Last time, we looked at the anatomy of Tyrannosaurus rex. This week, we will look at where it fits in relation to other dinosaurs.
The following cladograms (adapted from Tweet, 2013) illustrate Tyrannosaurus rex's position. The groups to which Tyrannosaurus rex belongs will be placed at the bottom.
Cladogram of Dinosauria (Owen, 1842)
O - Dinosauria
|---o Ornithischia (Seeley, 1888)
`---o Saurischia (Seeley, 1888)
The Ornithischia contains the horned dinosaurs, the armoured dinosaurs and the duck-billed dinosaurs. The Saurischia contains the meat-eating and the long-necked dinosaurs.
O - Saurischia
|--- o Herrerasauridae (Reig, 1963)
|--- o Sauropodomorpha (Huene, 1932)
`---o Theropoda (Marsh, 1881)
The Herrerasauridae contain a small group of dinosaurs that have bounced between the Sauropodomorpha and the Theropoda since...well, forever really (and have sometimes believed to be non-dinosaurs). The Sauropodomorpha contains the Sauropods and their relatives. Theropoda contains all the meat-eating dinosaurs.
O - Theropoda
|---o Daemonosaurus (Sues et al., 2011)
`---+---o Tawa (Nesbitt et al., 2009)
`---o Neotheropoda (Bakker, 1986)
Daemonosaurus and Tawa are the most basal ("primitive") well-established Theropods (if you discount the Herrerasaurids that is). The Neotheropoda contains all the other Theropods, being characterised by only four fingers (as opposed to five), a small fourth finger (and in some, no fourth finger at all), and three weight-bearing toes (sometimes with a fourth toe higher up that is not in contact with the ground).
O - Neotheropoda
| ---o Coelophysoidea (Holtz, 1994)
`---+---o Zupaysaurus (Arcucci and Coria, 2003)
`---+---o Dilophosauridae (Madsen and Welles, 2000)
`---o Averostra (Paul, 2002)
The Coelophysoids were basal theropods that were typically small with long, low bodies, long necks and heads and short hands. Zupaysaurus is somewhere in between the Coelophysoids and the Dilophosaurids. The Dilophosaurids were effectively larger versions of Coelophysoids with shorter necks and they had a sort of kink in their jaws like crocodiles do. The Averostra consists of all other theropods which, unlike the Coelophysoids and the Dilophosaurids have an extra hole in their upper jaw called the promaxillary fenestra.
O - Averostra
|---o Ceratosauria (Marsh, 1884)
`---o Tetanurae (Gauthier, 1986)
The Ceratosaurians are an arbitrary group of Averostrans that aren't Tetanurans but no-one can agree on their characteristics or members. The Tetanurae consists of theropods that lack a fourth digit on their hand, have all their upper jaw teeth situated in front of the eyes and have a strap-like shoulder-blade.
O - Tetanurae
|---o Chuandongocoelurus (He, 1984)
|---o Monolophosaurus (Zhao and Currie, 1993)
`---+---o Megalosauroidea (Huxley, 1869)
`---o Avetheropoda (Paul, 1988)
The Megalosauroids have long, low builds and short, stout arms (being used as fishing implements by some). The Avetheropods, as their name would suggest, contain the bird-like theropods.
O - Avetheropoda
|---o Carnosauria (Huene, 1920)
`---o Coelurosauria (Huene, 1914)
Traditionally, the big theropods were in the Carnosauria and the small theropods in Coelurosauria. This has now been abandoned. In general the Carnosaurs had large eyes, long, narrow skulls and thighs longer than their shins. The Coelurosaurs have a long sacrum, a stiffened tail (especially at the tip), a bowed ulna (one the forearm bones) and shins longer than their thighs.
