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 2nd 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 2nd 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 2nd 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 2nd 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.

Allosauroidea

Skeletal mount of Allosaurus fragilis, from here.


Belongs within: Tetanurae.

The Allosauroidea are a clade of large theropods known from the Late Jurassic and Cretaceous (Ortega et al. 2010). The best known species is Allosaurus fragilis, of which numerous remains have been recovered from the late Jurassic Morrison Formation of the United States; average members of this species reached about 8.5 metres in length.

<==Allosauroidea
    |  i. s.: Piveteausaurus Taquet & Welles 1977 APS03, HMC04
    |           `--P. divesensis (Walker 1964) HMC04 (see below for synonymy)
    |         Erectopus Huene 1921 APS03, HMC04
    |           |--E. sauvagei Huene 1932 HMC04
    |           `--E. superbus (Sauvage 1882) [=Megalosaurus superbus] HMC04
    |--Sinraptoridae OES10
    |    |  i. s.: Yangchuanosaurus Dong, Chang et al. 1978 HMC04
    |    |           |--Y. longqiaoensis D07
    |    |           |--Y. magnus Dong, Zhou & Zhang 1983 HMC04
    |    |           |--Y. shangyouensis Dong, Chang et al. 1978 HMC04 [=Metriacanthosaurus shangyouensis P88]
    |    |           `--Y. yandonensis D07
    |    |--Lourinhanosaurus Mateus 1998 OES10, HMC04
    |    |    `--L. antunesi Mateus 1998 HMC04
    |    `--+--Metriacanthosaurus Walker 1964 OES10, HMC04 [Metriacanthosaurinae]
    |       |    `--M. parkeri (Huene 1923) [=Megalosaurus parkeri] HMC04
    |       `--Sinraptor Currie & Zhao 1993 OES10, HMC04
    |            |--S. dongi Currie & Zhao 1993 [=S. heping (l. c.)] HMC04
    |            `--S. hepingensis (Gao 1992) [=Yangchuanosaurus hepingensis] HMC04
    `--+--Allosauridae HMC04
       |    |--Saurophaganax Chure 1996 [incl. Saurophagus Ray 1941] HMC04
       |    |    `--S. maximus (Ray 1941) HMC04 [=Saurophagus maximus HMC04, Allosaurus maximus D07]
       |    `--Allosaurus Marsh 1877 OES10, HMC04 (see below for synonymy)
       |         |--A. fragilis Marsh 1877 HMC04 (see below for synonymy)
       |         |--A. medius Marsh 1888 (n. d.) HMC04
       |         |--‘Creosaurus’ potens Lull 1911 (n. d.) HMC04
       |         |--A. tendagurensis Janensch 1925 HMC04
       |         `--A. whitei D07
       `--Carcharodontosauria OES10
            |--Neovenatoridae OES10
            |    |  i. s.: Siamotyrannus Buffetaut, Suteethorn & Ton 1996 HMC04
            |    |           `--S. isanensis Buffetaut, Suteethorn & Tong 1996 HMC04
            |    |--Neovenator Hutt, Martill & Barker 1996 OES10, HMC04
            |    |    `--N. salierii Hutt, Martill & Barker 1996 HMC04
            |    `--+--+--Aerosteon OES10
            |       |  `--Megaraptor Novas 1998 OES10, NM04
            |       |       `--M. namunhuaiquii Novas 1998 NM04
            |       `--+--Australovenator OES10
            |          `--Fukuiraptor Azuma & Currie 2000 OES10, HMC04 [incl. Katsuyamasaurus D07, Kitadanisaurus D07]
            |               `--F. kitadaniensis Azuma & Currie 2000 HMC04
            `--Carcharodontosauridae OES10
                 |  i. s.: Bahariasaurus Stromer 1934 F12, HMC04
                 |           `--B. ingens Stromer 1934 HMC04
                 |--Eocarcharia dinops OES10, F12
                 `--+--Concavenator Ortega, Escaso & Sanz 2010 OES10
                    |    `--*C. corcovatus Ortega, Escaso & Sanz 2010 OES10
                    `--+--Acrocanthosaurus Stovall & Langston 1950 OES10, HMC04
                       |    `--A. atokensis Stovall & Langston 1950 HMC04
                       `--+--Saochilong OES10
                          `--+--Tyrannotitan chubutensis OES10, F12
                             `--+--Carcharodontosaurus Stromer 1931 OES10, HMC04
                                |    `--C. saharicus (Depéret & Savornin 1927) (see below for synonymy) HMC04
                                `--+--Mapusaurus roseae OES10, F12
                                   `--Giganotosaurus Coria & Salgado 1995 XW12, HMC04
                                        `--G. carolinii Coria & Salgado 1995 HMC04

