:Carcharodontosaurus

File:Lost holotype teeth of Carcharodontosaurus saharicus.png being at the top (Fig. 1)]]

In 1924, two teeth of Carcharodontosaurus were unearthed from wall cuts in different foggaras near Timimoun, French Algeria. These sediments came from the Cretaceous-aged{{Cite journal |last1=Benyoucef |first1=Madani |last2=Pérez-García |first2=Adán |last3=Bendella |first3=Mohamed |last4=Ortega |first4=Francisco |last5=Vullo |first5=Romain |last6=Bouchemla |first6=Imad |last7=Ferré |first7=Bruno |date=2022 |title=The 'mid'-Cretaceous (Lower Cenomanian) Continental Vertebrates of Gara Samani, Algeria. Sedimentological Framework and Palaeodiversity |journal=Frontiers in Earth Science |volume=10 |page=927059 |doi=10.3389/feart.2022.927059 |bibcode=2022FrEaS..10.7059B |doi-access=free }} Continental intercalaire Formation.{{Cite journal |last1=Depéret |first1=Charles |last2=Savornin |first2=Justin |date=1927 |title=La faune de reptiles et de poisons albiens de Timimoun (Sahara algérien) |journal=Bulletin de la Société Géologique de France |volume=27 |pages=257–265 |url=https://patrimoine.sorbonne-universite.fr/viewer/2708/?offset=#page=293&viewer=picture&o=&n=0&q=}} The fossils were taken to the governor of Timimoun, Captain Burté, who gave them to French geologist Charles Depéret later that year. In 1925, Depéret and his colleague Justin Savornin described the teeth as coming of a new species of theropod dinosaur, Megalosaurus saharicus. These were the first fossils of theropods to be described from the region. The specific name saharicus refers to the Sahara Desert where the teeth had been found.{{Cite journal |last1=Depéret |first1=Charles |last2=Savornin |first2=Justin |date=1925 |title=Sur la découverte d'une faune de vertébrés albiens à Timimoun (Sahara occidental) |url=https://gallica.bnf.fr/ark:/12148/bpt6k3134w/f1108.item.r=saharicus |journal=Comptes Rendus de l'Académie des Sciences |volume=181 |pages=1108–1111}} The genus Megalosaurus was a wastebasket taxon, with many new species referred to it without justification, including M. saharicus.{{Cite journal |last1=Benson |first1=Roger B. J. |last2=Barrett |first2=Paul M. |last3=Powell |first3=H. Philip |last4=Norman |first4=David B. |date=2008 |title=The taxonomic status of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, UK |journal=Palaeontology |volume=51 |issue=2 |pages=419–424 |doi=10.1111/j.1475-4983.2008.00751.x |bibcode=2008Palgy..51..419B |s2cid=83324840 |doi-access=free }} It was later considered to be a species of Dryptosaurus in 1927, though this is unjustified.{{Cite book|last=von Huene|first=Friedrich|year=1956|title=Palaeontologie und Phylogenie der Niederen Tetrapoden|publisher=VEB Gustav Fischer Verlang|volume=1|place=Jena|pages=716|oclc=489883421}}{{Cite journal |last1=Sereno |first1=Paul C. |last2=Dutheil |first2=Didier B. |last3=Iarochene |first3=M. |last4=Larsson |first4=Hans C. E. |last5=Lyon |first5=Gabrielle H. |last6=Magwene |first6=Paul M. |last7=Sidor |first7=Christian A. |last8=Varricchio |first8=David J. |last9=Wilson |first9=Jeffrey A. |year=1996 |title=Predatory Dinosaurs from the Sahara and Late Cretaceous Faunal Differentiation |journal=Science |volume=272 |issue=5264 |pages=986–991 |doi=10.1126/science.272.5264.986 |pmid=8662584 |bibcode=1996Sci...272..986S |s2cid=39658297 |url=http://doc.rero.ch/record/13893/files/PAL_E831.pdf |archive-date=December 4, 2024 |access-date=March 20, 2024 |archive-url=https://web.archive.org/web/20241204204501/https://doc.rero.ch/record/13893/files/PAL_E831.pdf |url-status=live }} By accident, another species of Megalosaurus, M. africanus, was named by German paleontologist Friedrich von Huene based on the teeth. It is therefore considered a junior synonym of M. saharicus. Both syntypic teeth of M. saharicus have since been lost, possibly being kept in a collection in Algeria, Paris, or Lyon, and lack distinguishing characteristics from other carcharodontosaurids.{{Cite journal |last1=Brusatte |first1=Stephen L. |last2=Sereno |first2=Paul C. |date=December 12, 2007 |title=A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus |journal=Journal of Vertebrate Paleontology |volume=27 |issue=4 |pages=902–916 |doi=10.1671/0272-4634(2007)27[902:ANSOCD]2.0.CO;2 |s2cid=86202969 }} In 1960, French paleontologist Albert-Félix de Lapparent reported the discovery of more teeth and several caudal vertebrae from sites in Algeria belonging to Carcharodontosaurus,{{Cite journal |last=de Lapparent |first=Albert |date=1960 |title=Les dinosauriens du "Continental intercalaire" du Sahara central. |url=https://naturalhistory.si.edu/sites/default/files/media/translated_publications/Lapparent_60.pdf |journal=Memoirs of the Geological Society of France |volume=88A |pages=1–57}} though some of these fossils might belong to other genera. Later authors mentioned finds of teeth and isolated fossils from other provinces of Algeria.{{Cite journal|author1=Gabani, A. |author2=Mammeri, C. |author3=Adaci, M. |author4=Bensalah, M. |author5=Mahboubi, M. |date=2016 |title=Le Crétacé continental à vertébrés de la bordure sud du Plateau de Tinhert: considérations stratigraphiques et bilan paléontologique |journal=Mémoire du Service Géologique de l'Algérie |volume=19 |pages=39–61}}

File:Tameryraptor_(holotype,_SNSB-BSPG_1922_X_46).png in 1931, now the holotype of Tameryraptor]]

