After Carrano, Benson & Sampson, 2012
Note:
Phylogenetic position of megalosaurs has been, and probably still is, uncertain. These basal tetanurans are rather poorly defined and their phylogenetic position has been highly variable in different analyses. In some they have been associated with spinosaurids (Smith, Makovicky, Hammer & Currie, 2007; Carrano, Benson & Sampson, 2012), in others removed from the relations that group and treated as paraphyletic assemblage of basal Tetanurae (Holtz, 2000 (1998)).
The phylogenetic analysis of basal Tetanurae and Allosauroids by Carrano et al., on which this presentation is based, have once again reinstated megalosaurs as monophyletic group including pianitzkysaurids, megalosaurids and spinosaurids, with some orphan genera.
<==o Tetanurae Gauthier, 1986 [Avipoda Novas, 1992 (non Avepoda Paul, 2002] |-- †‘Dilophosaurus’ sinensis Hu, 1993 |-- †Chilesaurus diegosuarezi Novas, Salgado, Suarez, Agnolin, Ezcurra, Chimento, de la Cruz, Isasi, Vargas & Rubilar-Rogers, 2015; L.Jur. SWSAm. |-- †Cryolophosaurus ellioti Hammer & Hickerson, 1994 `--+-- †Chuandongocoelurus primitivus He, 1984 `--+-- †Monolophosaurus jiangi Zhao & Currie, 1995 `--o Orionides Carrano, Benson & Sampson, 2012 |--o †Megalosauroidea (Fitzinger, 1843) Walker, 1964 [Spinosauroidea (Stromer, 1915) Olshevsky, 1995; Torvosauroidea (Jensen, 1985) Sereno et al., 1994] | |--o †Piatnitzkysauridae Carrano, Benson & Sampson, 2012 | | |-- †Marshosaurus bicentesimus Madsen, 1976b | | `--+-- †Condorraptor currumili Rauhut, 2005 | | `-- †Piatnitzkysaurus floresi Bonaparte, 1979b | `--o †Megalosauria Bonaparte, 1850 | |?- †Sciurumimus albersdoerferi Rauhut, Foth, Tischlinger & Norell, 2012; L.Jur. CEu. | |?- †Streptospondylus altdorfensis Meyer, 1832 [†"Lealaps" gallicus Cope, 1867; Streptospondylus cuvieri (Owen, 1842) Huene, 1932; Eustreptospondylus divesensis Walker, 1964 partim] | |-- †Eustreptospondylus oxoniensis Walker, 1964 | `--+--o †Spinosauridae Stromer, 1915 [incl. Baryonychidae Charig & Milner, 1986; Irritatoridae Martill et al., 1996] | | |?- †Suchosaurus cultridens (Owen, 1841) Owen, 1842 [Crocodilus cultridens Owen, 1841] [Spinosauridae indet. sensu Carrano et al., 2012] | | |?- †Suchosaurus girardi Sauvage, 1897-1898 [Spinosauridae indet. sensu Carrano et al., 2012] | | |?- †Siamosaurus suteethorni Buffetaut & Ingavat, 1986 | | |?- †Ostafrikasaurus crassiserratus Buffetaut, 2011; L.Jur. EAf. | | |--o †Baryonychinae Charig & Milner, 1986 [Baryonychidae Charig & Milner, 1986] | | | |-- †Baryonyx walkeri Charig & Milner, 1986 | | | `-- †Suchomimus tenerensis Sereno et al., 1998 [incl. Cristatusaurus lapparenti Taquet & Russell, 1998 sensu Carrano et al., 2012] [cogeneric with Baryonyx?] | | `--o †Spinosaurinae Stromer, 1915 sensu Sereno et al., 1998 [incl. Irritatoridae Martill et al., 1996] | | |-- †Angaturama limai Kellner & Campos, 1996 | | |-- †Irritator challengeri Martill, Cruikshank, Frey, Small & Clarke, 1996 [incl.? Angaturama limai Kellner & Campos, 1996] [Irritatoridae Martill et al., 1996] | | `--o †Spinosaurus Stromer, 1915 | | |-- †S. aegypticus Stromer, 1915 | | `?