O - Coelurosauria
|---o Compsognathidae (Cope, 1871)
`---+---o "Bunch of well-known coelurosaurs that don't fit anywhere"
`---o Tyrannoraptora (Sereno, 1999)
The Compsognathids were tiny coelurosaurs. The Tyrannoraptora are all the coelurosaurs closer to birds than the Compsognathids. In between, are a large number of dinosaurs that don't fit anywhere else. In fact, some have suggested that the classification of basal coelurosaurs needs to be completely re-written given the number of taxa that don't have a home and the fact that neither the Compsognathids nor the Tyrannoraptorans have an agreed-upon set of characteristics.
O Tyrannoraptora
|---+---o Coelurus (Marsh, 1879)
| `---o Maniraptoriformes (Holtz, 1995)
`---o Tyrannosauroidea (Walker, 1964)
Coelurus is another critter that changes address every few years. Maniraptoriformes contains all the theropods closer to birds than to Tyrannosaurus. They are generally characterised by having bird-like shoulders and breastbones. The Tyrannosauroids are what we are interested in.
O Tyrannosauroidea
|--? Bagaraatan (Osmolska, 1996)
`---+---o Proceratosauridae (Rauhut et al., 2010)
|---o Dilong (Xu et al., 2004)
`---+---o Yutyrannus (Xu et al., 2012)
`---+---+---o Eotyrannus (Hutt et al., 2001)
| |---o Juratyrant (Brusatte and Benson, 2012)
| `---o Stokesosaurus (Madsen, 1974)
|---o Xiongguanlong (Li et al., 2010)
`---+---o Dryptosaurus (Marsh, 1877)
`---+---o Appalachiosaurus (Carr et al., 2005)
`---+---o Bistahieversor (Carr and Williamson, 2010)
`---o Tyrannosauridae (Osborn, 1905)
Bagaraatan might actually be a basal coelurosaurian instead (Unfortunately, because Tyrannosauroids branched off from the family tree so early on, basal coelurosaurians and basal tyrannosauroids are very similar). The Proceratosaurids had crests. The dinosaurs between the Proceratosaurids and the Tyrannosaurids, possessed three fingers on their hands and were relatively small (with the notable exception of Yutyrannus being 9m long). Quite a few of them also had feather-like coatings. The Tyrannosaurids were big and bad with the teeth at the front of the jaws D-shaped in cross-section and two fingers on their hands.
O Tyrannosauridae
|---o Albertosaurinae (Currie et al., 2003)
`---o Tyrannosaurinae (Matthew and Brown, 1922)
The Albertosaurines were generally smaller and possessed small horns in front of their eyes. The Tyrannosaurines were generally larger and possessed large bumps (called bosses) behind their eyes. The next tree is the last one, I promise.
O Tyrannosaurinae
|---o Alioramus (Kurzanov, 1976)
`---+---o Teratophoneus (Carr et al., 2011)
`---+---o Daspletosaurus (Russell, 1970)
`---+---o Tarbosaurus (Maleev, 1955)
`---o Tyrannosaurus (Osborn, 1905)
There are two types of Tyrannosaur researcher: the lumpers and the splitters. The lumpers regard all of these taxa to be synonymous with Tyrannosaurus whereas the splitters split them into into multiple genera and species (some of which we will look at in Part 3). I was hoping to cover them in this post, but considering this took me two weeks to do, I decided to leave it here. Next time, we'll look at life before the dinosaurs.
