Carcharodontosauridae

Carcharodontosaurids
Temporal range: Late Jurassic - early Late Cretaceous 154–90 Ma
Reconstructed Carcharodontosaurus skull, Science Museum of Minnesota
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: †Carcharodontosauria
Family: †Carcharodontosauridae
Stromer, 1931
Subgroups
Synonyms
  • Acrocanthosauridae Molnar, 2003

Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, carcharodontósauros: "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae as a family, which, in modern paleontology, indicates a clade within Carnosauria. Carcharodontosaurids include some of the largest land predators ever known: Giganotosaurus, Mapusaurus, Carcharodontosaurus, and Tyrannotitan all rivaled Tyrannosaurus in size. Estimates give a maximum weight of 8–10 metric tons (8.8–11.0 short tons) for the largest carcharodontosaurids, while the smallest carcharodontosaurids were estimated to have weighed at least 500 kilograms (1,100 lb).

Discovery and history

[edit]
Lost original teeth of Carcharodontosaurus (Megalosaurus) saharicus (Fig. 1-2), the lectotype being at the top (Fig. 1)

The earliest discovery of carcharodontosaurid fossils may date to 1835 with the discovery of Poekilopleuron in Jurassic-aged sediments in Normandy, France and it was then described in 1836 by French naturalist Jacques Amand Eudes-Deslongchamps.[8][9] However, the holotype (name-bearing) specimen, which consisted of an incomplete postcranial skeleton, was destroyed during World War II.[10] Due to the fragmentary and destroyed nature of Poekilopleuron's fossils, little can be ascertained about its phylogenetic position or validity. However, recent phylogenetic analyses have recovered it as a basal carcharodontosaurid or carcharodontosaurian,[11][12] though it has also been classified as a metriacanthosaurid,[13] megalosauroid,[14][15] or another position in Carnosauria.[16][17] If not Poekilopleuron, the oldest discovery of carcharodontosaurid fossils would be between 1909 and 1913 with the discovery of Veterupristisaurus in the Jurassic-aged Tendaguru Formation of Lindi, Tanzania. These fossils, consisting of several caudal (tail) vertebrae, were described by German paleontologist Werner Janensch in 1925 as belonging to an indeterminate large theropod.[18][19] However after further study in 2011, German paleontologist Oliver Rauhut concluded that these vertebrae belonged to a novel genus and species of carcharodontosaurid which he named Veterupristisaurus milneri.[19]

Fossils of Carcharodontosaurus itself were discovered in 1924 in several Cretaceous-aged[20] deposits of the Continental Intercalaire in Timimoun, Algeria.[21] These fossils, consisting of only two teeth, were then described by French geologists Charles Depéret and Justin Savornin in 1925 as belonging to a new species of theropod dinosaur, Megalosaurus saharicus,[22] though then Megalosaurus was a wastebasket taxon.[23] It was the first theropod described from North Africa and the first confirmed carcharodontosaurid to be named. Both teeth have since been lost and are now thought to be undiagnostic,[24] however many other teeth and bones were referred to the species from Cretaceous-aged sediments across North Africa.[25][24] A partial skeleton of a carcharodontosaurid was unearthed in marls near Ain Gedid, Egypt in 1914 by Austro-Hungarian paleontologist Richard Markgraf and deposited in the collections of the Paläontologisches Museum München in 1922.[26][27][28] This incomplete skeleton was described by German paleontologist Ernst Stromer in 1931, who recognized the uniqueness of the teeth and skeleton leading to him to erect a new genus name for M. saharicus, Carcharodontosaurus. This name was for their similarities, in sharpness and serrations, to the teeth of the great white shark (Carcharodon carcharias).[26] Carcharodontosaurus is the name-bearing genus of the family Carcharodontosauridae, whose name means "shark-toothed lizards". In describing this partial skeleton, Stromer designated the smaller of the two teeth described by Deperet and Savornin as the lectotype of C. saharicus.[29][26] However, this partial skeleton was later destroyed in World War II[28][30] and was recognized as a new genus and species of carcharodontosaurid, Tameryraptor markgrafi, in 2025.[28]

Partial holotype skull (white parts are reconstructed) and teeth of Giganotosaurus, Ernesto Bachmann Paleontological Museum