However, a partial skeleton later referred to C. saharicus was first found in marls near Ain Gedid, Egypt, in early April 1914 by Austro-Hungarian paleontologist Richard Markgraf. Marls from this region derive from the Cenomanian-aged Bahariya Formation, one of many Cretaceous-aged sites of North Africa.{{Cite journal |last1=Stromer |first1=Ernst |date=1931 |title=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. |trans-title=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. |journal=Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung |series=Neue Folge |language=de |volume=9 |pages=1–31|url=https://www.dinochecker.com/papers/Stromers-Egypt-expedition_Carcharodontosaurus_Stromer_1931.pdf|translator-last1=Carrano|translator-first1=Matthew}}{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=162}} In this formation, Markgraf did extensive collecting of dinosaur skeletons for his employer, German paleontologist Ernst Stromer of the Paläontologisches Museum München (Bavarian State Collection of Paleontology). Due to political tensions between the German Empire and then British-owned Egypt, this skeleton, since numbered as SNSB-BSPG 1922 X 46, took years to get to Germany. It was not until 1922 that they were transported overseas to Munich where they were described by Stromer in 1931.{{sfn|Nothdurft|Smith|2002|p=107–108}} Stromer recognized that the skeleton's teeth would match the characteristic dentition of those described by Depéret and Savornin, which led to Stromer conserving the species name saharicus. However, he found it necessary to erect a new genus for this species, Carcharodontosaurus, for their similarities, in sharpness and serrations, to the teeth of the great white shark (Carcharodon carcharias). In his 1931 and 1934 descriptions, Stromer designates the smaller of the two teeth originally described by Depéret and Savornin as the type specimen (name-bearing specimen) of the taxon.{{sfn|Stromer|1934|p=62}} Thus, this tooth, although lost, must be considered as the lectotype of C. saharicus. World War II would break out in 1939, leading SNSB-BSPG 1922 X 46 and other material from Bahariya to be destroyed during a British bombing raid on Munich during the night of April 24/25, 1944.{{Cite journal |last1=Smith |first1=Joshua B. |last2=Lamanna |first2=Matthew C. |last3=Mayr |first3=Helmut |last4=Lacovara |first4=Kenneth J. |date=2006 |title=New information regarding the holotype of Spinosaurus aegyptiacus Stromer, 1915 |journal=Journal of Paleontology |volume=80 |issue=2 |pages=400–406 |doi=10.1666/0022-3360(2006)080[0400:NIRTHO]2.0.CO;2 |s2cid=130989487 }}{{sfn|Nothdurft|Smith|2002|p=117}} An endocast was made and survived the war, being the only remaining relic of the specimen.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=162, 164}} However in 2025, this specimen was redescribed as the holotype of a distinct carcharodontosaurid genus, Tameryraptor.

Few discoveries of Carcharodontosaurus attributed material were made until 1995 when American paleontologist Paul Sereno found an incomplete skull during an expedition embarked on by the University of Chicago. This skull (SGM-Din 1) was found in the Cenomanian-aged rocks of the Lower Douira Formation, Kem Kem Beds, in Errachidia, southeastern Morocco. The specimen was taken to the University of Chicago and was first described in 1996 by Sereno and colleagues in Science. In 2007, SGM-Din 1 was officially designated as the neotype of C. saharicus due to the loss of other specimens and the similar age and geographic location to previously noted material. The taxonomy of Carcharodontosaurus was discussed by Chiarenza and Cau (2016),{{Cite journal |last1=Chiarenza |first1=Alfio Alessandro |last2=Cau |first2=Andrea |date=February 29, 2016 |title=A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa |journal=PeerJ |volume=4 |pages=e1754 |doi=10.7717/peerj.1754 |pmc=4782726 |pmid=26966675 |doi-access=free }} who suggested that the neotype of C. saharicus was similar but distinct from the skeleton described by Stromer in the morphology of the maxillary interdental plates. However, paleontologist Mickey Mortimer put forward that the suggested difference between the two specimens was actually due to damage to the neotype.{{Cite web |last=Mortimer |first=Mickey |date=2023 |title=Carnosauria |url=https://theropoddatabase.com/Carnosauria.htm#Carcharodontosaurussaharicus |url-status=live |archive-url=https://web.archive.org/web/20230528042437/https://theropoddatabase.com/Carnosauria.htm |archive-date=May 28, 2023 |access-date=June 6, 2023 |website=The Theropod Database}} Because the neotype designation was in accordance with the ICZN article 75.3 and 75.4, the describers of Tameryraptor agreed that SGM-Din 1 is a valid neotype.{{cite journal |last1=Kellermann |first1=Maximilian |last2=Cuesta |first2=Elena |last3=Rauhut |first3=Oliver W. M. |title=Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny |journal=PLOS ONE |date=January 14, 2025 |volume=20 |issue=1 |pages=e0311096 |doi=10.1371/journal.pone.0311096|doi-access=free |pmid=39808629 |pmc=11731741 |bibcode=2025PLoSO..2011096K }}

Several other fossils of C. saharicus have been unearthed from the Kem Kem Beds, such as dentary fragments, a cervical vertebra, and many teeth.{{Cite journal |last=Russell |first=Dale |date=1996 |title=Isolated Dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco |url=https://sciencepress.mnhn.fr/fr/periodiques/bulletin-du-museum-national-d-histoire-naturelle-4eme-serie-section-c-sciences-de-la-terre-paleontologie-geologie-mineralogie/18/2-3/os-isoles-de-dinosaures-du-cretace-moyen-du-tafilalt-maroc |journal=Bulletin du Muséum national d'Histoire naturelle, 4ème série – section C – Sciences de la Terre, Paléontologie, Géologie, Minéralogie |language=fr |volume=18 |issue=2–3}}{{cite book |last1=Gheerbrant |first1=Emmanuel |last2=Cappetta |first2=Henri |last3=Broin |first3=F. de Lapparent de |last4=Rage |first4=Jean Claude |last5=Tabuce |first5=Rodolphe |title=Mémoires de la société géologique de France |date=2017 |publisher=Société Géologique de France |isbn=978-2-85363-099-3 |pages=485–525 |chapter-url=https://mnhn.hal.science/mnhn-02264868/ |language=fr |chapter=Les faunes de vertébrés marins et terrestres du Paléogène du Bassin d'Ouarzazate, Maroc }}{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=167}} Sereno et al. also referred a multitude of cervical vertebrae described as the spinosaurids Sigilmassasaurus and "Spinosaurus B" to C. saharicus reasoning that stout cervicals would be needed to carry the skulls of carcharodontosaurids. Later research proved otherwise, with the vertebrae being placed in Spinosaurus aegyptiacus by Ibrahim et al. (2020).{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=169}} French paleontologist René Lavocat was the first to note the possible presence of Carcharodontosaurus in Morocco as early as 1954.{{Cite journal |last=Lavocat |first=Rene |date=1954 |title=Sur les dinosauriens du Continental Intercalaire des Kem-Kem de la Daoura |journal=Comptes Rendus 19th International Geological Congress |volume=1952 |pages=65–68}}