See also:
More dinosaurs
More Tyrannosaurus rex
References:
Arcucci, A. and Coria, R., (2003) 'A new Triassic carnivorous dinosaur from Argentina', Ameghiniana, 40 (2), pp. 217-228
Bakker, R. (1986) The Dinosaur Heresies, New York: William Morrow
Brusatte, S. and Benson, R. (2012) 'The systematics of Late Jurassic tyrannosauroids (Dinosauria: Theropoda) from Europe and North America', Acta Palaeontologica Polonica, 58 (1), pp. 47-54
Carr, T., Williamson, T. and Schwimmer, D. (2005) 'A new genus and species of tyrannosauroid from the Late Cretaceous (middle Campanian) Demopolis Formation of Alabama', Journal of Vertebrate Paleontology, 25 (1), pp. 119-143
Carr, T. and Williamson, T. (2010) 'Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea', Journal of Vertebrate Paleontology, 30 (1), pp. 1-16
Carr, T., Williamson, T., Britt, B. and Stadtman, K. (2011) 'Evidence for high taxonomic and morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits Formation of Utah', Naturwissenschaften, 98 (3), pp. 241-246
Cope, E. (1871) 'Supplement to the synopsis of the extinct Batrachia and Reptilia of North America', Proceedings of the American Philosophical Society, 12 (86), pp. 41-52
Currie, P., Hurum, H. and Sabath, K. (2003) 'Skull structure and evolution in tyrannosaurid dinosaurs', Acta Palaeontologica Polonica, 48 (2), pp. 227-234
Gauthier, J. (1986) 'Saurischian monophyly and the origin of birds', in Padian, K. (ed) 'The Origin of Birds and the Evolution of Flight', Memoirs of the California Academy of Science, 8, pp. 1-55
He, X. (1984) [The Vertebrate Fossils of Sichuan], Chengdu: Sichuan Scientific and Technical Publishing House
Holtz, T. (1994) 'The phylogenetic position of the Tyrannosauridae: Implications for theropod systematics', Journal of Paleontology, 68 (5), pp. 1100-1117
Holtz, T. and Padian, K. (1995) 'Definition and diagnosis of Theropoda and related taxa', Journal of Vertebrate Paleontology, 15 (supplement), p. 35A
Huene, F. von (1914) 'Das naturliche System der Saurischia' [The natural System of Saurischia], Zentralblatt fur Mineralogie, Geologie und Palaeontologie B, 1914, pp. 154-158
Huene, F. von (1920) 'Bemerkungen zur Systematik und Stammesgeschichte einiger Reptilien' [Notes on the Systematics and Phylogeny of some Reptiles], Zeitschrift fur Induktive Abstammungs-und Vererbungslehre, 22 (3), pp. 209-212
Huene, F. von (1932) 'Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte', [The fossil reptile order Saurischia, their development and history], Monographien zur Geologie und Palaeontologie, 1 (4), pp. 1-361
Hutt, S., Naish, D., Martill, D., Barker, M. and Newbery, P. (2001) 'A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England' Cretaceous Research, 22 (2), pp. 227-242
Huxley, T. (1869) 'On the upper jaw of Megalosaurus', Quarterly Journal of the Geological Society of London, 25 (1), pp. 311-314
Kurzanov, S. (1976) '[A new Late Cretaceous carnosaur from Nogon-Tsav, Mongolia], in Kramarenko, N., Luvsandansan, B., Voronin, Y., Barsbold, R., Rozhdestvensky, A., Trofimov, B. and Reshitov, V. (Eds) 'Palaeontology and Biostratigraphy of Mongolia', The Joint Soviet-Mongolian Palaeontological Expedition, Transactions, 3, pp. 93-104
Li, D., Norell, M., Gao, K-Q., Smith, N. and Makovicky, P. (2010) 'A longirostrine tyrannosauroid from the Early Cretaceous of China', Proceedings of the Royal Society B, 277 (1679), pp. 183-190
Madsen, J. (1974) 'A new theropod dinosaur from the Upper Jurassic of Utah', Journal of Paleontology, 48 (1), pp. 27-31
Madsen, J. and Welles, S. (2000) Ceratosaurus (Dinosauria, Theropoda) a revised osteology, Salt Lake City: Utah Geological Survey
Maleev, E. (1955) '[New carnivorous dinosaurs from the Upper Cretaceous of Mongolia], Doklady Akademii Nauk SSSR, 104 (5), pp. 779-783