Following the description of Carcharodontosaurus, few additional carcharodontosaurid discoveries were made or recognized as being carcharodontosaurid.[27] Fossils of carcharodontosaurids, such as those of Acrocanthosaurus, were often described as belonging to another family in Carnosauria or Theropoda,[31] or described as species of other genera, such as Shaochilong.[32][33] However, in the 1990s and 2000s, resurgent interest in the Cretaceous fauna of Gondwana led to the discovery of many additional carcharodontosaurids. Although isolated carcharodontosaurid fossils had been known from South America since the 1960s with the description of "Megalosaurus chubutensis",[34][35] the first major find came with the discovery of Giganotosaurus in 1993.[36][37] Giganotosaurus was found in Cretaceous-aged rocks in Neuquén, Argentina and named in 1995 by Argentine paleontologists Rodolfo Coria and Leonardo Salgado. This specimen of Giganotosaurus consisted of around 70% of the skeleton, including much of the vertebral column, skull, and limbs.[38][39][40] However, Coria and Salgado did not recognize the close relationship of Giganotosaurus to Carcharodontosaurus and Acrocanthosaurus.[40] Also in 1995, an incomplete skull of Carcharodontosaurus was unearthed from layers of the Lower Douira Formation, Kem Kem Beds, in Errachidia, southeastern Morocco.[27][24] This skull was then described by American paleontologist Paul Sereno and colleagues in 1996, who recognized that Giganotosaurus and Acrocanthosaurus were both members of Carcharodontosauridae, together forming a transcontinental clade.[27] These discoveries led to the modern understanding of the family, though further research and discoveries has led to the addition of more members to the clade from other continents.

Evolution

[edit]

Along with the spinosaurids, carcharodontosaurids were the largest predators in the early and middle Cretaceous throughout Gondwana, with species also present in North America (Acrocanthosaurus), Europe (Concavenator) and possibly Asia (Shaochilong).[41] Although not classified within the family Carcharodontosauridae, indeterminate remains of carcharodontosaurians are also found in Early Cretaceous deposits of Australia.[42] Carcharodontosaurids range throughout the Cretaceous from the Barremian (127-121 million years ago) to the Turonian (94-90 million years ago). Past the Turonian, they were replaced by the smaller abelisaurids in Gondwana and by tyrannosaurids in North America and Asia.[43][44] While some teeth and a maxilla discovered in Maastrichtian deposits of Brazil.[45][46] this identification has been subsequently rejected and the material assigned to abelisaurids after better examination,[47] and there are no reliable records of carcharodontosaurs in South America beyond the end of the Turonian.[43] In December 2011, Oliver W. M. Rauhut described a new genus and species of carcharodontosaurid from the Late Jurassic (late Kimmeridgian to earliest Tithonian faunal stage, about 154-150 million years ago) of Tendaguru Formation, southeastern Tanzania. This genus, Veterupristisaurus represents the oldest known carcharodontosaurid.[19]

Size comparison of seven carcharodontosaurids, though the inclusion of Shaochilong as a member of the family is under contention.

Classification

[edit]
Top to bottom: Acrocanthosaurus, Carcharodontosaurus, Meraxes, Giganotosaurus

The family Carcharodontosauridae was originally named by Ernst Stromer in 1931 to include the single newly discovered species Carcharodontosaurus saharicus. A close relative of C. saharicus, Giganotosaurus, was added to the family when it was described in 1995. Additionally, many paleontologists have included Acrocanthosaurus in this family (Sereno et al. 1996, Harris 1998, Holtz 2000, Rauhut 2003, Eddy & Clarke, 2011, Rauhut 2011), though others place it in the related family Allosauridae (Currie & Carpenter, 2000; Coria & Currie, 2002). Carcharodontosaurids are characterized by the following morphological characteristics : Dorsoventral depth of anterior maxillary interdental plates more than twice anteroposterior width, squared, sub-rectangular anterior portion of the dentary, teeth with wrinkled enamel surfaces, presence of four premaxillary alveoli and a premaxillary body taller than long in lateral aspect, opisthocoelous cervical vertebrae with neural spines more than 1.9 times the height of the centrum, large, textured rugosities on the lacrimal and postorbital formed by roofing and forming broad orbital shelves, and a proximomedially inclined femoral head.[48][49] With the discovery of Mapusaurus in 2006, Rodolfo Coria and Phil Currie erected a subfamily of Carcharodontosauridae, the Giganotosaurinae, to contain the most advanced South American species, which they found to be more closely related to each other than to the African and European forms. Coria and Currie did not formally refer Tyrannotitan to this subfamily, pending a more detailed description of that genus, but noted that based on characteristics of the femur, it may be a gigantosaurine as well.[50]