In 2007, a novel species of Carcharodontosaurus, C. iguidensis, was dubbed by paleontologists Steve Brusatte and Paul Sereno. Fossils of C. iguidensis had been uncovered during an expedition to the Echkar Formation of Iguidi, Niger, a partial maxilla (MNN IGU2) being designated the holotype. The species name iguidensis is after Iguidi, where the fossils were unearthed. Several other remains such as a braincase, a lacrimal, a dentary, a cervical vertebra, and a collection of teeth were referred to C. iguidensis based on size and supposed similarities to other Carcharodontosaurus bones. Chiarenza and Cau (2016) identified the referred material of C. iguidensis as belonging to Sigilmassasaurus (later referred to Spinosaurus sp.){{Cite journal |last1=Sereno |first1=Paul C. |last2=Myhrvold |first2=Nathan |last3=Henderson |first3=Donald M. |last4=Fish |first4=Frank E. |last5=Vidal |first5=Daniel |last6=Baumgart |first6=Stephanie L. |last7=Keillor |first7=Tyler M. |last8=Formoso |first8=Kiersten K. |last9=Conroy |first9=Lauren L. |date=November 30, 2022 |editor-last=Zhu |editor-first=Min |editor2-last=Rutz |editor2-first=Christian |editor3-last=Zhu |editor3-first=Min |editor4-last=Holtz |editor4-first=Thomas R. |editor5-last=Hone |editor5-first=David |title=Spinosaurus is not an aquatic dinosaur |journal=eLife |volume=11 |pages=e80092 |doi=10.7554/eLife.80092 |pmc=9711522 |pmid=36448670 |doi-access=free }} and a non-carcharodontosaurine, and therefore chose to limit C. iguidensis to the holotype pending future research. Another carcharodontosaurid from the Kem Kem Beds, Sauroniops pachytholus, was dubbed in 2012 based on a single frontal,{{Cite journal |last1=Cau |first1=Andrea |last2=Dalla Vecchia |first2=Fabio M. |last3=Fabbri |first3=Matteo |date=March 1, 2013 |title=A thick-skulled theropod (Dinosauria, Saurischia) from the Upper Cretaceous of Morocco with implications for carcharodontosaurid cranial evolution |journal=Cretaceous Research |volume=40 |pages=251–260 |doi=10.1016/j.cretres.2012.09.002 |bibcode=2013CrRes..40..251C }} and has been proposed to be synonymous with C. saharicus.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=171}} This proposed synonymy has been disputed by others.{{Cite journal |last1=Candeiro |first1=Carlos Roberto dos Anjos |last2=Brusatte |first2=Stephen Louis |last3=Vidal |first3=Luciano |last4=Pereira |first4=Paulo Victor Luiz Gomes da Costa |date=July 26, 2018 |title=Paleobiogeographic evolution and distribution of Carcharodontosauridae (Dinosauria, Theropoda) during the middle Cretaceous of North Africa |journal=Papéis Avulsos de Zoologia |volume=58 |pages=e20185829 |doi=10.11606/1807-0205/2018.58.29 |s2cid=53353652 |hdl=20.500.11820/c4ca0a5c-4f8e-4136-8355-8bd32d6ea544 |hdl-access=free }}{{Cite journal |last1=Paterna |first1=Alessandro |last2=Cau |first2=Andrea |date=October 11, 2022 |title=New giant theropod material from the Kem Kem Compound Assemblage (Morocco) with implications on the diversity of the mid-Cretaceous carcharodontosaurids from North Africa |journal=Historical Biology |volume=35 |issue=11 |pages=2036–2044 |doi=10.1080/08912963.2022.2131406 |s2cid=252856791 }} The South American genus Giganotosaurus was synonymized with Carcharodontosaurus by Figueiredo (1998){{Cite news |last=Figueiredo |date=1998 |title=Os dinossáurios carnívoros: A sua descrição e modo de vida |pages=1–4 |work=Centro Portugues de Geo-historia e Prehistoria}} and Paul (2010),{{Cite book |last=Paul |first=Gregory S. |url=http://worldcat.org/oclc/985402380 |title=The Princeton Field Guide to Dinosaurs |publisher=Princeton University Press |year=2016 |isbn=978-1-78684-190-2 |pages=103–104 |oclc=985402380 |author-link=Gregory S. Paul}} but no authors have since followed this assessment.

{{location map+|Africa|relief=yes|width=300|float=|caption=Fossil localities of Carcharodontosaurus

Legend: 8px C. saharicus 8px C. iguidensis 8px Possible specimens|places={{location map~ | Africa| label =Continental Interclaire Formation | position = none | lat=27.0333| long=1.0833| mark= Steel pog.svg}}

{{location map~ | Africa| label =Kem Kem Beds | position = none | lat=31.5317| long=-4.6656| mark= Steel pog.svg}}

{{location map~ | Africa| label =Douiret Formation | position = none | lat=32.691| long= 10.261| mark= Steel pog.svg}}

{{location map~ | Africa| label =Continental Interclaire Formation (type locality) | position = none | lat=29.25| long= 0.25| mark= Steel pog.svg}}

{{location map~ | Africa| label =Continental Interclaire Formation | position = none | lat=28.369| long= 9.381| mark= Steel pog.svg}}

{{location map~ | Africa| label =Gara Samani Formation | position = none | lat=32.7022| long= -0.0067| mark= Steel pog.svg}}

{{location map~ | Africa| label = Elrhaz Formation | position = none | lat=16.1292| long=10.222| mark= Orange_pog.svg}}

{{location map~ | Africa| label = Echkar Formation| position = none | lat=17.9333| long= 5.6167| mark= Red pog.svg}}|alt=Map of sites preserving Carcharodontosaurus.}}