- †S. maroccanus Russell, 1996 [cospecific with S. aegypticus? ] | `--o †Megalosauridae Fitzinger, 1843 [Megalosauridae Huxley, 1869; Torvosauridae; Eustreptospondylidae Paul, 1988] | |--o †Afrovenatorinae Carrano, Benson & Sampson, 2012 | | |-- †Afrovenator abakensi Sereno, Wilson, Larsson, Dutheil & Sues, 1994 | | |-- †Dubreuillosaurus valesdunensis (Allain, 2002) Allain, 2005 [†Poekilopleuron? valesdunensis Allain, 2002] | | |-- †Leshansaurus qianweiensis Li, Peng, Ye, Jiang & Huang, 2009; L.Jur. EAs. | | |-- †Magnosaurus nethercombensis (Huene, 1926b) [†"Megalosaurus" nethercombensis Huene, 1926b] | | |-- †Piveteausaurus divesensis (Walker, 1964) [Eustreptospondylus divesensis Walker, 1964 partim] [see Spinosauroidea] | | `-- †Poekilopleuron bucklandii Eudes-Deslongchamps, 1838 [taxonomic history] | `--o †Megalosaurinae (Fitzinger, 1843) Paul, 1988 | |-- †Duriavenator hesperis (Waldman, 1974) Benson, 2008 [Megalosaurus hesperis Waldman, 1974] | `--+-- †Megalosaurus bucklandii Ritgen, 1926 Mantell, 1827 [Megalosaurus Buckland, 1824] L.Jur. WEu. | `--o †Torvosaurus Galton & Jensen, 1979a; L.Jur. WNAm. SWEu. | |-- †T. tanneri Galton & Jensen, 1979a | `-- †T. gurneyi Hendrickx & Mateus, 2014 `--o Avetheropoda Paul, 1988 [Neotetanurae Sereno et al., 1994] (avetheropods) |-- †Allosauroidea (Marsh, 1878) Currie & Zhao, 1994 [Carnosauria Huene, 1920 (in partim); Allosauria Paul, 1988] (allosaurs and relatives; allosaurit) `-- Coelurosauria von Huene, 1914 sensu Holtz, 1999 (coelurosaurs; keelurosaurit)
Tetanurae incertae sedis:
†Becklespinax altispinax (Huene, 1923) [non Paul, 1988?] sensu Olshevsky, 1991 [Megalosaurus dunkeri Dames, 1884; Altispinax dunkeri (Dames, 1884) Huene, 1923; Acrocanthosaurus altispinax (Huene, 1923) sensu Paul, 1988]
†“Brontoraptor” sp. Redman, 1995 [nomen nudum]
†Cruxicheiros newmanorum Benson & Radley, 2010; M.Jur. WEu.
†Edmarka rex Bakker, Krails, Siegwarth & Filla, 1992
†Kaijiangosaurus lini He, 1984
†Megalosaurus? cambrensis (Newton, 1899) [Zanclodon cambrensis Newton, 1899]
†Megalosaurus ingens Janensch, 1920 [Tetanurae indet. sensu Carrano et al., 2012]
†“Saltriosaurus” sp. Dal Sasso, 2000 [nomen nudum]
†Sarcosaurus andrewsi Huene, 1932 [“Megalosaurus” andrewsi]
†Sigilmassasaurus brevicollis Russell, 1996
†Chingkankousaurus fragilis Young, 1958a
†Iliosuchus incognitus Huene, 1932 [Iliosuchidae Paul, 1998]
†Kelmayisaurus petrolicus Dong, 1973
Alternative historical phylogenies of basal Tetanurae (mostly as represented in Carrano et al., 2012):
After Rauhut, Foth, Tischlinger & Norell, 2012<==o Theropoda |-- †Syntarsus kayentakatae `--+-- †Dilophosaurus `--+-- †Ceratosauria `--o Tetanurae |-- †Chuandongocoelurus `--+-- Coelurosauria `--o †Carnosauria |-- †Allosauroidea `--o †Megalosauroidea |--+-- †Xuanhanosaurus | `--+-- †Marshosaurus | `--+-- †Condorraptor | `-- †Piatnitzkysaurus `--+-- †Monolophosaurus `--+-- †Spinosauridae `--o †Megalosauridae |-- †Sciurumimus `--+-- †Duriavenator |-- †Eustreptospondylus |-- †Magnosaurus |-- †Streptospondylus `--+-- †Afrovenator |-- †Dubreuillosaurus `--+-- †Torvosaurus `-- †Megalosaurus reference(s):