Restoration of the crushed skull of Concavenator

In 1998 Paul Sereno defined Carcharodontosauridae as a clade, consisting of Carcharodontosaurus and all species closer to it than to either Allosaurus, Sinraptor, Monolophosaurus, or Cryolophosaurus. Therefore, this clade is by definition outside of the clade Allosauridae.

The cladogram below follows the analyses by Canale et al. (2022) from their description of the large carcharodontosaurine Meraxes.[3]

Carcharodontosauridae

In his 2024 review of theropod relationships, Cau recovered the following results for the Carcharodontosauridae.[51]

Carcharodontosauridae
Neovenator

Carcharodontosaurus iguidensis (holotype maxilla)

Acrocanthosaurus

Eocarcharia (referred maxilla)

Meraxes

Carcharodontosaurus iguidensis (referred cranial material)

Carcharodontosaurus saharicus (neotype)

Carcharodontosaurus saharicus (described by Stromer in 1931)

While describing Tameryraptor in their phylogenetic analyses Kellermann, Cuesta & Rauhut (2025) recovered the following results. They recovered Lusovenator and Veterupristisaurus as Late Jurassic carcharodontosaurids. Taurovenator and Kelmayisaurus were recovered variably within or outside Carcharodontosauridae. Siamraptor and Concavenator were recovered as sister taxa outside of Carcharodontosauridae but still basal members of Carcharodontosauria in all analyses. Datanglong was consistently recovered as an indeterminate carcharodontosaurid along with Veterupristisaurus. Neovenator, Eocarcharia, and Poekilopleuron were recovered outside of Carcharodontosauridae in all analyses, with Neovenator consistently recovered within Carcharodontosauria and sometimes as the sister taxon of Chilantaisaurus. Shaochilong was recovered as a tyrannosauroid rather than a carcharodontosaurid.[28]

Mounted Acrocanthosaurus skeleton (NCSM 14345) at the North Carolina Museum of Natural Sciences.
Carcharodontosauridae

Paleobiology

[edit]

Growth

[edit]

Osteohistological analysis of the Meraxes holotype specimen suggests the individual could have been between 39 and 53 years old when it died, having reached skeletal maturity approximately 4 years prior to its death (between 35 and 49 years old), making it the longest-lived non-avian theropod currently known. Meraxes was determined to have grown to large size by extending its growth period (hypermorphosis), rather than increasing its relative growth rate (acceleration) through development as in Tyrannosaurus, to which it was compared.[3][52]