Marsh, O. (1877) 'Notice of some new dinosaurian reptiles from the Jurassic Formation', American Journal of Science, 3 (14), pp. 514-516
Marsh, O. (1878) 'Notice of new Jurassic reptiles', American Journal of Science, 3 (18), pp. 501-505
Marsh, O. (1881) 'Principle characters of American Jurassic dinosaurs. Part V.', American Journal of Science, 3 (21), pp. 417-423
Marsh, O. (1884) 'Principle characters of American Jurassic dinosaurs. Part VIII. The Order Theropoda', American Journal of Science, 3 (27), pp. 329-340
Matthew, W. and Brown, B. (1922) 'The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta', Bulletin of the American Museum of Natural History, 46, pp. 367-385
Nesbitt, S., Smith, N., Irmis, R., Turner, R., Downs, A. and Norell, M. (2009) 'A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs', Science, 326 (5959), pp. 1530-1533
Osborn, H. (1905) 'Tyrannosaurus and other Cretaceous carnivorous dinosaurs', Bulletin of the American Museum of Natural History, 21, pp. 259-265
Osmolska, H. (1996) 'An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia', Acta Palaeontologica Polonica, 41 (1), pp. 1-38
Owen, R. (1842) 'Report on British fossil reptiles. Part II', in, British Association for the Advancement of Science, Report of the Eleventh Meeting of the British Association for the Advancement of Science, London: John Murray, pp. 60-204
Paul, G. (1988) Predatory Dinosaurs of the World: A Complete Illustrated Guide, New York: Simon and Schuster
Paul, G. (2002) Dinosaurs of the Air, Baltimore: The Johns Hopkins University Press
Rauhut, O., Milner, A. and Moore-Fay, S. (2010) 'Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England', Zoological Journal of the Linnean Society, 158 (1), pp. 155-195
Reig, O. (1963) 'La prescencia de dinosaurios saurisquios en los "Estrados de Ischigualasto" (Mesotriásico superior) de las Provincias de San Juan y La Rioja (Republica Argentina)' [The presence of saurischian dinosaurs in the "Ischigualasto formation" (Late Middle Triassic) in the provinces of San Juan and La Rioja (Republic of Argentina)], Ameghiniana, 3 (1), pp. 3-20
Russell, D. (1970) 'Tyrannosaurs from the Late Cretaceous of western Canada', National Museum of Natural Sciences Publications in Paleontology, 1 (1), pp. 1-34
Seeley, H. (1888) 'The classification of the Dinosauria', Proceedings of the Royal Society of London, 43 (258-265), pp. 165-171
Sereno, P. (1999) 'The evolution of Dinosaurs', Science, 284 (5423), pp. 2137-2147
Sues, H-D., Nesbitt, S., Berman, D. and Henrici, A., (2011) 'A late-surviving basal theropod dinosaur from the Latest Triassic of North America', Proceedings of the Royal Society B, 278 (1723), pp. 3459-3464
Tweet, J. Thescelosaurus, [Online] Available at: http://www.thescelosaurus.com/index.html, Accessed on: 13/06/2013
Walker, A. (1964) 'Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs', Philosophical Transactions of the Royal Society B, 248 (744), pp. 53-134
Xu, X., Norell, M., Kuang, X., Wang, X., Zhao, Q. and Jia, C. (2004) 'Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids', Nature, 431 (7009), pp. 680-684
Xu, X., Wang, K., Zhang, K., Ma, Q., Xing, L., Sullivan, C., Hu, D., Cheng, S. and Wang, S. (2012) 'A gigantic feathered dinosaur from the Lower Cretaceous of China, Nature, 484 (7392), pp. 92-95
Zhao, X-J. and Currie, P. (1993) 'A large crested theropod from the Jurassic of Xinjiang, People's Republic of China', in Currie, P. (ed) 'Results from the Sino-Canadian Dinosaur Project', Canadian Journal of Earth Sciences, 30 (10), pp. 2027-2036
The following cladograms (adapted from Tweet, 2013) illustrate Tyrannosaurus rex's position. The groups to which Tyrannosaurus rex belongs will be placed at the bottom.