Allosaurus Marsh 1877 OES10, HMC04 [incl. Antrodemus Leidy 1870 HMC04, Apatodon N85, Creosaurus Marsh 1878 HMC04, Empaterias N85, Epanterias Cope 1878 HMC04, Epantherias N85, Labrosaurus Marsh 1879 HMC04]

Allosaurus fragilis Marsh 1877 HMC04 [=Antrodemus fragilis P88; incl. Epanterias amplexus Cope 1878 HMC04, Allosaurus amplexus P88, Creosaurus atrox Marsh 1878 HMC04, Al. atrox HMC04, Hypsirhophus discurus Cope 1878 HMC04, Al. ferox Marsh 1896 HMC04, Labrosaurus ferox HMC04, Al. lucaris Marsh 1878 HMC04, Laelaps trihedrodon Cope 1877 HMC04, Al. trihedrodon D07, Poicilopleuron valens Leidy 1870 HMC04, Antrodemus valens N85]

Carcharodontosaurus saharicus (Depéret & Savornin 1927) [=Megalosaurus saharicus; incl. M. africanus Huene 1956] HMC04

Piveteausaurus divesensis (Walker 1964) HMC04 [=Eustreptospondylus divesensis HMC04, Proceratosaurus divesensis P88]

*Type species of generic name indicated

REFERENCES

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

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

[F12] Fanti, F. 2012. Cretaceous continental bridges, insularity, and vicariance in the Southern Hemisphere: which route did the dinosaurs take? In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 883–911. Springer.

[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 2nd ed. pp. 71–110. University of California Press: Berkeley.

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

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

[OES10] Ortega, F., F. Escaso & J. L. Sanz. 2010. A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain. Nature 467: 203–206.

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

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

Last updated: 11 July 2018.

Megalosauroidea

Lower jaw of Megalosaurus bucklandii, copyright Ghedoghedo.


Belongs within: Tetanurae.

The Megalosauroidea is a clade of large theropods known from the Middle Jurassic (Aalenian) to the mid-Cretaceous (Cenomanian) (Holtz et al. 2004). In some references this clade is referred to as the Spinosauroidea but the name Megalosauroidea has priority. Megalosaurus bucklandii from the Middle Jurassic of England was one of the first fossil dinosaurs recognised; many other predatory theropods have been subsequently referred to this genus but most (if not all) are currently agreed to be either misplaced or not diagnostic.

Synapomorphies (from Holtz et al. 2004, as Spinosauroidea): Premaxillary body rostral to external nares longer than portion of body ventral to nares, angle between rostral margin and alveolar margin less than 70°; maxila rostral ramus present, rostrocaudally as long or longer than dorsoventrally; lacrimal rostral ramus dorsoventrally pinched and narrow; deltopectoral crest greater than 45% of humerus length, well developed; humerus with well-developed olecranon fossa on caudal face of distal end.

