Neotheropoda

Skeletal mounts of Coelophysis bauri, copyright Firsfron.

Belongs within: Theropoda.
Contains: Ceratosauria, Tetanurae.

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

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

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

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

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

*Type species of generic name indicated

REFERENCES

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

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

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

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

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

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

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

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

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

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

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

Last updated: 11 July 2018.

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 2^nd 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 2^nd 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 2^nd 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 2^nd ed. pp. 111–136. University of California Press: Berkeley.

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

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

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

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

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

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

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

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

Last updated: 11 July 2018.

Troodontidae

Skeletal reconstruction of Saurornithoides junior, copyright Jaime A. Headden.

Belongs within: Maniraptora.

The Troodontidae are a group of small, cursorial theropods known from the Cretaceous of Asia and North America with a single possible representative, Koparion douglassi, also known from the Upper Jurassic (Makovicky & Norell 2004). Certain fragmentary taxa from the late Cretaceous of Europe, such as Bradycneme draculae and Elopteryx nopcsai, have also been suggested to be troodontids but are not universally accepted as such.

Characters (from Makovicky & Norell 2004, possibly excluding Jinfengopteryx): Dentary bearing groove on at least part of labial face containing exits for neurovascular foramina; quadrate with pneumatic foramen present caudally; basicranium with basisphenoid recess absent; internarial bar dorsoventrally flattened; large number of teeth present; mesial dentary teeth closely packed; laterosphenoid with depression present on ventral surface of postorbital process; reduced basal tubera usually lying directly ventral to occipital condyle; transverse processes elongate; distal caudals with sulcus on dorsal midline in place of neural spine; metatarsus asymmetrical with slender metatarsal II markedly shorter than metatarsals III and IV, metatarsal IV robust.

<==Troodontidae [Saurornithoididae]
    |--Jinfengopteryx GC13
    `--+--Mei Xu & Norell 2004 GC13, XN04
       |    `--*M. long Xu & Norell 2004 XN04
       `--+--Sinovenator Xu, Norell et al. 2002 MN04
          |    `--*S. changii Xu, Norell et al. 2002 XN02
          `--+--Sinusonasus Xu & Wang 2004 GC13, XW04
             |    `--*S. magnodens Xu & Wang 2004 XW04
             `--+--Byronosaurus Norell, Makovicky & Clark 2000 GC13, MN04
                |    `--B. jaffei Norell, Makovicky & Clark 2000 MN04
                `--+--Sinornithoides Russell & Dong 1993 GC13, MN04
                   |    `--S. youngi Russell & Dong 1993 MN04
                   `--+--Zanabazar GC13
                      |--Troodon Leidy 1856 (see below for synonymy) MN04
                      |    |--*T. formosus Leidy 1856 N85 (see below for synonymy)
                      |    |--‘Pectinodon’ asiamericanus Nessov 1995 (n. d.) MN04
                      |    `--T. isfarensis Nessov 1995 (n. d.) MN04
                      `--Saurornithoides Osborn 1924 MN04
                           |--S. asiamericanus D07
                           |--S. isfarensis D07
                           |--S. junior Barsbold 1974 MN04
                           `--S. mongoliensis Osborn 1924 MN04 [=Troodon mongoliensis P88]

Troodontidae incertae sedis:
  Borogovia Osmólska 1987 MN04
    `--B. gracilicrus Osmólska 1987 MN04
  Bradycneme draculae Harrison & Walker 1975 (n. d.) MN04
  Heptasteornis Harrison & Walker 1975 (n. d.) MN04
    `--H. andrewsi Harrison & Walker 1975 [=Troodon andrewsi] P88
  Koparion Chure 1994 H98
    `--K. douglassi Chure 1994 MN04
  Tochisaurus Kurzanov & Osmólska 1991 MN04
    `--T. nemegtensis Kurzanov & Osmólska 1991 MN04
  Elopteryx [Elopterygidae] MN04
    `--E. nopcsai Andrews 1913 (n. d.) MN04
  ‘Laelaps’ cristatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus cristatus O69]
  ‘Laelaps’ explanatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus explanatus O69]
  ‘Laelaps’ laevifrons Cope 1876 (n. d.) MN04 [=Dromaeosaurus laevifrons O69]
  Paronychodon lacustris Cope 1876 (n. d.) MN04
  Zapsalis abradens Cope 1876 (n. d.) MN04

Troodon Leidy 1856 [incl. Pectinodon Carpenter 1982, Polyodontosaurus Gilmore 1932, Stenonychosaurus Sternberg 1932] MN04

*Troodon formosus Leidy 1856 N85 [incl. Pectinodon bakkeri Carpenter 1982 MN04, Polyodontosaurus grandis Gilmore 1932 MN04, Stenonychosaurus inequalis Sternberg 1932 MN04]

*Type species of generic name indicated

REFERENCES

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

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[H98] Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5–61.