References

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  1. ^ Stephen L. Brusatte; Roger B. J. Benson; Xing Xu (2012). "A reassessment of Kelmayisaurus petrolicus, a large theropod dinosaur from the Early Cretaceous of China" (PDF). Acta Palaeontologica Polonica. 57 (1): 65–72. doi:10.4202/app.2010.0125. S2CID 53387460. Archived from the original (PDF) on 2011-12-18. Retrieved 2011-04-28.
  2. ^ Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (2025-01-14). "Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny". PLOS One. 20 (1): e0311096. Bibcode:2025PLoSO..2011096K. doi:10.1371/journal.pone.0311096. ISSN 1932-6203. PMC 11731741. PMID 39808629.
  3. ^ a b c d Canale, J.I.; Apesteguía, S.; Gallina, P.A.; Mitchell, J.; Smith, N.D.; Cullen, T.M.; Shinya, A.; Haluza, A.; Gianechini, F.A.; Makovicky, P.J. (2022-07-07). "New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod arm reduction". Current Biology. 32 (14): 3195–3202.e5. Bibcode:2022CBio...32E3195C. doi:10.1016/j.cub.2022.05.057. PMID 35803271.
  4. ^ Rauhut, Oliver W M; Bakirov, Aizek A; Wings, Oliver; Fernandes, Alexandra E; Hübner, Tom R (August 1, 2024). "A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan". Zoological Journal of the Linnean Society. 201 (4). doi:10.1093/zoolinnean/zlae090. ISSN 0024-4082.
  5. ^ a b Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (14 January 2025). "Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny". PLOS ONE. 20 (1): e0311096. Bibcode:2025PLoSO..2011096K. doi:10.1371/journal.pone.0311096. PMC 11731741. PMID 39808629.
  6. ^ Novas, Fernando E. (2013). "Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia". Cretaceous Research. 45: 174–215. Bibcode:2013CrRes..45..174N. doi:10.1016/j.cretres.2013.04.001. hdl:11336/102037.
  7. ^ Rolando, Alexis M. Aranciaga; Motta, Matías J.; Agnolín, Federico L.; Tsuihiji, Takanobu; Miner, Santiago; Brissón-Egli, Federico; Novas, Fernando E. (2024-10-09). "A new carcharodontosaurid specimen sheds light on the anatomy of South American giant predatory dinosaurs". The Science of Nature. 111 (6): 56. Bibcode:2024SciNa.111...56R. doi:10.1007/s00114-024-01942-4. ISSN 1432-1904. PMID 39382666.
  8. ^ Eudes-Deslongchamps J.-A. (1836). "Plusieurs parties d'un mémoire sur un très-grand animal fossile découvert, l'an dernier, dans les carrières de la Maladrerie à un quart de lieu de Caen". Analyse des Travaux de la Société Pendant l'Année Académique 1835–1836, Séance Publique de la Société Linnéenne de Normandie Tenue à Vire le 24 Mai 1836: 14–25.
  9. ^ Brignon, Arnaud (2018). "Nouvelles données historiques sur les premiers dinosaures trouvés en France". BSGF - Earth Sciences Bulletin (in French). 189 (1): 4. doi:10.1051/bsgf/2018003. ISSN 0037-9409.
  10. ^ Allain, Ronan; Chure, Daniel J. (2002). "Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy". Palaeontology. 45 (6): 1107–1121. Bibcode:2002Palgy..45.1107A. doi:10.1111/1475-4983.00277. ISSN 1475-4983.
  11. ^ Oliver W. M. Rauhut; Diego Pol (2019). "Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs". Scientific Reports. 9 (1): Article number 18826. Bibcode:2019NatSR...918826R. doi:10.1038/s41598-019-53672-7. PMC 6906444. PMID 31827108.
  12. ^ Rauhut, Oliver W M; Bakirov, Aizek A; Wings, Oliver; Fernandes, Alexandra E; Hübner, Tom R (August 1, 2024). "A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan". Zoological Journal of the Linnean Society. 201 (4). doi:10.1093/zoolinnean/zlae090. ISSN 0024-4082.
  13. ^ Cau, Andrea (2024). "A Unified Framework for Predatory Dinosaur Macroevolution" (PDF). Bollettino della Società Paleontologica Italiana. 63 (1): 1-19. doi:10.4435/BSPI.2024.08 (inactive 2024-11-20).{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)