• Lapparent (1951, 1960) described several Carcharodontosaurus teeth from the Continental intercalaire Formation of Guermessa, Tunisia.{{cite journal|last1=Buffetaut|first1=Éric|last2=Ouaja|first2=Mohamed|title=A new specimen of Spinosaurus (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae|journal=Bulletin de la Société géologique de France|year=2002|volume=173|issue=5|pages=415–421|doi=10.2113/173.5.415|s2cid=53519187|url=https://doc.rero.ch/record/14728/files/PAL_E1854.pdf}}
• A postorbital bone and several postcranial remains assigned to Carcharodontosaurus were found in the Elrhaz Formation of northern Niger. Taquet (1976) noted that the postorbital was similar to that of Acrocanthosaurus, a relative of Carcharodontosaurus,{{sfn|Taquet|1976|p=53}} while the postcranial fossils could belong to other theropods.
• Two braincase fragments, 137 teeth, two caudal vertebrae, and a manual phalanx from the Echkar Formation were referred to as Carcharodontosaurus by Lapparent (1960). A pedal phalanx had also been described as Carcharodontosaurus but it likely is from a spinosaurid instead.{{Cite journal |last1=Ibrahim |first1=Nizar |last2=Sereno |first2=Paul C. |last3=Dal Sasso |first3=Cristiano |last4=Maganuco |first4=Simone |last5=Fabbri |first5=Matteo |last6=Martill |first6=David M. |last7=Zouhri |first7=Samir |last8=Myhrvold |first8=Nathan |last9=Iurino |first9=Dawid A. |date=September 26, 2014 |title=Semiaquatic adaptations in a giant predatory dinosaur |journal=Science |volume=345 |issue=6204 |pages=1613–1616 |doi=10.1126/science.1258750 |pmid=25213375 |bibcode=2014Sci...345.1613I |s2cid=34421257 |doi-access=free }}
• Many vertebrae, including two associated dorsals, were found in the Early Cretaceous strata of the Irhazer Group of Agadez, Niger. Lapparent mentioned these fossils as C. saharicus in 1960, though they may belong to other theropod genera.
• Caudal vertebrae from the Tefidet and teeth from Akarazeras sites of the Continental intercalaire Formation of Agadez, Niger were recorded by Lapparent (1960) and Taquet (1976) respectively.{{sfn|Taquet|1976|p=53}} The vertebrae could be from other theropods.
• From an unknown locale in the Continental intercalaire of the Sahara Desert, Lapparent (1960) documented eight vertebrae, a humerus, and a manual phalanx as coming from C. saharicus. These elements could be from other theropods.
• Teeth and a caudal vertebra from the Chenini Formation of southern Tunisia have been referred to Carcharodontosaurus.{{cite journal |last1=Schlüter |first1=T |last2=Schwarzhans |first2=W |date=1978 |title=Eine Bonebed-Lagerstätte aus dem Wealden Süd Tunesiens (Umgebung Ksar Krerachfa) |journal=Berliner Geowiss. Abhandlungen A |volume=8 |pages=53–65 }} However, the caudal vertebra is now labeled Carcharodontosauridae indet.{{cite journal |last1=Fanti |first1=Federico |last2=Cau |first2=Andrea |last3=Martinelli |first3=Agnese |last4=Contessi |first4=Michela |title=Integrating palaeoecology and morphology in theropod diversity estimation: A case from the Aptian-Albian of Tunisia |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |date=September 2014 |volume=410 |pages=39–57 |doi=10.1016/j.palaeo.2014.05.033 |bibcode=2014PPP...410...39F }}
• In 2015, a large neural arch of a dorsal vertebra from the Kem Kem Beds was informally described as belonging to a new genus and species of megaraptoran dubbed "Osteoporosia gigantea". This specimen is owned by the head of a Polish theme park chain who described it as belonging to a {{Convert|15|m|ft}} long carnosaur similar to Mapusaurus and Carcharodontosaurus.{{Cite web |last=Singer |date=2015 |title=JuraPark na tropie nowych dinozaurow z Maroka. |url=https://jurapark.pl/jurapark-na-tropie-nowych-dinozaurow-z-maroka/ |url-status=live |archive-url=https://web.archive.org/web/20151206224352/https://jurapark.pl/jurapark-na-tropie-nowych-dinozaurow-z-maroka/ |archive-date=December 6, 2015 |access-date=June 27, 2023 |website=Jurapark}} However, it was much smaller than proposed and may belong to C. saharicus or Sauroniops based on its carcharodontosaurid traits and origin.{{Cite book |last1=Molina-Pérez |first1=Rubén |title=Dinosaur Facts and Figures: The Theropods and Other Dinosauriformes |last2=Larramendi |first2=Asier |last3=Connolly |first3=David |last4=Cruz |first4=Gonzalo Ángel Ramírez |date=2019 |publisher=Princeton University Press |isbn=978-0-691-19059-4 }}{{pn|date=October 2024}}