[MN04] Makovicky, P. J., & M. A. Norell. 2004. Troodontidae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 184–195. University of California Press: Berkeley.

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

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

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

[XN04] Xu, X., & M. A. Norell. 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature 431: 838–841.

[XN02] Xu, X., M. A. Norell, X.-L. Wang, P. J. Makovicky & X.-C. Wu. 2002. A basal troodontid from the Early Cretaceous of China. Nature 415: 780–784.

[XW04] Xu X. & Wang X. 2004. A new troodontid (Theropoda: Troodontidae) from the Lower Cretaceous Yixian Formation of western Liaoning, China. Acta Geologica Sinica (English Edition) 78: 22–26.

Last updated: 3 July 2018.

Oviraptorosauria

Reconstruction of Caudipteryx zoui, by Matt Martyniuk.

Belongs within: Maniraptora.

The Oviraptorosauria are a group of cursorial theropods known from the Cretaceous of Eurasia and North America. Mostly less than two metres in length, they are characterised by deep, short skulls with the dentition reduced or absent. The ecology of oviraptorosaurs has been subject to much debate; they may have included herbivorous or omnivorous forms.

The majority of oviraptorosaurs belong to a clade Caenagnathoidea, which has been defined as the least inclusive clade including Oviraptor philoceratops and Caenagnathus collinsi. This clade is supported by several synapomorphies, including the presence of two longitudinal ridges and a toothlike ventral process on the palatal shelf of the maxilla, and a jaw joint close to the midline of the skull (Osmólska et al. 2004). Non-caenagnathoid oviraptorosaurs include the early Cretaceous Caudipteryx, the type species of which, C. zoui, is known from fossils in which a fan of feathers is preserved at the end of the tail.

Synapomorphies (from Osmólska et al. 2004): Premaxilla with crenulated ventral margin; parietal at least as long as frontal; ascending process of quadratojugal bordering more than three-quarters of infratemporal fenestra; U-shaped mandibular symphysis present; dentary edentulous; pubic foot with cranial process longer than caudal process.

<==Oviraptorosauria
    |  i. s.: Thecocoelurus Huene 1923 OCB04
    |           `--*T. daviesi (Seeley 1888) OCB04 [=Thecospondylus daviesi OCB04, Coelurus daviesi N02]
    |--+--Protarchaeopteryx Ji & Ji 1997 GC13, P04
    |  |    `--P. robusta Ji & Ji 1997 P04
    |  `--Incisivosaurus Xu, Cheng et al. 2002 XT07, OCB04
    |       `--*I. gauthieri Xu, Cheng et al. 2002 OCB04
    `--+--Similicaudipteryx GC13
       `--+--Avimimus Kurzanov 1981 [Avimimidae] OCB04
          |    `--A. portentosus Kurzanov 1981 OCB04
          `--+--Caudipteryx Ji, Currie et al. 1998 XT07, OCB04 [Caudipterygidae]
             |    |--C. dongi Zhou & Wang 2000 OCB04
             |    `--C. zoui Ji, Currie et al. 1998 OCB04
             |--Microvenator Ostrom 1970 GC13, OCB04
             |    `--M. celer Ostrom 1970 OCB04
             `--Caenagnathoidea [Caenagnathinae, Oviraptoroidea] OCB04
                  |  i. s.: Shixinggia oblita N10
                  |--Caenagnathidae [Elmisauridae] OCB04
                  |    |--Hagryphus giganteus N10
                  |    |--Elmisaurus Osmólska 1981 MOW02, OCB04
                  |    |    `--E. rarus Osmólska 1981 OCB04 [=Chirostenotes rarus P88]
                  |    |--Caenagnathasia Currie, Godfrey & Nessov 1993 OCB04
                  |    |    `--C. martinsoni Currie, Godfrey & Nessov 1993 1994 OCB04
                  |    |--Nomingia Barsbold, Osmólska et al. 2000 OCB04
                  |    |    `--N. gobiensis Barsbold, Osmólska et al. 2000 OCB04
                  |    `--Chirostenotes Gilmore 1924 OCB04 (see below for synonymy)
                  |         |--C. elegans (Parks 1933) OCB04 (see below for synonymy)
                  |         `--C. pergracilis Gilmore 1924 (see below for synonymy) OCB04
                  `--Oviraptoridae XT07
                       |  i. s.: Nemegtomaia barsboldi (Lü et al. 2004) [=Nemegtia barsboldi] N10
                       |--Gigantoraptor Xu, Tan et al. 2007 XT07
                       |    `--*G. erlianensis Xu, Tan et al. 2007 XT07
                       `--+--Oviraptorinae OCB04
                          |    |--Oviraptor Osborn 1924 OCB04
                          |    |    `--O. philoceratops Osborn 1924 OCB04
                          |    |--Rinchenia Barsbold 1997 OCB04
                          |    |    `--R. mongoliensis (Barsbold 1986) [=Oviraptor mongoliensis] OCB04
                          |    `--Citipati Clark, Norell & Barsbold 2001 OCB04
                          |         `--C. osmolskae Clark, Norell & Barsbold 2001 OCB04
                          `--Ingeniinae OCB04
                               |--+--Conchoraptor Barsbold 1986 OCB04
                               |  |    `--C. gracilis Barsbold 1986 OCB04
                               |  `--Khaan Clark, Norell & Barsbold 2001 OCB04
                               |       `--K. mckennai Clark, Norell & Barsbold 2001 OCB04
                               `--+--‘Ingenia’ Barsbold 1981 non Gerlach 1957 OCB04
                                  |    `--I. yanshini Barsbold 1981 OCB04 [=Oviraptor (Ingenia) yanshini P88]
                                  `--Heyuannia Lü 2003 ZZ08, OCB04
                                       `--H. huangi Lü 2003 OCB04