  14. ^ Rauhut, Oliver W.M.; Hübner, Tom R.; Lanser, Klaus-Peter (2016). "A new megalosaurid theropod dinosaur from the late Middle Jurassic (Callovian) of north-western Germany: Implications for theropod evolution and faunal turnover in the Jurassic". Palaeontologia Electronica. 19 (2): 1–65. Retrieved 13 January 2022.
  15. ^ Hui Dai; Roger Benson; Xufeng Hu; Qingyu Ma; Chao Tan; Ning Li; Ming Xiao; Haiqian Hu; Yuxuan Zhou; Zhaoying Wei; Feng Zhang; Shan Jiang; Deliang Li; Guangzhao Peng; Yilun Yu; Xing Xu (2020). "A new possible megalosauroid theropod from the Middle Jurassic Xintiangou Formation of Chongqing, People's Republic of China and its implication for early tetanuran evolution". Scientific Reports. 10 (1): Article number 139. Bibcode:2020NatSR..10..139D. doi:10.1038/s41598-019-56959-x. PMC 6954265. PMID 31924836.
  16. ^ Allain R (2001). "Redescription de Streptospondylus altdorfensis, le dinosaure théropode de Cuvier, du Jurassique de Normandie [Redescription of Streptospondylus altdorfensis, Cuvier's theropod dinosaur from the Jurassic of Normandy]" (PDF). Geodiversitas. 23 (3): 349–367. Archived from the original (PDF) on 2012-06-02. Retrieved 2011-02-24.
  17. ^ Benson, R.B.J., Carrano, M.T and Brusatte, S.L. (2010). "A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic". Naturwissenschaften. 97 (1): 71–78. Bibcode:2010NW.....97...71B. doi:10.1007/s00114-009-0614-x. PMID 19826771. S2CID 22646156.{{cite journal}}: CS1 maint: multiple names: authors list (link) Supporting Information
  18. ^ W. Janensch. (1925). Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas [The coelurosaurs and theropods of the Tendaguru Formation of German East Africa]. Palaeontographica, Supplement VII. (1) 1(1):1-100
  19. ^ a b c Rauhut, Oliver W. M. (2011). "Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania)". Special Papers in Palaeontology. 86: 195–239. Archived from the original on 2020-02-01. Retrieved 2020-03-17.
  20. ^ Benyoucef, Madani; Pérez-García, Adán; Bendella, Mohamed; Ortega, Francisco; Vullo, Romain; Bouchemla, Imad; Ferré, Bruno (2022-07-06). "The "mid"-Cretaceous (Lower Cenomanian) Continental Vertebrates of Gara Samani, Algeria. Sedimentological Framework and Palaeodiversity". Frontiers in Earth Science. 10 (927059). Bibcode:2022FrEaS..10.7059B. doi:10.3389/feart.2022.927059. ISSN 2296-6463.
  21. ^ Depéret, Charles; Savornin, Justin (1927). "La faune de reptiles et de poisons albiens de Timimoun (Sahara algérien)". Bulletin de la Société Géologique de France. 27: 257–265.
  22. ^ Depéret, Charles; Savornin, Justin (1925). "Sur la découverte d'une faune de vertébrés albiens à Timimoun (Sahara occidental)". Comptes Rendus de l'Académie des Sciences. 181: 1108–1111.
  23. ^ Benson, Roger B. J.; Barrett, Paul M.; Powell, H. Philip; Norman, David B. (2008). "The Taxonomic Status of Megalosaurus Bucklandii (dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, Uk". Palaeontology. 51 (2): 419–424. Bibcode:2008Palgy..51..419B. doi:10.1111/j.1475-4983.2008.00751.x. ISSN 1475-4983.
  24. ^ a b c Brusatte, Stephen L.; Sereno, Paul C. (December 12, 2007). "A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus". Journal of Vertebrate Paleontology. 27 (4): 902–916. doi:10.1671/0272-4634(2007)27[902:ANSOCD]2.0.CO;2. S2CID 86202969.
  25. ^ de Lapparent, Albert (1960). "Les dinosauriens du "Continental intercalaire" du Sahara central" (PDF). Memoirs of the Geological Society of France. 88A: 1–57.
  26. ^ a b c Stromer, Ernst (1931). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîjestufe (unterstes Cenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov. gen" [Results of Prof. E. Stromer's research trips in the deserts of Egypt. II. Vertebrate remains from the Baharîje stage (lower Cenomanian). 10. A skeletal remains of Carcharodontosaurus nov. gen.] (PDF). Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung. Neue Folge (in German). 9. Translated by Carrano, Matthew: 1–31.
