. J. Butler, M. D. Ezcurra, P. M. Barrett, M. R. Stocker, K. D. Angielczyk, R. M. H. Smith, C. A. Sidor, G. Niedźwiedzki, A. G. Sennikov & A. J. Charig. 2017. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544: 484–487.

[NS10] Nesbitt, S. J., C. A. Sidor, R. B. Irmis, K. D. Angielczyk, R. M. H. Smith & L. A. Tsuji. 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464: 95–98.

[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.

[R72] Romer, A. S. 1972. The Chañares (Argentina) Triassic reptile fauna. XVI. Thecodont classification. Breviora 395: 1–24.

Pholidosauridae

Reconstruction of Dyrosaurus phosphaticus, copyright Nobu Tamura.

Belongs within: Neosuchia.

The Pholidosauridae are a group of crocodyliforms known from the Cretaceous, most of which are slender-snouted, fish-eating specialists adapted to near-shore marine habitats (Sereno et al. 2001).

<==Pholidosauridae
    |--Pholidosaurus decipiens SL01, GPS06
    `--+--+--Terminonaris ropbusta SL01, GPS06
       |  `--Sarcosuchus Broin & Tarquet 1966 SL01, D07
       |       |--S. hartti SL01
       |       `--S. imperator Broin & Taquet 1966 SL01
       `--+--Sokotosuchus ianwilsoni GPS06
          `--Dyrosauridae GPS06
               |--Congosaurus LB02
               |--Rhabdognathus SF03
               |--Hyposaurus rogersii GPS06
               |--Chenanisuchus lateroculi GPS06
               `--Dyrosaurus Thomas 1893 SL01, D07
                    |--D. minor SF03
                    `--D. phosphaticus GPS06

*Type species of generic name indicated

REFERENCES

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

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

Thalattosuchia

Reconstruction of Geosaurus giganteus scaled to a human diver, from Young et al. (2012).

Belongs within: Neosuchia.

The Thalattosuchia are a clade of marine crocodyliforms known from the Jurassic. Basal members of the clade had an extremely long and tubular hyperdentate snout though this became striking modified in some derived members of the clade (Gasparini et al. 2006).

<==Thalattosuchia
    |--Teleosauridae SF03
    |    |--Steneosaurus bollensis LB02, GPS06
    |    |--Machimosaurus LB02
    |    |--Pelagosaurus Brown 1841 LB02, SF03
    |    |    `--P. typus GPS06
    |    `--Teleosaurus Deslongchamps 1866 D07
    |         |--T. chapmanni D07
    |         |--T. hastifer D07
    |         `--T. mandelslohi D07
    `--Metriorhynchidae GPS06
         |--Metriorhynchus Meyer 1830 GPS06, D07
         |    |--M. casamiquelai GPS06
         |    |--M. moreli D07
         |    `--M. superciliosus GPS06
         `--+--Dakosaurus GPS06
            |    |--D. andiniensis GPS06
            |    `--D. maximus GPS06
            `--Geosaurus Cuvier 1842 GPS06, D07
                 |--G. araucanensis GPS06
                 |--G. cynodus Gervais 1867–1869 C07
                 |--G. giganteus D07
                 |--G. gracilis D07
                 |--G. suevicus GPS06
                 `--G. vignaudi D07

*Type species of generic name indicated

REFERENCES

[C07] Case, E. C. 1907. Revision of the Pelycosauria of North America. Carnegie Institute of Washington Publication 55: 1–176, 35 pls.

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

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

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).

Neosuchia

Spectacled caiman Caiman crocodilus, copyright Berrucomons.

Belongs within: Metasuchia.
Contains: Thalattosuchia, Pholidosauridae, Crocodylidae.

The Neosuchia are a clade of crocodylomorphs that has been defined as including all taxa closer to Crocodylus niloticus than to Notosuchus terrestris. This clade includes all living crocodilians.