Chirostenotes Gilmore 1924 OCB04 [incl. Caenagnathus Sternberg 1940 OCB04, Macrophalangia Sternberg 1932 N85]

Chirostenotes elegans (Parks 1933) OCB04 [=Ornithomimus elegans OCB04; incl. Caenagnathus sternbergi Cracraft 1971 OCB04, Chirostenotes sternbergi D07]

Chirostenotes pergracilis Gilmore 1924 [incl. Caenagnathus collinsi Sternberg 1940, Macrophalangia canadensis Sternberg 1932] OCB04

*Type species of generic name indicated

REFERENCES

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

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[MOW02] Maryańska, T., H. Osmólska & M. Wolsan. 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica 47 (1): 97–116.

[N02] Naish, D. 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria) Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings of the Geologists' Association 113: 153–163.

[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.

[OCB04] Osmólska, H., P. J. Currie & R. Barsbold. 2004. Oviraptorosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 165–183. University of California Press: Berkeley.

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

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

[XT07] Xu, X., Q. Tan, J. Wang, X. Zhao & L. Tan. 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature 447: 844–847.

[ZZ08] Zhang, F., Z. Zhou, X. Xu, X. Wang & C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105–1108.

Last updated: 11 July 2018.

Tetanurae

Skeletal mount of Monolophosaurus jiangi, copyright Kabacchi.

Belongs within: Neotheropoda.
Contains: Megalosauroidea, Allosauroidea, Coelurosauria.

The clade Tetanurae includes all theropods more closely related to birds than to Ceratosaurus. The name of the clade refers to the development of a stiffened tail; members of the clade are also characterised by the loss of an external fourth digit on the hand. The clade is first definitely recorded from the beginning of the Middle Jurassic (Rauhut 2005).

Most basal members of the Tetanurae are large carnivorous species, the 'carnosaurs' of earlier authors. However, Chilesaurus diegosuarezi from the Late Jurassic (Tithonian) of Chile is a smaller herbivorous form (Novas et al. 2015); this species could be described as superficially similar to one of the basal sauropodomorphs. Other basal tetanurans include Condorraptor currumili and Piatnitzkysaurus floresi from the Middle Jurassic (Callovian) of Argentinean Patagonia, and Marshosaurus bicentesimus from the Late Jurassic (Kimmeridgian) of Utah (Holtz et al. 2004; Rauhut 2005); Piatnitzkysaurus is known from a relatively complete skeleton whereas Condorraptor and Marshosaurus are known from more fragmentary remains. Monolophosaurus jiangi from the Middle Jurassic of China is also known from a complete skull and pelvis and much of the vertebral column; this species is characterised by a distinctive hollow crest along the dorsal midline of the skull (Holtz et al. 2004). The Allosauroidea and Coelurosauria are united in a clade Avetheropoda to the exclusion of megalosauroids by several features including the presence of palatine recesses, an expanded ectopterygoid, laterally displaced zygapophyses on the cervical vertebrae, a widely open ventral floor to the pelvic canal, and L-shaped haemal arches in at least the distal half of the tail (Holtz et al. 2004).