  27. ^ a b c d Sereno, Paul C.; Dutheil, Didier B.; Iarochene, M.; Larsson, Hans C. E.; Lyon, Gabrielle H.; Magwene, Paul M.; Sidor, Christian A.; Varricchio, David J.; Wilson, Jeffrey A. (1996). "Predatory Dinosaurs from the Sahara and Late Cretaceous Faunal Differentiation" (PDF). Science. 272 (5264): 986–991. Bibcode:1996Sci...272..986S. doi:10.1126/science.272.5264.986. PMID 8662584. S2CID 39658297. Archived (PDF) from the original on December 4, 2024. Retrieved March 20, 2024.
  28. ^ a b c d Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (2025-01-14). "Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny". PLOS ONE. 20 (1): e0311096. Bibcode:2025PLoSO..2011096K. doi:10.1371/journal.pone.0311096. ISSN 1932-6203. PMC 11731741. PMID 39808629.
  29. ^ Stromer 1934, p. 62.
  30. ^ Smith, Joshua B.; Lamanna, Matthew C.; Mayr, Helmut; Lacovara, Kenneth J. (2006). "New information regarding the holotype of Spinosaurus aegyptiacus Stromer, 1915". Journal of Paleontology. 80 (2): 400–406. doi:10.1666/0022-3360(2006)080[0400:NIRTHO]2.0.CO;2. S2CID 130989487.
  31. ^ Stovall, J. Willis; Langston, Wann (1950). "Acrocanthosaurus atokensis, a New Genus and Species of Lower Cretaceous Theropoda from Oklahoma". The American Midland Naturalist. 43 (3): 696–728. doi:10.2307/2421859. ISSN 0003-0031. JSTOR 2421859.
  32. ^ Brusatte, Stephen L.; Benson, Roger B. J.; Chure, Daniel J.; Xu, Xing; Sullivan, Corwin; Hone, David W. E. (2009-09-01). "The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids". Naturwissenschaften. 96 (9): 1051–1058. Bibcode:2009NW.....96.1051B. doi:10.1007/s00114-009-0565-2. ISSN 1432-1904.
  33. ^ Hu, S.-Y. (1964). "Carnosaurian remains from Alashan, Inner Mongolia." Vertebrata PalAsiatica, 8: 42–63. [In Chinese, with English summary]
  34. ^ Corro, G. del (1966). "Un nuevo dinosaurio Carnivoro del Chubut (Argentina)". Comunicaciones del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" e Instituto Nacional de Investigación de las Ciencias Naturales, Paleontología. 1: 1–4.
  35. ^ Ezcurra, Martin Daniel; Novas, Fernando Emilio (2016). "Theropod dinosaurs from Argentina". Contribuciones del MACN. 6: 139–156. ISSN 1666-5503.
  36. ^ Pons, M. "Ernesto Bachmann Dinosaurs Museum – El Chocón". Welcome Argentina. Archived from the original on August 8, 2016. Retrieved August 3, 2016.
  37. ^ Hajek, D. (2015). "Bankrolling A Dinosaur Dig And Unearthing A Giant: The Giganotosaurus". NPR.org. Retrieved August 3, 2016.
  38. ^ Coria, R. A.; Sagado, L. (1994). "A giant theropod from the middle Cretaceous of Patagonia, Argentina". Journal of Vertebrate Paleontology. 14 (3): 22A. doi:10.1080/02724634.1994.10011592. JSTOR 4523584.
  39. ^ Coria, Rodolfo A.; and Currie, Philip J. (2003-01-14). "The braincase of Giganotosaurus carolinii (Dinosauria: Theropoda) from the Upper Cretaceous of Argentina". Journal of Vertebrate Paleontology. 22 (4): 802–811. doi:10.1671/0272-4634(2002)022[0802:TBOGCD]2.0.CO;2. ISSN 0272-4634.
  40. ^ a b Coria, Rodolfo A.; Salgado, Leonardo (1995). "A new giant carnivorous dinosaur from the Cretaceous of Patagonia". Nature. 377 (6546): 224–226. Bibcode:1995Natur.377..224C. doi:10.1038/377224a0. ISSN 1476-4687.
  41. ^ Brusatte, S.; Benson, R.; Chure, D.; Xu, X.; Sullivan, C.; Hone, D. (2009). "The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids" (PDF). Naturwissenschaften. 96 (9): 1051–8. Bibcode:2009NW.....96.1051B. doi:10.1007/s00114-009-0565-2. hdl:20.500.11820/33528c2e-0c9c-4160-8693-984f077ee5d0. PMID 19488730. S2CID 25532873. Archived (PDF) from the original on 2021-08-27. Retrieved 2020-08-24.