Cladogram of Dinosauria (Owen, 1842)
O - Dinosauria
|---o Ornithischia (Seeley, 1888)
`---o Saurischia (Seeley, 1888)
The Ornithischia contains the horned dinosaurs, the armoured dinosaurs and the duck-billed dinosaurs. The Saurischia contains the meat-eating and the long-necked dinosaurs.
O - Saurischia
|--- o Herrerasauridae (Reig, 1963)
|--- o Sauropodomorpha (Huene, 1932)
`---o Theropoda (Marsh, 1881)
The Herrerasauridae contain a small group of dinosaurs that have bounced between the Sauropodomorpha and the Theropoda since...well, forever really (and have sometimes believed to be non-dinosaurs). The Sauropodomorpha contains the Sauropods and their relatives. Theropoda contains all the meat-eating dinosaurs.
O - Theropoda
|---o Daemonosaurus (Sues et al., 2011)
`---+---o Tawa (Nesbitt et al., 2009)
`---o Neotheropoda (Bakker, 1986)
Daemonosaurus and Tawa are the most basal ("primitive") well-established Theropods (if you discount the Herrerasaurids that is). The Neotheropoda contains all the other Theropods, being characterised by only four fingers (as opposed to five), a small fourth finger (and in some, no fourth finger at all), and three weight-bearing toes (sometimes with a fourth toe higher up that is not in contact with the ground).
O - Neotheropoda
| ---o Coelophysoidea (Holtz, 1994)
`---+---o Zupaysaurus (Arcucci and Coria, 2003)
`---+---o Dilophosauridae (Madsen and Welles, 2000)
`---o Averostra (Paul, 2002)
The Coelophysoids were basal theropods that were typically small with long, low bodies, long necks and heads and short hands. Zupaysaurus is somewhere in between the Coelophysoids and the Dilophosaurids. The Dilophosaurids were effectively larger versions of Coelophysoids with shorter necks and they had a sort of kink in their jaws like crocodiles do. The Averostra consists of all other theropods which, unlike the Coelophysoids and the Dilophosaurids have an extra hole in their upper jaw called the promaxillary fenestra.
O - Averostra
|---o Ceratosauria (Marsh, 1884)
`---o Tetanurae (Gauthier, 1986)
The Ceratosaurians are an arbitrary group of Averostrans that aren't Tetanurans but no-one can agree on their characteristics or members. The Tetanurae consists of theropods that lack a fourth digit on their hand, have all their upper jaw teeth situated in front of the eyes and have a strap-like shoulder-blade.
O - Tetanurae
|---o Chuandongocoelurus (He, 1984)
|---o Monolophosaurus (Zhao and Currie, 1993)
`---+---o Megalosauroidea (Huxley, 1869)
`---o Avetheropoda (Paul, 1988)
The Megalosauroids have long, low builds and short, stout arms (being used as fishing implements by some). The Avetheropods, as their name would suggest, contain the bird-like theropods.
O - Avetheropoda
|---o Carnosauria (Huene, 1920)
`---o Coelurosauria (Huene, 1914)
Traditionally, the big theropods were in the Carnosauria and the small theropods in Coelurosauria. This has now been abandoned. In general the Carnosaurs had large eyes, long, narrow skulls and thighs longer than their shins. The Coelurosaurs have a long sacrum, a stiffened tail (especially at the tip), a bowed ulna (one the forearm bones) and shins longer than their thighs.
O - Coelurosauria
|---o Compsognathidae (Cope, 1871)
`---+---o "Bunch of well-known coelurosaurs that don't fit anywhere"
`---o Tyrannoraptora (Sereno, 1999)
The Compsognathids were tiny coelurosaurs. The Tyrannoraptora are all the coelurosaurs closer to birds than the Compsognathids. In between, are a large number of dinosaurs that don't fit anywhere else. In fact, some have suggested that the classification of basal coelurosaurs needs to be completely re-written given the number of taxa that don't have a home and the fact that neither the Compsognathids nor the Tyrannoraptorans have an agreed-upon set of characteristics.