Synapomorphies (from Novas et al. 2015): Metacarpal I closely appressed to proximal half of metacarpal II; metacarpal IV absent; antitrochanter posterior to acetabulum absent or poorly developed; brevis fossa narrow and with subparallel margins; femur with ratio between maximum transverse width of the distal end and total length of the bone of 0.20-0.25; femoral head horizontally directed; well-developed extensor groove present on anterior side of distal femur; tibia with lateral condyle of the proximal end anteriorly displaced from the posterior margin of the medial condyle but well on the posterior half of the bone in proximal view; tibia with deep and broad notch separating the posterior condyles of the proximal end; ridge on lateral side of tibia for connection with fibula present and clearly separated from proximal articular surface.

<==Tetanurae [Allosaurinae, Avipoda, Carnosauria, Eustreptospondylidae, Eustreptospondylinae, Torvosauridae, Torvosauroidea]
    |--Chilesaurus Novas, Salgado et al. 2015 NS15
    |    `--*C. diegosuarezi Novas, Salgado et al. 2015 NS15
    `--+--+--Marshosaurus Madsen 1976 OES10, HMC04
       |  |    `--M. bicentesimus Madsen 1976 HMC04
       |  `--+--Condorraptor Rauhut 2005 NS15, R05
       |     |    `--*C. currumili Rauhut 2005 R05
       |     `--Piatnitzkysaurus Bonaparte 1979 NS15, HMC04
       |          `--P. floresi Bonaparte 1979 HMC04
       `--+--Megalosauroidea NS15
          `--+--Monolophosaurus Zhao & Currie 1993 NS15, HMC04
             |    `--M. jiangi Zhao & Currie 1993 HMC04
             `--Avetheropoda [Allosauria, Neotetanurae] HMC04
                  |  i. s.: Bahariasaurus Stromer 1934 HMC04
                  |           `--B. ingens Stromer 1934 HMC04
                  |         Ozraptor Long & Molnar 1998 HMC04
                  |           `--O. subotaii Long & Molnar 1998 HMC04
                  |         Quilmesaurus Coria 2001 HMC04
                  |           `--Q. curriei Coria 2001 HMC04
                  |         Shanyangosaurus Xue, Zhang & Bi 1996 HMC04
                  |           `--S. niupanggouensis Zue, Zhang & Bi 1996 HMC04
                  |         Valdoraptor Olshevsky 1991 HMC04
                  |           `--V. oweni (Lydekker 1889) [=Megalosaurus oweni] HMC04
                  |--Allosauroidea HMC04
                  `--Coelurosauria HMC04

Tetanurae incertae sedis:
  Becklespinax Olshevsky 1991 HMC04
    `--B. altispinax (Paul 1988) [=Acrocanthosaurus altispinax] HMC04
  Chilantaisaurus Hu 1964 HMC04
    |--C. sibiricus (Riabinin 1914) [=Allosaurus sibiricus] HMC04
    `--C. tashuikouensis Hu 1964 HMC04
  Kaijiangosaurus He 1984 HMC04
    `--K. lini He 1984 HMC04
  Kelmayisaurus Dong 1973 HMC04
    `--K. petrolicus Dong 1973 HMC04
  Sigilmassaurus Russell 1996 [Sigilmassauridae] HMC04
    `--S. brevicollis Russell 1996 HMC04
  ‘Zanclodon’ cambrensis Newton 1899 HMC04
  Chienkosaurus ceratosauroides (Young 1942) (n. d.) HMC04
  Chuandongocoelurus primitivus He 1984 (n. d.) HMC04
  Dandakosaurus indicus Yadagiri 1982 (n. d.) HMC04
  Embasaurus minax Riabinin 1931 (n. d.) HMC04
  Inosaurus tedreftensis Lapparent 1960 (n. d.) HMC04
  Betasuchus Huene 1932 (n. d.) HMC04
    `--*B. bredai (Seeley 1883) [=Megalosaurus bredai] HMC04
  Scrotum humanum Brookes 1763 (n. d.) HMC04
  Siamosaurus suteethorni Buffetaut & Ingavat 1986 (n. d.) HMC04
  Sinocoelurus fragilis Young 1942 (n. d.) HMC04
  Teinurosaurus Nopcsa 1928 HMC04, O69 [=Caudocoelus Huene 1932 HMC04]
    `--*T. sauvagei (Huene 1932) (n. d.) O69 [=*Caudocoelus sauvagei HMC04]
  Wakinosaurus satoi Okazaki 1992 (n. d.) HMC04
  Walgettosuchus woodwardi Huene 1932 (n. d.) [=Magnosaurus woodwardi] HMC04
  Szechuanosaurus Young 1942 HMC04, D07
    |--S. campi Young 1942 HMC04
    |--S. yandonensis Dong, Chang et al. 1978 HMC04
    `--S. zigongensis Gao 1993 HMC04
  Xuanhanosaurus Dong 1984 HMC04
    `--X. qilixiaensis Dong 1984 HMC04