• A maxillary tooth recovered from the Villar del Arzobispo Formation of Spain was referred to Carcharodontosaurus in 1966.{{cite journal|language=es|first1=M.|last1=Crusafont-Pairó|first2=R.|last2=Adrover|year=1966|title=El primer representante de la clase mamíferos hallado en el Mesozoico de España|journal=Teruel|volume=35|pages=139–143}} However, it lacks the traits of carcharodontosaurid teeth and instead is more similar to that of other allosauroids.{{Cite journal |last1=Kuhne |first1=W. G. |last2=Crusafont-Pairo |first2=M. |date=1968 |title=Mamíferos del Wealdiense de Uña, cerca de Cuenca |url=https://www.raco.cat/index.php/ActaGeologica/article/download/74602/97493 |journal=Acta Geológica Hispánica |volume=3 |issue=5 |pages=133–134}}{{Cite journal |last1=Gascó |first1=Francisco |last2=Cobos |first2=Alberto |last3=Royo-Torres |first3=Rafael |last4=Mampel |first4=Luis |last5=Alcalá |first5=Luis |date=June 1, 2012 |title=Theropod teeth diversity from the Villar del Arzobispo Formation (Tithonian–Berriasian) at Riodeva (Teruel, Spain) |journal=Palaeobiodiversity and Palaeoenvironments |volume=92 |issue=2 |pages=273–285 |doi=10.1007/s12549-012-0079-3 |bibcode=2012PdPe...92..273G |s2cid=129930988 }}
• Bond and Bromley (1970) described teeth deriving from the Gokwe Formation of Zimbabwe as being similar to Carcharodontosaurus, with Mickey Mortimer assigning them to the genus tentatively.{{Cite journal |last1=Bond |first1=Geoffrey |last2=Bromley |first2=K. |date=December 1970 |title=Sediments with the remains of dinosaurs near Gokwe, Rhodesia |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=8 |issue=4 |pages=313–327 |bibcode=1970PPP.....8..313B |doi=10.1016/0031-0182(70)90104-5 |s2cid=128716797 }} However, later studies have found them to be indeterminate.{{Cite journal|author1=Munyikwa, D.|author2=Sampson, S. D.|author3=Rogers, R. R.|author4=Forster, C. A. |author5=Curry, K. A. |author6=Curtice, B. D. |date=1998 |title=Vertebrate palaeontology and geology of the Gokwe Formation, Zimbabwe |journal=Journal of African Earth Sciences |volume=27 |issue=1 |pages=142}}
• Teeth from the Alcantara Formation of Brazil were placed in Carcharodontosaurus in 2002,{{Cite journal |last1=Medeiros |first1=Manuel |last2=Schultz |first2=Cesar |date=2002 |title=A fauna dinossauriana da 'Laje do Coringa', Cretaceo medio de Nordeste do Brasil |id={{BHL page|57292482}} |journal=Arquivos do Museo Nacional, Rio de Janeiro |volume=60 |issue=3 |pages=155–162 }} but this has been disputed based on its geographic origin.
• Partial specimens from Wadi Milk Formation were originally assigned to Carcharodontosaurus, but were now considered to be indeterminate carcharodontosaurids, some of which are similar to the genus.
• Fossils from the Campanian Quseir Formation of western Egypt have been tentatively assigned to Spinosaurus and Carcharodontosaurus,{{cite book |last1=Churcher |first1=C. S. |chapter=A note on the Late Cretaceous vertebrate fauna of the Dakhleh Oasis |pages=55–68 |editor1-last=Churcher |editor1-first=C. S. |editor2-last=Mills |editor2-first=Anthony J. |title=Reports from the Survey of the Dakhleh Oasis, Western Desert of Egypt, 1977-1987 |date=1999 |publisher=Oxbow Books |isbn=978-1-900188-49-4 }} but these specimens were never described in detail and thus classified as Theropoda indet.{{cite book|last1=Augustin|first1=F.J.|last2=Hartung|first2=J.|last3=Kampouridis|first3=P.|editor1=Hamimi, Z.|editor2=Khozyem, H.|editor3=Adatte, T.|editor4=Nader, Fadi H.|editor5=Oboh-Ikuenobe, F.|editor6=Zobba, Mohamed K.|editor7=Atfy, Haytham El|year=2023|chapter=Dinosaur Faunas of Egypt—The Terrestrial Late Cretaceous Vertebrate Record|title=The Phanerozoic Geology and Natural Resources of Egypt|pages=253–284|series=Advances in Science, Technology & Innovation|publisher=Springer, Cham|doi=10.1007/978-3-030-95637-0_9|isbn=978-3-030-95636-3}}

File:Longest theropods.svg, C. saharicus in orange, far right]]

Stromer hypothesized that C. saharicus was around the same size as the tyrannosaurid Gorgosaurus, which would place it at around {{convert|8|-|9|m}} long, based on his specimen SNSB-BSPG 1922 X 46 (now Tameryraptor). This individual was around 15% smaller than the neotype,{{Sfn|Nothdurft|Smith|2002|p=109}} the latter was estimated to be {{convert|12|-|12.5|m}} in length and approximately {{convert|5|-|7|MT|ST}} in body mass.{{cite journal |last1=Henderson |first1=D.M. |last2=Nicholls |first2=R. |year=2015 |title=Balance and Strength—Estimating the Maximum Prey-Lifting Potential of the Large Predatory Dinosaur Carcharodontosaurus saharicus |journal=The Anatomical Record |volume=298 |issue=8 |pages=1367–1375 |doi=10.1002/ar.23164 |pmid=25884664 |s2cid=19465614|doi-access=free }}{{cite journal |last1=Seebacher |first1=Frank |title=A new method to calculate allometric length-mass relationships of dinosaurs |journal=Journal of Vertebrate Paleontology |date=March 26, 2001 |volume=21 |issue=1 |pages=51–60 |doi=10.1671/0272-4634(2001)021[0051:ANMTCA]2.0.CO;2 }}{{Cite book |title=Tyrannosaurid Paleobiology |last1=Hurlburt |first1=G. S. |last2=Ridgely |first2=R. C. |last3=Witmer |first3=L. M. |date=July 5, 2013 |publisher=Indiana University Press |isbn=978-0-253-00947-0 |editor-last=Parrish |editor-first=M. J. |pages=134–154 |chapter=Relative size of brain and cerebrum in Tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators |access-date=October 20, 2013 |editor-last2=Molnar |editor-first2=R. E. |editor-last3=Currie |editor-first3=P. J. |editor-last4=Koppelhus |editor-first4=E. B. |chapter-url=https://www.researchgate.net/publication/256536375}} This makes Carcharodontosaurus saharicus one of the largest known theropod dinosaurs and one of the largest terrestrial carnivores. C. iguidensis was much smaller, only reaching {{convert|10|m}} in length and {{convert|4|MT|ST}} in body mass.

File:Carcharodontosaurus_saharicus_skull_reconstruction.png

The largest and most complete skull of C. saharicus would measure {{Convert|1.6|m|ft}} when complete, around the same size as the largest Tyrannosaurus skulls. No skulls of the genus preserve premaxillae, complete posterior skull regions, or mandibles. Skulls of carcharodontosaurids tend to be more slender and lightly built than those of later tyrannosaurids, which have robust builds and adaptations for crushing. The neotype cranium tapers towards the front in side view creating a triangular outline. This is similar to that of other carcharodontosaurids like Mapusaurus and Giganotosaurus. Its skull was lighter than that of tyrannosaurids, with the antorbital fenestra composing over 30% of the total skull length as well as being surrounded by {{Dinogloss|fossae}} in the maxillae (upper jaw bone), nasals (nose bone), jugals (cheekbone), and lacrimals (front orbit bone). Akin to other genera, its nasal is elongated and its exposed side is covered in a rugose surface. These bumps were likely extended by keratin sheaths, creating a horn-like structure as in Ceratosaurus. A similar rugosity is found on the lacrimal which would also be lengthened by keratin, forming a similar element.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=165}} The most distinctive trait of Carcharodontosaurus{{'}} skull is the sculpted exterior of the maxillae, which is unique to the genus. C. iguidensis has antorbital fossae limited to the proximity of the antorbital fenestra, a crest running along the medial (right) face of the maxilla, and a process along its midline. These traits are missing in C. saharicus, differentiating the two species.