|
After Benson et al., 2010<==o Theropoda |-- †“Syntarsus” kayentakatae `--+-- †Dilophosaurus `--+--+-- †Ceratosaurus | `-- †Abelisauridae `--o Tetanurae |--+-- †Xuanhanosaurus | `--+-- †Marshosaurus | `--+-- †Piatnitzkysaurus | `-- †Condoraptor `--+--+-- †Monolophosaurus | `--+--o †Spinosauridae | | |--+-- †Baryonyx | | | `-- †Suchomimus | | `--+-- †Irritator | | `-- †Spinosaurus | `--+--+-- †Eustreptospondylus | | `--+-- †Streptospondylus | | `-- †Magnosaurus | `--+-- †Duriavenator | `--+--+-- †Torvosaurus | | `-- †Megalosaurus | `--+-- †Afrovenator | `-- †Dubreuillosaurus `--+-- Coelurosauria `--o †Allosauroidea |--+--+-- †Lourinhanosaurus | | `-- †Poekilopleuron | `--+-- †Metriacanthosaurus | `-- †Sinraptor `--+-- †Allosaurus `--+--o †Neovenatoridae | |-- †Neovenator | `--o †Megaraptora | |-- †Chilantaisaurus | |-- †Orkoraptos | |-- †Fukuiraptor | |-- †Aerosteon | |-- †Megaraptor | `-- †Australovenator `--+-- †Eocarcharia `--+-- †Acrocanthosaurus `--+-- †Shaochilong `--+-- †Tyrannotitan `--+-- †Carcharodontosaurus |-- †Giganotosaurus `-- †Mapusaurus Reference:
|
After Smith et al., 2007<==o Theropoda |-- †Coelophysoidea `--+-- †Zupaysaurus `--+-- “dilophosaurs” `--+-- †Neoceratosauria `--o Tetanurae |--+-- †Piatnitzkysaurus | `-- †Condoraptor `--+--+--+-- †Afrovenator | | `-- †Dubreuillosaurus | `--+-- †Torvosaurus | `--+--+-- †Eustreptospondylus | | `-- †Streptospondylus | `--+-- †Baryonyx | |-- †Irritator | `-- †Suchomimus `--+-- †Monolophosaurus `--+-- Coelurosauria `--o †Allosauroidea |--+-- †Acrocanthosaurus | `--+-- †Neovenator | `-- †Allosaurus `--+-- †Sinraptor `--+-- †Tyrannotitan `--+-- †Carcharodontosaurus |-- †Giganotosaurus `-- †Mapusaurus |
After Xu et al., 2009<==o Theropoda |-- †Coelophysoidea `--+-- †Zupaysaurus `--+-- “dilophosaurs” `--+-- †Ceratosauria `--o Tetanurae |--+-- †Piatnitzkysaurus | `-- †Condoraptor `--+--+--+-- †Afrovenator | | `-- †Dubreuillosaurus [= †Poekilopleuron? valesdunensis] | `--+-- †Torvosaurus | `--+-- †Megaraptor | |-- †Streptospondylus | |-- †Eustreptospondylus | |-- NMV P186076 | |-- †Baryonyx | |-- †Irritator | `-- †Suchomimus `--+-- †Monolophosaurus `--+-- Coelurosauria `--o †Allosauroidea |-- †Allosaurus |-- †Neovenator `--+-- †Acrocanthosaurus |-- †Sinraptor `--+-- †Tyrannotitan |-- †Carcharodontosaurus `--+-- †Giganotosaurus `-- †Mapusaurus |
After Holtz et al., 2004<==o Theropoda |-- †Ceratosauria `--o Tetanurae |-- †Szechuanosaurus? zigongensis `--+-- †Xuanhanosaurus `--+--+--o †Spinosauridae | | |-- †Baryonychidae | | `--+-- †Irritator | | `-- †Spinosaurus | `--+--+-- †Torvosaurus | | `--+-- †Megalosaurus | | `-- †Poekilopleuron | `--+-- †Piatnitzkysaurus | `--+-- †Eustreptospondylus | `--+-- †Poekilopleuron? valesdunensis | `-- †Afrovenator `--+-- Coelurosauria `--o †Allosauroidea |-- †Lourinhanosaurus |-- †Monolophosaurus |-- †Allosaurus |-- †Sinraptoridae |--+-- †Siamotyrannus | `-- †Fukuiraptor `--o †Carcharodontosauridae |-- †Neovenator `--+-- †Acrocanthosaurus `--+-- †Giganotosaurus `-- †Carcharodontosaurus |