*Type species of generic name indicated

REFERENCES

[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 2^nd 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.

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

[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.

[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.

Last updated: 6 August 2017.

Ceratosauria

Skeletal reconstruction of Elaphrosaurus bambergi by Jaime Headden.

Belongs within: Neotheropoda.

The Ceratosauria is a relatively basal clade of theropods containing all species more closely related to Ceratosaurus than to birds. Though Ceratosaurus itself came from the Jurassic of North America, the ceratosaurians were a primarily Gondwanan clade, with representatives from South America, Africa and India. As well as larger forms such as Ceratosaurus and the Abelisauridae, the ceratosaurs also included smaller forms such as Elaphrosaurus and the Noasauridae. The Chinese Limusaurus inextricabilis and the African Elaphrosaurus are Late Jurassic cursorial forms that have been described as convergent to the later Ornithomimosauria. Deltadromeus agilis is a relatively large theropod (about eight metres in length) known from the mid-Cretaceous Kem Kem beds of northern Africa which may be a basal ceratosaur though some authors have placed it in the ceratosaur sister-clade Tetanurae.

<==Ceratosauria [Neoceratosauria]
    |--Spinostropheus XC09
    |--Deltadromeus Sereno, Duthiel et al. 1996 XC09, HMC04
    |    `--D. agilis Sereno, Duthiel et al. 1996 HMC04
    |--+--Limusaurus Xu, Clark et al. 2009 XC09
    |  |    `--*L. inextricabilis Xu, Clark et al. 2009 XC09
    |  `--Elaphrosaurus Janensch 1920 TR04
    |       |--E. bambergi Janensch 1920 TR04
    |       |--E. gautieri Lapparent 1960 (n. d.) MKC04
    |       `--E. iguidiensis Lapparent 1960 (n. d.) MKC04
    `--Ceratosauroidea WS03
         |--Ceratosauridae P88
         |    |--Sarcosaurus Andrews 1921 P88
         |    |    |--S. andrewsi Huene 1932 TR04
         |    |    `--S. woodi Andrews 1921 TR04
         |    `--Ceratosaurus Marsh 1884 TR04
         |         |--C. dentisulcatus Madsen & Welles 2000 TR04
         |         |--C. magnicornis Madsen & Welles 2000 TR04
         |         |--C. meriani D07
         |         |--C. nasicornis Marsh 1884 TR04
         |         `--C. roechlingi Janensch 1925 (n. d.) TR04
         `--Abelisauroidea [Abelisauria] XC09
              |  i. s.: Genusaurus Accarie, Beaudoin et al. 1995 WS03, TR04
              |           `--G. sisteronis Accarie, Beaudoin et al. 1995 TR04
              |         Ligabueino Bonaparte 1996 F12, TR04
              |           `--L. andesi Bonaparte 1996 TR04
              |         Velocisaurus Bonaparte 1991 F12, TR04
              |           `--V. unicus Bonaparte 1991 TR04
              |         Pycnonemosaurus nevesi F12
              |         Berberosaurus liassicus F12
              |         Kryptops palaios F12
              |--Noasauridae [Noasaurinae] XC09
              |    |--Noasaurus Bonaparte & Powell 1980 TR04
              |    |    `--N. leali Bonaparte & Powell 1980 TR04
              |    `--Masiakasaurus Sampson, Carrano & Forster 2001 TR04
              |         `--*M. knopfleri Sampson, Carrano & Forster 2001 SCF01
              `--Abelisauridae [Abelisaurinae] XC09
                   |  i. s.: Tarascosaurus Le Loeuff & Buffetaut 1991 APS03, TR04
                   |           `--T. salluvicus Le Loeuff & Buffetaut 1991 TR04
                   |         Indosuchus Huene & Matley 1933 WS03
                   |           `--*I. raptorius Huene & Matley 1933 WS03
                   |         Indosaurus Huene & Matley 1933 WS03
                   |           `--*I. matleyi Huene & Matley 1933 WS03
                   |         Xenotarsosaurus Martínez, Gimenez et al. 1986 WS03, TR04
                   |           `--X. bonapartei Martínez, Gimenez et al. 1986 TR04
                   |         Majungasaurus Lavocat 1955 WS03, TR04
                   |           `--M. crenatissimus (Depéret 1896) [=Megalosaurus crenatissimus] TR04
                   |         Rugops Sereno, Wilson & Conrad 2004 XC09, D07
                   |           `--R. primus D07
                   |         Ekrixinatosaurus novasi XC09, F12
                   |         Aucasaurus Coria, Chiappe & Dingus 2002 XC09, TR04
                   |           `--A. garridoi Coria, Chiappe & Dingus 2002 TR04
                   |         Genyodectes serus Woodward 1901 (n. d.) P88, TR04
                   |--Ilokelesia Coria & Salgado 1998 TR04
                   |    `--I. aguadagrandensis Coria & Salgado 1998 TR04
                   `--+--Abelisaurus Bonaparte & Novas 1985 TR04
                      |    `--A. comahuensis Bonaparte & Novas 1985 TR04
                      `--Carnotaurinae TR04
                           |--Rajasaurus Wilson, Sereno et al. 2003 WS03
                           |    `--*R. narmadensis Wilson, Sereno et al. 2003 WS03
                           `--+--Majungatholus Sues & Taquet 1979 TR04
                              |    `--M. atopus Sues & Taquet 1979 TR04
                              `--Carnotaurus Bonaparte 1985 TR04
                                   `--C. sastrei Bonaparte 1985 TR04