{{multiple image

| align = left

| total_width = 240

| perrow = 2/2

| image1 = Right maxilla of Carcharodontosaurus.jpg

| alt1 = Right maxilla of C. saharicus

| image2 = Carcharodontosaurus nasal and lacrimal bone.jpg

| alt2 = Nasal and lacrimal of C. saharicus

| image3 = Carcharodontosaurus jugal bones.jpg

| alt3 = Jugals of C. saharicus

| image4 = Carcharodontosaurus postorbital bones.jpg

| alt4 = Postorbitals of C. saharicus

| footer = Skull bones of the neotype specimen; right maxilla (upper left), nasal and lacrimal (upper right), jugals (lower left), and postorbitals (lower right)

}}

14 teeth sockets are present in each maxilla. Parts of the braincase are known though much of their morphology is the same as Giganotosaurus{{'}}. However, C. saharicus has a much more prominent nuchal crest, which overhangs the skull roof. The frontal bones are firmly fused, a characteristic evident in most theropods.{{cite journal |last1=Coria |first1=Rodolfo A. |last2=Currie |first2=Philip J. |title=The braincase of Giganotosaurus carolinii (Dinosauria: Theropoda) from the Upper Cretaceous of Argentina |journal=Journal of Vertebrate Paleontology |date=January 14, 2003 |volume=22 |issue=4 |pages=802–811 |doi=10.1671/0272-4634(2002)022[0802:TBOGCD]2.0.CO;2 }} The jugals are broad and triangle-shaped. The lower jaw articulation was placed farther back behind the occipital condyle (where the neck is attached to the skull) compared to other theropods. Two dentary (lower jaw bone) fragments which were referred to C. saharicus by Ibrahim et al. (2020) have deep and expanded alveoli (tooth sockets), traits found in other large theropods.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=167}} If like Tyrannotitan and Giganotosaurus, the dentary would have 16 alveoli (tooth sockets).{{Cite journal |last1=Novas |first1=Fernando E. |last2=de Valais |first2=Silvina |last3=Vickers-Rich |first3=Pat |last4=Rich |first4=Tom |date=May 1, 2005 |title=A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids |journal=Naturwissenschaften |volume=92 |issue=5 |pages=226–230 |doi=10.1007/s00114-005-0623-3 |pmid=15834691 |bibcode=2005NW.....92..226N |hdl=11336/103474 |s2cid=24015414 |hdl-access=free }}

Estimations of the tooth count of Carcharodontosaurus vary, but a recent estimate of 30 dentary, 8 premaxillary, and 24 maxillary teeth for a total of 62 teeth was made.{{Cite journal |last1=Beevor |first1=Thomas |last2=Quigley |first2=Aaron |last3=Smith |first3=Roy E. |last4=Smyth |first4=Robert S. H. |last5=Ibrahim |first5=Nizar |last6=Zouhri |first6=Samir |last7=Martill |first7=David M. |date=January 1, 2021 |title=Taphonomic evidence supports an aquatic lifestyle for Spinosaurus |journal=Cretaceous Research |volume=117 |pages=104627 |doi=10.1016/j.cretres.2020.104627 |bibcode=2021CrRes.11704627B |s2cid=224888268 |url=https://researchportal.port.ac.uk/portal/en/publications/taphonomic-evidence-supports-an-aquatic-lifestyle-for-spinosaurus(e7fb2358-2ac6-4b6c-9697-225a525e8366).html }} Carcharodontosaurid teeth are some of the largest of any dinosaur group, with a maxillary tooth from SNSB-BSPG 1922 X 46 being {{Convert|6.8|cm|in}} tall and {{Convert|3.5|cm|in}} wide.{{Cite web |title=Discoveries {{!}} Paul Sereno - Paleontologist {{!}} The University of Chicago |url=https://paulsereno.uchicago.edu/discoveries/carcharodontosaurus/#:~:text=Carcharodontosaurus%20is%20Africa%27s%20answer%20to,-inch-long%20serrated%20teeth. |access-date=June 30, 2023 |website=paulsereno.uchicago.edu}} However, they are extremely thin, with most being under a centimeter thick. Serrations are numerous on the anterior and posterior margins, with over 18 to 20 serrations per centimeter of edge in C. saharicus and up to 32 per centimeter in C. iguidensis. Its teeth are straight, laterally flattened, and spindle-shaped in cross-section. However, dentition towards the back of the mouth became more recurved than those in the maxilla. The posterior margin of these crowns are recurved and convex at its termination. Bowed enamel wrinkles are present on both dorsoventral sides of the crowns. These wrinkles curve towards the marginal serrations, composing a band-shape along the ends.{{Cite journal |last1=Brusatte |first1=Stephen L. |last2=Benson |first2=Roger B. J. |last3=Carr |first3=Thomas D. |last4=Williamson |first4=Thomas E. |last5=Sereno |first5=Paul C. |date=December 12, 2007 |title=The systematic utility of theropod enamel wrinkles |journal=Journal of Vertebrate Paleontology |volume=27 |issue=4 |pages=1052–1056 |doi=10.1671/0272-4634(2007)27[1052:TSUOTE]2.0.CO;2 |s2cid=85615205 }}

{{multiple image

| align = right

| direction = vertical

| image1 = Carcharodontosaurus braincase.jpg

| image2 = Endocasts of Carcharodontosaurus.jpg

| footer = Braincase of the neotype (above), with endocasts of same (A–D) and MB. R. 2056 (E–F) below

| alt1 = Braincase of the C. saharicus neotype

| alt2 = Endocast of the lost Tameryraptor skull.

| total_width = 170

}}

In 2001, Hans C. E. Larsson published a description of the inner ear and endocranium of Carcharodontosaurus saharicus. Starting from the portion of the brain closest to the tip of the animal's snout is the forebrain, which is followed by the midbrain. The midbrain is angled downwards at a 45-degree angle and towards the rear of the animal. This is followed by the hindbrain, which is roughly parallel to the forebrain and forms a roughly 40-degree angle with the midbrain. Overall, the brain of C. saharicus would have been similar to that of a related dinosaur, Allosaurus fragilis. Larsson found that the ratio of the cerebrum to the volume of the brain overall in Carcharodontosaurus was typical for a non-avian reptile. Carcharodontosaurus also had a large optic nerve.{{cite book | last1 = Larsson | first1 = H.C.E. | date = 2001 | chapter = Endocranial anatomy of Carcharodontosaurus saharicus (Theropoda: Allosauroidea) and its implications for theropod brain evolution | title = Mesozoic Vertebrate Life | editor-first1 = D.H. | editor-last1 = Tanke | editor-last2 = Carpenter | editor-first2 = K. | editor-last3 = Skrepnick | editor-first3 = M. W. | publisher = Indiana University Press | pages = 219–236|isbn=978-0-253-33907-2|chapter-url=https://archive.org/details/mesozoicvertebra0000unse/page/19/mode/1up|url=https://archive.org/details/mesozoicvertebra0000unse/page/n3/mode/1up|url-access=registration}}

The three semicircular canals of the inner ear of Carcharodontosaurus saharicus—when viewed from the side—had a subtriangular outline. This subtriangular inner-ear configuration is present in Allosaurus, lizards, and turtles, but not in birds. The semi-"circular" canals themselves were very linear, which explains the pointed silhouette. In life, the floccular lobe of the brain would have projected into the area surrounded by the semicircular canals, just like in other non-avian theropods, birds, and pterosaurs.