  42. ^ Kotevski, Jake; Duncan, Ruairidh J.; Ziegler, Tim; Bevitt, Joseph J.; Vickers-Rich, Patricia; Rich, Thomas H.; Evans, Alistair R.; Poropat, Stephen F. (2025). "Evolutionary and paleobiogeographic implications of new carcharodontosaurian, megaraptorid, and unenlagiine theropod remains from the upper Lower Cretaceous of Victoria, southeast Australia". Journal of Vertebrate Paleontology. 0: e2441903. doi:10.1080/02724634.2024.2441903. ISSN 0272-4634.
  43. ^ a b Meso, J.G.; Juárez Valieri, R.D.; Porfiri, J.D.; Correa, S.A.S.; Martinelli, A.G.; Casal, G.A.; Canudo, J.I.; Poblete, F.; Dos Santos, D. (2021-09-01). "Testing the persistence of Carcharodontosauridae (Theropoda) in the Upper Cretaceous of Patagonia based on dental evidence". Cretaceous Research. 125: 104875. Bibcode:2021CrRes.12504875M. doi:10.1016/j.cretres.2021.104875. hdl:11336/183886. ISSN 0195-6671. Archived from the original on 2021-04-30. Retrieved 2021-04-30.
  44. ^ Novas, de Valais, Vickers-Rich, and Rich. (2005). "A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids." Naturwissenschaften,
  45. ^ Rodrigo P. Fernandes de Azevedo; Felipe Medeiros Simbras; Miguel Rodrigues Furtado; Carlos Roberto A. Candeiro & Lílian Paglarelli Bergqvist (2013). "First Brazilian carcharodontosaurid and other new theropod dinosaur fossils from the Campanian–Maastrichtian Presidente Prudente Formation, São Paulo State, southeastern Brazil". Cretaceous Research. 40: 131–142. Bibcode:2013CrRes..40..131F. doi:10.1016/j.cretres.2012.06.004.
  46. ^ Carlos Roberto Candeiro; Philip Currie; Lílian Bergqvist (2012). "Theropod teeth from the Marília Formation (late Maastrichtian) at the Paleontological Site of Peirópolis in Minas Gerais State, Brazil". Revista Brasileira de Geociências. 42 (2): 323–330. doi:10.5327/z0375-75362012000200008 (inactive 1 November 2024).{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)
  47. ^ Delcourt, R.; Grillo, O.N. (2017). "Carcharodontosaurids remained extinct in the Campanian-Maastrichtian: Reassessment of a fragmentary maxilla from Presidente Prudente Formation, Brazil". Cretaceous Research. 84: 515–524. doi:10.1016/j.cretres.2017.09.008.
  48. ^ Ortega, F.; Escaso, F.; Sanz, J. L. (2010). "A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain" (PDF). Nature. 467 (7312): 203–206. Bibcode:2010Natur.467..203O. doi:10.1038/nature09181. PMID 20829793. S2CID 4395795. Archived from the original (PDF) on 2018-10-08. Retrieved 2016-12-06.
  49. ^ Eddy, DR; Clarke, JA (2011). "New Information on the Cranial Anatomy of Acrocanthosaurus atokensis and Its Implications for the Phylogeny of Allosauroidea (Dinosauria: Theropoda)". PLOS ONE. 6 (3): e17932. Bibcode:2011PLoSO...617932E. doi:10.1371/journal.pone.0017932. PMC 3061882. PMID 21445312.
  50. ^ Coria, R.A.; Currie, P.J. (2006). "A new carcharodontosaurid (Dinosauria, Theropoda) from the Upper Cretaceous of Argentina". Geodiversitas. 28 (1): 71–118.
  51. ^ Cau, Andrea (2024). "A Unified Framework for Predatory Dinosaur Macroevolution" (PDF). Bollettino della Società Paleontologica Italiana. 63 (1): 1–19. doi:10.4435/BSPI.2024.08 (inactive 2024-11-20). Archived from the original (PDF) on 2024-04-27. Retrieved 2024-05-02.{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)
  52. ^ Cullen, Thomas M.; Canale, Juan I.; Apesteguía, Sebastián; Smith, Nathan D.; Hu, Dongyu; Makovicky, Peter J. (2020-11-25). "Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution". Proceedings of the Royal Society B: Biological Sciences. 287 (1939): 20202258. doi:10.1098/rspb.2020.2258. ISSN 0962-8452. PMC 7739506. PMID 33234083.
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