O Tyrannoraptora
|---+---o Coelurus (Marsh, 1879)
| `---o Maniraptoriformes (Holtz, 1995)
`---o Tyrannosauroidea (Walker, 1964)
Coelurus is another critter that changes address every few years. Maniraptoriformes contains all the theropods closer to birds than to Tyrannosaurus. They are generally characterised by having bird-like shoulders and breastbones. The Tyrannosauroids are what we are interested in.
O Tyrannosauroidea
|--? Bagaraatan (Osmolska, 1996)
`---+---o Proceratosauridae (Rauhut et al., 2010)
|---o Dilong (Xu et al., 2004)
`---+---o Yutyrannus (Xu et al., 2012)
`---+---+---o Eotyrannus (Hutt et al., 2001)
| |---o Juratyrant (Brusatte and Benson, 2012)
| `---o Stokesosaurus (Madsen, 1974)
|---o Xiongguanlong (Li et al., 2010)
`---+---o Dryptosaurus (Marsh, 1877)
`---+---o Appalachiosaurus (Carr et al., 2005)
`---+---o Bistahieversor (Carr and Williamson, 2010)
`---o Tyrannosauridae (Osborn, 1905)
Bagaraatan might actually be a basal coelurosaurian instead (Unfortunately, because Tyrannosauroids branched off from the family tree so early on, basal coelurosaurians and basal tyrannosauroids are very similar). The Proceratosaurids had crests. The dinosaurs between the Proceratosaurids and the Tyrannosaurids, possessed three fingers on their hands and were relatively small (with the notable exception of Yutyrannus being 9m long). Quite a few of them also had feather-like coatings. The Tyrannosaurids were big and bad with the teeth at the front of the jaws D-shaped in cross-section and two fingers on their hands.
O Tyrannosauridae
|---o Albertosaurinae (Currie et al., 2003)
`---o Tyrannosaurinae (Matthew and Brown, 1922)
The Albertosaurines were generally smaller and possessed small horns in front of their eyes. The Tyrannosaurines were generally larger and possessed large bumps (called bosses) behind their eyes. The next tree is the last one, I promise.
O Tyrannosaurinae
|---o Alioramus (Kurzanov, 1976)
`---+---o Teratophoneus (Carr et al., 2011)
`---+---o Daspletosaurus (Russell, 1970)
`---+---o Tarbosaurus (Maleev, 1955)
`---o Tyrannosaurus (Osborn, 1905)
There are two types of Tyrannosaur researcher: the lumpers and the splitters. The lumpers regard all of these taxa to be synonymous with Tyrannosaurus whereas the splitters split them into into multiple genera and species (some of which we will look at in Part 3). I was hoping to cover them in this post, but considering this took me two weeks to do, I decided to leave it here. Next time, we'll look at life before the dinosaurs.