Few postcranial elements are confidently known from Carcharodontosaurus, though many isolated bones from the Sahara have been referred to the genus without detailed study.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=169}}{{sfn|Taquet|1976|p=53}} However, the description of other carcharodontosaurids from North Africa such as Tameryraptor and Sauroniops has put into question the referral of carcharodontosaurid remains that lack overlap with the C. saharicus neotype. Like other carcharodontosaurids, it was robust with small forelimbs, an elongated tail, and short neck. The most complete specimen was SNSB-BSPG 1922 X 46, but it was destroyed and is now the holotype of Tameryraptor. A single cervical vertebra was referred to the genus by Dale A. Russell in 1996, the only described postcranial element recovered from the Kem Kem Beds that may belong to Carcharodontosaurus.{{Cite journal |last1=Evers |first1=Serjoscha W. |last2=Rauhut |first2=Oliver W. M. |last3=Milner |first3=Angela C. |last4=McFeeters |first4=Bradley |last5=Allain |first5=Ronan |date=2015-10-20 |title=A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the "middle" Cretaceous of Morocco |journal=PeerJ |language=en |volume=3 |pages=e1323 |doi=10.7717/peerj.1323 |doi-access=free |issn=2167-8359 |pmc=4614847 |pmid=26500829}} This cervical vertebra is stout and {{Dinogloss|opisthocoelus}} (concave posterior ends).{{cite journal |last=Harris |first=Jerald D. |year=1998 |title=A reanalysis of Acrocanthosaurus atokensis, its phylogenetic status, and paleobiological implications, based on a new specimen from Texas |journal=New Mexico Museum of Natural History and Science Bulletin |volume=13 |pages=1–75}} Its preserved length is {{Convert|148|mm|in}} As in Giganotosaurus, the vertebra is topped by low neural spines joined with sturdy {{Dinogloss|transverse processes}} which hung over the {{Dinogloss|pleurocoels}} (shallow depressions on the sides of centra), which would contain pneumatic air sacs to lighten the vertebrae. The centrum however lacks the keels observed in other carcharodontosaurids, possibly due to it being an anterior cervical vertebra.{{cite journal|first=Dale A.|last=Russell|title=Isolated Dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco|journal=Bulletin du Muséum National d'Histoire Naturelle|year=1996|volume=18|issue=2–3|series=4|pages=569–593|issn=0181-0642|url=https://www.biodiversitylibrary.org/item/249335#page/175/mode/1up}}

File:Carcharodontosaurus.png of C. saharicus]]

Carcharodontosaurus is the type genus of the family Carcharodontosauridae and subfamily Carcharodontosaurinae. This subfamily contains Carcharodontosaurus itself as well as the other carcharodontosaurines Giganotosaurus, Mapusaurus, Meraxes, and Tyrannotitan; however, these genera make up an independent tribe: Giganotosaurini. Carcharodontosauridae was a clade created by Stromer for Carcharodontosaurus and Bahariasaurus, though the name remained unused until the recognition of other members of the group in the late 20th century. He noted the likeness of Carcharodontosaurus bones to the American theropods Allosaurus and Tyrannosaurus, leading him to consider the family part of Theropoda.{{sfn|Stromer|1934|pp=60–61}}

Paul Sereno's description of Carcharodontosaurus fossils in 1996 led to the realization of a transcontinental clade of carcharodontosaurids. As more carcharodontosaurids were discovered, their interrelationships became even clearer. The group was defined as all allosauroids closer to Carcharodontosaurus than Allosaurus or Sinraptor by the paleontologist Thomas R. Holtz and colleagues in 2004.{{Cite journal |last1=Novas |first1=Fernando E. |last2=Agnolín |first2=Federico L. |last3=Ezcurra |first3=Martín D. |last4=Porfiri |first4=Juan |last5=Canale |first5=Juan I. |date=October 1, 2013 |title=Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia |journal=Cretaceous Research |volume=45 |pages=174–215 |doi=10.1016/j.cretres.2013.04.001 |bibcode=2013CrRes..45..174N |hdl=11336/102037|s2cid=129504278 |hdl-access=free }} Carcharodontosaurus is more poorly known than most other carcharodontosaurids, with Meraxes and Giganotosaurus represented by nearly complete skeletons.{{Cite journal |last1=Canale |first1=Juan I. |last2=Apesteguía |first2=Sebastián |last3=Gallina |first3=Pablo A. |last4=Mitchell |first4=Jonathan |last5=Smith |first5=Nathan D. |last6=Cullen |first6=Thomas M. |last7=Shinya |first7=Akiko |last8=Haluza |first8=Alejandro |last9=Gianechini |first9=Federico A. |last10=Makovicky |first10=Peter J. |date=July 7, 2022 |title=New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod arm reduction |journal=Current Biology |volume=32 |issue=14 |pages=3195–3202.e5 |doi=10.1016/j.cub.2022.05.057 |pmid=35803271 |s2cid=250343124 |doi-access=free |bibcode=2022CBio...32E3195C }} Carcharodontosaurians have been recognized from the Late Jurassic to the Mid-Cretaceous of every continent except Antarctica.{{Cite journal |last1=Coria |first1=Rodolfo A. |last2=Currie |first2=Philip J. |last3=Ortega |first3=Francisco |last4=Baiano |first4=Mattia A. |date=July 1, 2020 |title=An Early Cretaceous, medium-sized carcharodontosaurid theropod (Dinosauria, Saurischia) from the Mulichinco Formation (upper Valanginian), Neuquén Province, Patagonia, Argentina |journal=Cretaceous Research |volume=111 |pages=104319 |doi=10.1016/j.cretres.2019.104319 |bibcode=2020CrRes.11104319C |hdl=11336/122794 |s2cid=214475057 |hdl-access=free }}{{Cite journal |last1=Brusatte |first1=Stephen L. |last2=Benson |first2=Roger B.J. |last3=Xu |first3=Xing |date=2012 |title=A Reassessment of Kelmayisaurus petrolicus, a Large Theropod Dinosaur from the Early Cretaceous of China |journal=Acta Palaeontologica Polonica |volume=57 |issue=1 |pages=65–72 |doi=10.4202/app.2010.0125 |s2cid=53387460 |doi-access=free |hdl=20.500.11820/95de36fb-46b2-4acc-bd2e-1d5a496fc78c |hdl-access=free }}{{Cite journal |last1=Kotevski |first1=Jake |last2=Duncan |first2=Ruairidh J. |last3=Ziegler |first3=Tim |last4=Bevitt |first4=Joseph J. |last5=Vickers-Rich |first5=Patricia |last6=Rich |first6=Thomas H. |last7=Evans |first7=Alistair R. |last8=Poropat |first8=Stephen F. |title=Evolutionary and paleobiogeographic implications of new carcharodontosaurian, megaraptorid, and unenlagiine theropod remains from the upper Lower Cretaceous of Victoria, southeast Australia |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2024.2441903?s=04&fbclid=IwY2xjawIjLKdleHRuA2FlbQIxMQABHUyzuupFFj1lvGMHco5BrUwax1YTsiLx9T244449qie-QMvdFC7YqHssZw_aem_8X4FfXyB4WaNw3rjXwBrfg#abstract |journal=Journal of Vertebrate Paleontology |date=2025 |volume=0 |pages=e2441903 |doi=10.1080/02724634.2024.2441903 |issn=0272-4634|url-access=subscription }}