<==Neosuchia [Metamesosuchia] SF03
    |  i. s.: Susisuchus Salisbury, Frey et al. 2003 [Susisuchidae] SF03
    |           `--*S. anatoceps Salisbury, Frey et al. 2003 SF03
    |         Brillanceausuchus Michard et al. 1990 SF03, LB02
    |           `--B. babouriensis SF03
    |         Gilchristosuchus palatinus SF03
    |--Atoposauridae GPS06
    |    |--Theriosuchus pusillus GPS06
    |    |--Montsechosuchus depereti [=Alligatorium depereti] BC88
    |    `--Alligatorium [incl. Alligatorellus, Atoposaurus] BC88
    |         `--A. meyeri BC88
    `--+--+--Thalattosuchia GPS06
       |  `--Pholidosauridae GPS06
       `--+--Goniopholidae GPS06
          |    |--Sunosuchus SL01
          |    |--Eutretauranosuchus delfsi GPS06
          |    `--Goniopholis CRA04
          |         |--G. gracilidens [=Nannosuchus gracilidens] BC88
          |         `--G. paulistanus Roxo 1936 CRA04
          `--+--Shamosuchus Mook 1924 BC88, SF03 [incl. Paralligator BC88]
             |--Bernissartia Dollo 1885 GPS06, N85
             |    `--B. fagesii Dollo 1883 SF03
             `--Eusuchia GPS06
                  |--Borealosuchus formidabilis GPS06
                  |--Leidyosuchus BC88
                  |--Hylaeochampsa Owen 1874 GPS06, BC88
                  |    `--H. vectiana GPS06
                  |--Stomatosuchus Stromer 1925 [Stomatosuchidae] BC88
                  |    `--S. inermis SF03
                  `--Crocodylia BC88
                       |  i. s.: Diplocynodon RB05
                       |           |--D. darwini RRO00
                       |           |--D. hantoniensis RRO00
                       |           `--D. styriacus RB05
                       |         Pallimnarchus pollens F71
                       |         Rimasuchus Storrs 2003 LA04
                       |           `--*R. lloydi (Fourtau 1918) [=Crocodylus lloydi] LA04
                       |--+--Gavialidae AS09
                       |  |    |--Gavialis gangeticus GPS06
                       |  |    `--Tomistoma Müller 1846 AS09, LA04 [Tomistominae]
                       |  |         |--T. coppensi Pickford 1994 LA04
                       |  |         |--T. dowsoni Fourtau 1918 LA04
                       |  |         |--T. lusitanica Antunes 1967 LA04
                       |  |         `--T. schlegelii USDI77
                       |  `--Eosuchus lerichei GPS06
                       `--+--+--Crocodylidae GPS06
                          |  `--Osteolaemus tetraspis BC88, USDI77
                          |       |--O. t. tetraspis USDI77
                          |       `--O. t. osborni USDI77
                          `--Alligatoroidea D07
                               |  i. s.: Deinosuchus Colbert & Bird 1958 D07 [incl. Phobosuchus N85]
                               |           |--D. hatchery D07
                               |           |--D. riograndensis D07
                               |           `--D. rugosus D07
                               `--Alligatorinae ML14
                                    |--Navajosuchus mooki ML14
                                    `--+--+--Stangerochampsa mccabei AS09
                                       |  `--Alligator GPS06
                                       |       |--A. mississippiensis Daudin 1801 NS10
                                       |       `--A. sinensis SF03
                                       `--+--Paleosuchus BC88
                                          |    |--P. palpebrosus DS86
                                          |    `--P. trigonatus DS86
                                          `--Caiman BC88
                                               |  i. s.: C. sclerops GWN71
                                               |         C. yacare USDI77
                                               |--C. latirostris BC88
                                               `--+--C. crocodilus BC88
                                                  |    |--C. c. crocodilus B81
                                                  |    |--C. c. apaporiensis USDI77
                                                  |    `--C. c. fuscus B81
                                                  `--Melanosuchus niger BC88, DS86

Inorganic: Caiman crocodilus protominilorientalus Okamura 1987 O87

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

[BC88] Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[B81] Brooks, D. R. 1981. Hennig's parasitological method: a proposed solution. Systematic Zoology 30 (3): 229–249.

[CRA04] Carvalho, I. de S., L. C. B. Ribeiro & L. dos S. Avilla. 2004. Uberabasuchus terrificus sp. nov., a new Crocodylomorpha from the Bauru Basin (Upper Cretaceous), Brazil. Gondwana Research 7 (4): 975–1002.

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

[DS86] Dixon, J. R., & P. Soini. 1986. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum: Milwaukee.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[GWN71] Gorman, G. C., A. C. Wilson & M. Nakanishi. 1971. A biochemical approach towards the study of reptilian phylogeny: evolution of serum albumin and lactic dehydrogenase. Systematic Zoology 20 (2): 167–185.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[LA04] Llinás Agrasar, E. 2004. Crocodile remains from the Burdigalian (lower Miocene) of Gebel Zelten (Libya). Geodiversitas 26 (2): 309–321.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[NS10] Nesbitt, S. J., C. A. Sidor, R. B. Irmis, K. D. Angielczyk, R. M. H. Smith & L. A. Tsuji. 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464: 95–98.

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

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

[RRO00] Rossmann, T., M. Rauhe & F. Ortega. 2000. Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions. Paläontologische Zeitschrift 74: 379–392.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3-47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

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.

Anchisauria

Reconstruction of Melanorosaurus readi, copyright Steveoc 86.

Belongs within: Sauropodomorpha.

The Anchisauria is a clade of prosauropods defined by Galton & Upchurch (2004) as the smallest clade including Anchisaurus and Melanorosaurus.

Characters (from Galton & Upchurch 2004): Length of prefrontal subequal to that of frontal; frontal excluded from margin of supratemporal fossa; at least five teeth present on premaxilla; ratio of forelimb length to hindlimb length greater than 0.60; pubis with pubic foramen completely visible in cranial view; femoral shaft straight in cranial or caudal view; femoral fourth trochanter displaced to caudomedial margin of shaft; proximal end of metatarsal II hourglass-shaped.

<==Anchisauria GU04
    |--Anchisauridae GU04
    |    |--Ammosaurus Marsh 1891 GU04
    |    |    `--A. major (Marsh 1889) [=Anchisaurus major; incl. An. solus Marsh 1892] GU04
    |    `--Anchisaurus Marsh 1885 GU04 (see below for synonymy)
    |         `--*A. polyzelus (Hitchcock 1865) Y03 (see below for synonymy)
    `--Melanorosauridae GU04
         |--Riojasaurus Bonaparte 1969 GU04
         |    `--R. incertus Bonaparte 1969 GU04
         `--+--Melanorosaurus Haughton 1924 GU04
            |    |--M. readi Haughton 1924 GU04
            |    `--M. thabanensis Gauffre 1993 GU04
            `--+--Camelotia Galton 1985 GU04
               |    `--C. borealis Galton 1985 GU04
               `--Lessemsaurus Bonaparte 1999 GU04
                    `--L. sauropoides Bonaparte 1999 GU04

Anchisaurus Marsh 1885 [=Amphisaurus Marsh 1882 (preoc.), Megadactylus Hitchcock 1865 (preoc.), Yaleosaurus Huene 1932] GU04

*Anchisaurus polyzelus (Hitchcock 1865) Y03 [=Megadactylus polyzelus GU04; incl. A. colurus Marsh 1891 GU04, Yaleosaurus colurus N85]

*Type species of generic name indicated

REFERENCES

[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.

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

[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.

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.