See also:
More dinosaurs
More Tyrannosaurus rex
References:
Arcucci, A. and Coria, R., (2003) 'A new Triassic carnivorous dinosaur from Argentina', Ameghiniana, 40 (2), pp. 217-228
Bakker, R. (1986) The Dinosaur Heresies, New York: William Morrow
Brusatte, S. and Benson, R. (2012) 'The systematics of Late Jurassic tyrannosauroids (Dinosauria: Theropoda) from Europe and North America', Acta Palaeontologica Polonica, 58 (1), pp. 47-54
Carr, T., Williamson, T. and Schwimmer, D. (2005) 'A new genus and species of tyrannosauroid from the Late Cretaceous (middle Campanian) Demopolis Formation of Alabama', Journal of Vertebrate Paleontology, 25 (1), pp. 119-143
Carr, T. and Williamson, T. (2010) 'Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea', Journal of Vertebrate Paleontology, 30 (1), pp. 1-16
Carr, T., Williamson, T., Britt, B. and Stadtman, K. (2011) 'Evidence for high taxonomic and morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits Formation of Utah', Naturwissenschaften, 98 (3), pp. 241-246
Cope, E. (1871) 'Supplement to the synopsis of the extinct Batrachia and Reptilia of North America', Proceedings of the American Philosophical Society, 12 (86), pp. 41-52
Currie, P., Hurum, H. and Sabath, K. (2003) 'Skull structure and evolution in tyrannosaurid dinosaurs', Acta Palaeontologica Polonica, 48 (2), pp. 227-234
Gauthier, J. (1986) 'Saurischian monophyly and the origin of birds', in Padian, K. (ed) 'The Origin of Birds and the Evolution of Flight', Memoirs of the California Academy of Science, 8, pp. 1-55
He, X. (1984) [The Vertebrate Fossils of Sichuan], Chengdu: Sichuan Scientific and Technical Publishing House
Holtz, T. (1994) 'The phylogenetic position of the Tyrannosauridae: Implications for theropod systematics', Journal of Paleontology, 68 (5), pp. 1100-1117
Holtz, T. and Padian, K. (1995) 'Definition and diagnosis of Theropoda and related taxa', Journal of Vertebrate Paleontology, 15 (supplement), p. 35A
Huene, F. von (1914) 'Das naturliche System der Saurischia' [The natural System of Saurischia], Zentralblatt fur Mineralogie, Geologie und Palaeontologie B, 1914, pp. 154-158
Huene, F. von (1920) 'Bemerkungen zur Systematik und Stammesgeschichte einiger Reptilien' [Notes on the Systematics and Phylogeny of some Reptiles], Zeitschrift fur Induktive Abstammungs-und Vererbungslehre, 22 (3), pp. 209-212
Huene, F. von (1932) 'Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte', [The fossil reptile order Saurischia, their development and history], Monographien zur Geologie und Palaeontologie, 1 (4), pp. 1-361
Hutt, S., Naish, D., Martill, D., Barker, M. and Newbery, P. (2001) 'A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England' Cretaceous Research, 22 (2), pp. 227-242
Huxley, T. (1869) 'On the upper jaw of Megalosaurus', Quarterly Journal of the Geological Society of London, 25 (1), pp. 311-314
Kurzanov, S. (1976) '[A new Late Cretaceous carnosaur from Nogon-Tsav, Mongolia], in Kramarenko, N., Luvsandansan, B., Voronin, Y., Barsbold, R., Rozhdestvensky, A., Trofimov, B. and Reshitov, V. (Eds) 'Palaeontology and Biostratigraphy of Mongolia', The Joint Soviet-Mongolian Palaeontological Expedition, Transactions, 3, pp. 93-104
Li, D., Norell, M., Gao, K-Q., Smith, N. and Makovicky, P. (2010) 'A longirostrine tyrannosauroid from the Early Cretaceous of China', Proceedings of the Royal Society B, 277 (1679), pp. 183-190
Madsen, J. (1974) 'A new theropod dinosaur from the Upper Jurassic of Utah', Journal of Paleontology, 48 (1), pp. 27-31
Madsen, J. and Welles, S. (2000) Ceratosaurus (Dinosauria, Theropoda) a revised osteology, Salt Lake City: Utah Geological Survey
Maleev, E. (1955) '[New carnivorous dinosaurs from the Upper Cretaceous of Mongolia], Doklady Akademii Nauk SSSR, 104 (5), pp. 779-783
Marsh, O. (1877) 'Notice of some new dinosaurian reptiles from the Jurassic Formation', American Journal of Science, 3 (14), pp. 514-516
Marsh, O. (1878) 'Notice of new Jurassic reptiles', American Journal of Science, 3 (18), pp. 501-505
Marsh, O. (1881) 'Principle characters of American Jurassic dinosaurs. Part V.', American Journal of Science, 3 (21), pp. 417-423
Marsh, O. (1884) 'Principle characters of American Jurassic dinosaurs. Part VIII. The Order Theropoda', American Journal of Science, 3 (27), pp. 329-340
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