<==Megalosauroidea [Spinosauroidea] NS15
    |  i. s.: Altispinax Huene 1923 N85, HMC04
    |           `--*A. dunkeri (Dames 1884) [=Megalosaurus dunkeri] HMC04
    |         Magnosaurus Huene 1932 OES10, HMC04
    |           `--M. nethercombensis (Huene 1926) [=Megalosaurus nethercombensis] HMC04
    |--+--Dubreuillosaurus NS15
    |  |--‘Poekilopleuron’ valesdunensis Allain 2002 HMC04
    |  `--Afrovenator Sereno, Wilson et al. 1994 NS15, HMC04
    |       `--A. abakensis Sereno, Wilson et al. 1994 HMC04
    `--+--+--+--Streptospondylus Meyer 1832 NS15, HMC04
       |  |  |    |--S. altdorfensis Meyer 1832 [incl. Laelaps gallicus Cope 1867] HMC04
       |  |  |    `--S. cuvieri Owen 1842 (n. d.) HMC04 [=Megalosaurus cuvieri P88]
       |  |  `--Eustreptospondylus Walker 1964 NS15, HMC04
       |  |       `--E. oxoniensis Walker 1964 [=Magnosaurus oxoniensis] HMC04
       |  `--Spinosauridae HMC04
       |       |--Baryonychinae HMC04
       |       |    |--Baryonyx Charig & Milner 1986 NS15, HMC04
       |       |    |    `--B. walkeri Charig & Milner 1986 HMC04
       |       |    |--Suchomimus Sereno, Beck et al. 1998 HMC04
       |       |    |    `--S. tenerensis Sereno, Beck et al. 1998 HMC04
       |       |    `--Cristatusaurus Taquet & Russell 1998 HMC04
       |       |         `--C. lapparenti Taquet & Russell 1998 HMC04
       |       `--Spinosaurinae HMC04
       |            |--Irritator Martill, Cruickshank et al. 1996 HMC04
       |            |    `--I. challengeri Martill, Cruickshank et al. 1996 HMC04
       |            |--Angaturama Kellner & Campos 1996 HMC04
       |            |    `--A. limai Kellner & Campos 1996 HMC04
       |            `--Spinosaurus Stromer 1915 HMC04
       |                 |--S. aegyptiacus Stromer 1915 HMC04
       |                 `--S. maroccanus Russell 1996 HMC04
       `--Megalosauridae [Megalosaurinae] B08
            |--Torvosaurus Galton & Jensen 1979 XC09, HMC04 [incl. Edmarka Bakker, Kralis et al. 1992 HMC04]
            |    `--T. tanneri Galton & Jensen 1979 (see below for synonymy) HMC04
            |--Poekilopleuron Eudes-Deslongchamps 1838 HMC04 (see below for synonymy)
            |    |--*P. bucklandii Eudes-Deslongchamps 1838 AC02 (see below for synonymy)
            |    `--P. schmidti Kiprijanov 1883 (n. d.) HMC04
            |--Duriavenator Benson 2008 B08
            |    `--*D. hesperis (Waldman 1974) [=Megalosaurus hesperis] B08
            `--Megalosaurus Buckland 1824 HMC04
                 |--*M. bucklandii Ritgen 1826 AC02, HMC04
                 |--M. chubutensis Corro 1974 (n. d.) HMC04
                 |--M. dabukaensis D07
                 |--M. hungaricus Nopcsa 1901 (n. d.) HMC04
                 |--M. inexpectatus Corro 1966 (n. d.) HMC04
                 |--M. ingens Janensch 1920 (n. d.) HMC04 [=Ceratosaurus ingens P88]
                 |--M. insignis Eudes-Deslongchamps 1870 (n. d.) HMC04
                 |--M. lonzeensis Dollo 1883 (n. d.) HMC04
                 |--M. lydekkeri Huene 1926 (n. d.) [=Magnosaurus lydekkeri] HMC04
                 |--M. meriani Greppin 1870 (n. d.) HMC04
                 |--M. pannoniensis Seeley 1881 (n. d.) HMC04
                 |--M. phillipsi D07
                 |--M. pombali Lapparent & Zbyszewski 1957 (n. d.) HMC04
                 |--M. terquemi Huene 1926 (n. d.) HMC04
                 `--M. tibetensis D07

Poekilopleuron Eudes-Deslongchamps 1838 HMC04 [=Poecilopleuron (l. c.) AC02, Poecilopleurum (l. c.) AC02, Poicilopleuron (l. c.) AC02, Poikilopleuron (l. c.) AC02]

Poekilopleuron bucklandii Eudes-Deslongchamps 1838 AC02 [=Poecilopleuron bucklandi (l. c.) AC02, Poecilopleurum bucklandi (l. c.) AC02, Poekilopleuron bucklandi (l. c.) AC02, Poicilopleuron bucklandi (l. c.) AC02, Poikilopleuron bucklandi (l. c.) AC02, Megalosaurus poikilopleuron Huene 1926 AC02, Poekilopleuron poikilopleuron P88]

Torvosaurus tanneri Galton & Jensen 1979 [=Megalosaurus tanneri; incl. Edmarka rex Bakker, Kralis et al. 1992] HMC04

*Type species of generic name indicated

REFERENCES

[AC02] Allain, R., & D. J. Chure. 2002. Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Palaeontology 45 (6): 1107–1121.

[B08] Benson, R. B. J. 2008. A redescription of 'Megalosaurus' hesperis (Dinosauria, Theropoda) from the Inferior Oolite (Bajocian, Middle Jurassic) of Dorset, United Kingdom. Zootaxa 1931: 57–67.

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

[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 2nd ed. pp. 71–110. University of California Press: Berkeley.

[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.

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

[OES10] Ortega, F., F. Escaso & J. L. Sanz. 2010. A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain. Nature 467: 203–206.

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

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

Last updated: 11 July 2018.

Tyrannosauroidea

Skeletal mount of Tyrannosaurus rex, copyright Scott Robert Anselmo.


Belongs within: Coelurosauria.

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

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

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

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

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

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

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

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

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

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

*Type species of generic name indicated

REFERENCES

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

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

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

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

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

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

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

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

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

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

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

Last updated: 11 July 2018.

Ornithurae

Reconstruction of Enaliornis barretti, copyright Nobu Tamura.


Belongs within: Ornithothoraces.
Contains: Aves.

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

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

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

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

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

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

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

*Type species of generic name indicated

REFERENCES

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

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

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

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

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

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

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

Last updated: 6 August 2017.