After Rauhut, 2003<==o Theropoda |-- †Coelophysoidea `--+-- †Dilophosaurus `--+-- †Ceratosauria `--+-- †Szechuanosaurus? zigongensis |-- †Piatnitzkysaurus `--+-- Coelurosauria `--o †Allosauroidea |-- †Magnosaurus |-- †Poekilopleuron |-- †Monolophosaurus |--+-- †Torvosaurus | `--+-- †Chilantaisaurus | `-- †Baryonychidae `--+-- †Afrovenator |-- †Allosaurus `--+-- †Sinraptoridae `--+-- †Neovenator `-- †Carcharodontosauridae |
After Allain, 2002<==o Theropoda |-- †Herrerasaurus `--+-- †Ceratosauria `--o Tetanurae |--+--+-- †Cristatusaurus | | `-- †Baryonyx | `--+-- †Torvosaurus | `--+-- †Afrovenator | `--+-- †Eustreptospondylus | `-- †Poekilopleuron? valesdunensis `--+-- Coelurosauria `--o †Allosauroidea |--+-- †Allosaurus | `-- †Acrocanthosaurus `--+-- †Sinraptor `--+-- †Giganotosaurus `-- †Carcharodontosaurus reference:
|
After Holtz, 2000 (1998)<==o Theropoda |-- †Ceratosauria `--o Tetanurae |-- †Spinosauridae `--+-- †Megalosaurus `--+-- †Eustreptospondylus `--+-- †Torvosaurus `--+-- †Piatnitzkysaurus `--+-- †Afrovenator `--+-- Coelurosauria `--o †Allosauroidea |-- †Monolophosaurus `--+--+-- †Yangchuanosaurus | `-- †Sinraptor `--+--+-- †Neovenator | `-- †Allosaurus `--+-- †Acrocanthosaurus `--+-- †Giganotosaurus `-- †Carcharodontosaurus Reference:
|
After Sereno et al., 1998<==o Theropoda |-- †Ceratosauria `--+-- Neotetanurae `--o †Torvosauroidea |--o †Torvosauridae | |-- †Eustreptospondylus | `-- †Torvosaurus `--o †Spinosauridae |--+-- †Baryonyx | `-- †Suchomimus `--+-- †Irritator `-- †Spinosaurus Reference:
|
After Charig & Milner, 1997<==o Theropoda |-- †Ceratosauria `--+-- †Torvosaurus `--+-- †Megalosaurus `--+-- †Baryonyx `--+-- Coelurosauria `--o †Allosauroidea |-- †Allosaurus `-- †Acrocanthosaurus |
After Sereno et al., 1996<==o Theropoda |-- †Ceratosauria `--+--o †Megalosauroidea | |-- †Afrovenator | `--+-- †Torvosauridae | `-- †Spinosauridae `--+-- Coelurosauria `--o Allosauroidea |-- †Sinraptoridae |-- †Cryolophosaurus |-- †Monolophosaurus |-- †Allosaurus `--+-- †Giganotosaurus `--+-- †Acrocanthosaurus `-- †Carcharodontosaurus Reference:
|
After Coria & Salgado, 1995<==o Theropoda |-- †Herrerasauridae `--+-- †Ceratosauria `--+-- †Torvosauroidea `--+-- †Giganotosaurus `--+-- †Allosauroidea `-- Coelurosauria |
After Sereno et al., 1994<==o Theropoda |-- †Eoraptor `--+-- †Herrerasaurus `--+-- †Ceratosauria `--+--+-- †Afrovenator | `--+-- †Torvosauridae | `-- †Spinosauridae `--+-- Coelurosauria `--+-- †Sinraptoridae `-- †Allosauridae |
After Holtz, 1994<==o Theropoda |-- †Ceratosauria `--+-- †Torvosaurus `--+-- †Megalosaurus `--+-- Coelurosauria `--+-- †Allosaurus `-- †Acrocanthosaurus |