Ceratosauria incertae sedis:
  Laevisuchus Huene 1932 TR04
    `--*L. indicus Huene 1932 WS03, TR04
  Coeluroides Huene & Matley 1933 TR04, WS03
    `--*C. largus Huene 1932 (n. d.) WS03, TR04
  Dryptosauroides Huene & Matley 1933 TR04, WS03
    `--*D. grandis Huene 1932 (n. d.) WS03, TR04
  Halticosaurus [Halticosauridae] TR04
    `--H. longotarsus Huene 1907-1908 (n. d.) TR04
  Jubbulpuria Huene & Matley 1933 TR04, WS03
    `--*J. tenuis Huene & Matley 1933 (n. d.) WS03
  ‘Labrosaurus’ stechowi Janensch 1925 (n. d.) TR04
  ‘Labrosaurus’ sulcatus Marsh 1896 (n. d.) TR04
  Ornithomimoides Huene & Matley 1933 TR04, WS03
    |--O. barasimlensis Huene & Matley 1933 (n. d.) TR04
    `--O. mobilis Huene & Matley 1933 (n. d.) TR04
  Orthogoniosaurus matleyi Das-Gupta 1930 (n. d.) TR04
  Pterospondylus trielbae Jaekel 1913-1914 (n. d.) TR04
  Tanystrosuchus Kuhn 1963 (n. d.) TR04
    `--*T. posthumus (Huene 1907–1908) (n. d.) [=Tanystrophaeus posthumus] TR04
  Tanystrophaeus willistoni Cope 1887 (n. d.) TR04

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

[MKC04] Makovicky, P. J., Y. Kobayashi & P. J. Currie. 2004. Ornithomimosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 137–150. University of California Press: Berkeley.

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

[SCF01] Sampson, S. D., M. T. Carrano & C. A. Forster. 2001. A bizarre predatory dinosaur from the Late Cretaceous of Madagascar. Nature 409: 504–506.

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

[WS03] Wilson, J. A., P. C. Sereno, S. Srivastava, D. K. Bhatt, A. Khosla & A. Sahni. 2003. A new abelisaurid (Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian) of India. Contributions from the Museum of Paleontology, The University of Michigan 31 (1): 1–42.

[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.