After Russell & Dong, 1993<==o Theropoda |-- †Coelophysis `--+--+-- †Ornithomimidae | `--+-- †Therizinosauridae | `--+-- †Troodontidae | `-- †Oviraptorosauria `--+-- †Yangchuanosaurus `--+-- †Baryonyx `--+-- †Allosaurus `--+-- †Dromaeosauridae `-- †Tyrannosauridae |
After Novas, 1992<==o Theropoda |-- Ceratosauria `--+-- †Eustreptospondylus `--+-- †Pianitzkysaurus `--+-- †Compsognathus `--+-- †Carnosauria `-- Coelurosauria |
After Paul, 1988<==o Theropoda |--+-- †Coelophysidae | `--+-- †Baryonyx | `-- †Spinosaurus `--+-- †Ceratosauridae `--+--+-- †Megalosaurus | `-- †Abelisauridae `--+-- †Eustreptospondylus `--+-- †Pianitzkysaurus `--+-- †Gasosaurus `--+-- †Marshoaurus `--+-- †Metriacanthosaurus `--+-- †Compsognatha `--+-- Protoavia `--o †Allosauria |-- †Coeluridae `--o †Allosauridae |-- †Ornitholestinae `==o †Allosaurinae |-- †Allosaurus `--+-- †Chilantaisaurus `--+-- †Acrocanthosaurus `-- †Tyrannosauridae |
After Paul, 1984<==o Theropoda |-- †Megalosaurus / †Torvosaurus / †Poekilopleuron |-- †Procompsognathus `--+-- †Coelophysidae `--+-- †Ceratosaurus `--+-- †Eustreptospondylus `--+-- †Yangchuanosaurus `--+--o Allosauridae | |-- †Allosaurus | `-- †Ornitholestinae `-- †Tyrannosaurs & †Dromaeosaurs |
Reference(s):
- Allain, R., 2001: Redescription de Streptospondylus altdorfensis, le dinosaure théropode de Cuvier, du Jurassique de Normandie.
[Redescription of Streptospondylus altdorfensis, Cuvier's theropod from the Jurassic of Normandy]
–Geodiversitas: Vol. 23, #3, pp. 349-367 - Allain, R., 2002: Discovery of megalosaur (Dinosauria, Theropoda) in the middle Bathonian of Normandy (France) and its implications for the phylogeny of basal Tetanurae.
–Journal of Vertebrate Paleontology: Vol. 22, #3, pp. 548-563 - Allain, R., 2005: The postcranial anatomy of the megalosaur Dubreuillosaurus valesdunensis (Dinosauria Theropoda) from the Middle Jurassic of Normandy, France.
–Journal of Vertebrate Paleontology: Vol. 25, #4, pp. 850-858 - Allain, R. & Chure, D. J., 2002: Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy.
–Palaeontology: Vol. 45, #6, pp. 1107-1121 - Benton, M. J., 1990: Origin and interrelationships of Dinosaurs. 11-30
in Weishampel, D. B., Dodson, P., & Osmólska, H. (eds.), 1990: The Dinosauria.
–University of California Press, Berkley, Los Angeles, Oxford, 1990 xvi-733 - Benson, R., Carrano, M. & Brusatte, S., 2010: A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic.
–Naturwissenschaften: Vol. 97, #1, pp. 71-78 [doi: 10.1007/s00114-009-0614-x] - Benson, R. B. J. & Radley, J. D., 2010: A new large-bodied theropod dinosaur from the Middle Jurassic of Warwickshire, United Kingdom.
–Acta Palaeontologica Polonica: Vol. 55, #1, pp. 35-42 [doi: 10.4202/app.2009.0083] - Buffetaut, E., 2011: An early spinosaurid dinosaur from the Late Jurassic of tendaguru (Tanzania) and the evolution of the spinosaurid dentition.
–Oryctos: Vol. 10, pp. 1-8 - Carrano, M. T., Benson, R. B. J. & Sampson, S. D., 2012: The phylogeny of Tetanurae (Dinosauria: Theropoda).
–Journal of Systematic Palaeontology: Vol. 10, #2, pp. 211-300 [doi: 10.1080/14772019.2011.630927] - Charig, A. J. & Milner, A. C., 1986: Baryonyx, a remarkable new theropod dinosaur.
–Nature: Vol. 324, 27 November, pp. 359-361 - Hendrickx, C. & Mateus, O., 2014: Torvosaurus gurneyi n. sp., the Largest Terrestrial Predator from Europe, and a Proposed Terminology of the Maxilla Anatomy in Nonavian Theropods.
–Plos One: Vol. 9, #3, pp. e88905 (1-25) [doi: 10.1371/journal.pone.0088905] - Holtz, T. R. Jr., 1998 [preview]: Spinosaurs as crocodile mimics.
–Science: Vol. 282, 13 November, pp. 1276-1277 - Holtz, T. R. Jr., 2000 (1998): A new phylogeny of the carnivorous dinosaurs.
–Gaia: Vol. 15, #1, pp. 5-61 - Keesey, T. M., 1996-2000: Tetanurae – Three-fingered meat-eaters
–iNet: The Dinosauricon: http://dinosauricon.com/taxa/tetanurae.html - Novas, F. E. 1996: Dinosaur monophyly.
–Journal of Vertebrate Paleontology: vol. 16, #4, pp. 723-741. - Novas, F. E., Salgado, L., Suarez, M., Agnolin, F. L., Ezcurra, M. D., Chimento, N. R., de la Cruz, R., Isasi, M. P., Vargas, A. O. & Rubilar-Rogers, D., 2015: An enigmatic plant-eating theropod from the Late Jurassic period of Chile.
–Nature: in press [doi: 10.1038/nature14307] - Molnar, R., E., 1990: Problematic Theropoda: “Carnosaurs”. 306-317
in Weishampel, D. B., Dodson, P., & Osmólska, H. (eds.), 1990: The Dinosauria.
–University of California Press, Berkley, Los Angeles, Oxford, 1990 xvi-733 - Molnar, R. E., Kurzanov, S. M., & Dong, Z., 1990: Carnosauria. 169-209
in Weishampel, D. B., Dodson, P., & Osmólska, H. (eds.), 1990: The Dinosauria.
–University of California Press, Berkley, Los Angeles, Oxford, 1990 xvi-733 - Padian, K., Hutchinson, J. R. & Holtz, T. R. Jr., 1999: Phylogenetic defenitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda).
–Journal of Vertebrate Paleontology: Vol. 19, #1, pp. 69-80 - Paul, G. S. 1988: The Predatory Dinosaurs of the World.
–Simon and Schuster Co., New York, 1988. 464. - Rauhut, O. W. M., 2005: Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia.
–Palaeontology: Vol. 48, #1, pp. 87-111 - Rauhut, O. W. M., Foth, C., Tischlinger, H. & Norell, M. A., 2012: Exceptionally preserved juvenile megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic of Germany.
–Proceedings of the National Academy of Sciences: Vol. 109, #29, pp. 11746-11751 [doi: 10.1073/pnas.1203238109] - Dal Sasso, C., Maganuco, S., Buffetaut, E. & Mendez, M. A., 2005: New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its size and affinities.
–Journal of Vertebrate Paleontology: Vol. 25, #4, pp. 888-896 - Sereno, P. C., Dutheil, D. B., Iarochene, M., Larsson, H. C. E., Lyon, G. H., Magwene, P. M., Sidor, C. A., Varricchio, D. J. & Wilson, J. A., 1996: Predatory dinosaurs from the Sahara and Late Cretaceous faunal differation.
–Science: Vol. 272, pp. 986-991 - Sereno, P. C., Beck, A. L., Dutheil, D. B., Gabo, B., Larsson, H. C. E., Lyon, G. H., Marcot, J. D., Rauhut, O. W. M., Sadleir, R. W., Sidor, C. A., Verricchio, D. J., Wilson, G. P. & Wilson, J. A., 1998: A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids.
–Science: Vol. 282, 13 November, pp. 1298-1302 - Sues, H.-D., Frey, E., Martill, D. M. & Scott, D. M., 2002: Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil.
–Journal of Vertebrate Paleontology: Vol. 22, #3, pp. 535-547