In their phylogenetic analyses, Kellermann, Cuesta & Rauhut (2025) recovered C. iguidensis as a non-carcharodontosaurine member of the Carcharodontosauridae outside the genus Carcharodontosaurus, suggesting that this species belongs to a different genus. Similar results were recovered by Cau & Paterna (2025) in their analysis of large Cretaceous theropods from Africa, who also argued for the removal of C. iguidensis from the genus.{{Cite journal |last1=Cau |first1=Andrea |author1-link=Andrea Cau |last2=Paterna |first2=Alessandro |date=May 2025 |title=Beyond the Stromer's Riddle: the impact of lumping and splitting hypotheses on the systematics of the giant predatory dinosaurs from northern Africa |url=https://www.italianjournalofgeosciences.it/297/article-1220/beyond-the-stromer-s-riddle-the-impact-of-lumping-and-splitting-hypotheses-on-the-systematics-of-the-giant-predatory-dinosaurs-from-northern-africa.html |journal=Italian Journal of Geosciences |volume=144 |issue=2 |pages=1–24 |doi=10.3301/IJG.2025.10|url-access=subscription }}

The analyses of Kellermann, Cuesta & Rauhut (2025) found support for a sister taxon relationship of carcharodontosaurids and metriacanthosaurids, which the authors named as a new clade, Carcharodontosauriformes. The results of their analysis using merged OTUs (operational taxonomic units) are displayed in the cladogram below:

{{clade

|{{clade

|1=Metriacanthosauridae

|label2=Carcharodontosauridae

|2={{clade

|1={{clade

|1=Lusovenator

|2=Lajasvenator }}

|3={{clade

|1=Acrocanthosaurus

|2={{clade

|1=Tameryraptor

|2={{clade

|1=Carcharodontosaurus iguidensis

|label2=Carcharodontosaurinae

|2={{clade

|1={{clade

|1=Taurovenator

|2=Carcharodontosaurus saharicus

|label3=Giganotosaurini

|3={{clade

|1=Meraxes

|2={{clade

|1=Tyrannotitan

|2={{clade

|1=Mapusaurus

|2=Giganotosaurus

}} }} }} }} }} }} }} }} }}|label1=Carcharodontosauriformes}}

Rodolfo Coria and Leonardo Salgado suggested that the convergent evolution of gigantism in theropods could have been linked to common conditions in their environments or ecosystems.{{Cite journal |last1=Coria |first1=Rodolfo A. |last2=Salgado |first2=Leonardo |date=1995 |title=A new giant carnivorous dinosaur from the Cretaceous of Patagonia |journal=Nature |volume=377 |issue=6546 |pages=224–226 |doi=10.1038/377224a0 |bibcode=1995Natur.377..224C |s2cid=30701725 }} Sereno and colleagues found that the presence of carcharodontosaurids in Africa (Carcharodontosaurus), North America (Acrocanthosaurus), and South America (Giganotosaurus), showed the group had a transcontinental distribution by the Early Cretaceous period. Dispersal routes between the northern and southern continents appear to have been severed by ocean barriers in the Late Cretaceous, which led to more distinct, provincial faunas, by preventing exchange.{{Cite journal |last=Currie |first=Philip J. |date=May 17, 1996 |title=Out of Africa: Meat-Eating Dinosaurs That Challenge Tyrannosaurus rex |journal=Science |volume=272 |issue=5264 |pages=971–972 |doi=10.1126/science.272.5264.971 |bibcode=1996Sci...272..971C |s2cid=85110425 }} Previously, it was thought that the Cretaceous world was biogeographically separated, with the northern continents being dominated by tyrannosaurids, South America by abelisaurids, and Africa by carcharodontosaurids.{{cite journal |last1=Coria |first1=Rodolfo A. |last2=Salgado |first2=Leonardo |date=June 1996 |title=Dinosaurios carnívoros de Sudamérica |url=http://www.investigacionyciencia.es/revistas/investigacion-y-ciencia/numero/237/dinosaurios-carnvoros-de-sudamrica-6530 |journal=Investigación y Ciencia |language=es |issue=237 |pages=39–40}} The subfamily Carcharodontosaurinae, in which Carcharodontosaurus belongs, appears to have been restricted to the southern continent of Gondwana (formed by South America and Africa), where they were probably the apex predators. The South American tribe Giganotosaurini may have been separated from their African relatives through vicariance, when Gondwana broke up during the Aptian–Albian ages of the Early Cretaceous.{{cite journal |last1=Canale |first1=J. I. |last2=Novas |first2=F. E. |last3=Pol |first3=D. |date=2014 |title=Osteology and phylogenetic relationships of Tyrannotitan chubutensis Novas, de Valais, Vickers-Rich and Rich, 2005 (Theropoda: Carcharodontosauridae) from the Lower Cretaceous of Patagonia, Argentina |journal=Historical Biology |volume=27 |issue=1 |pages=1–32 |doi=10.1080/08912963.2013.861830 |s2cid=84583928 |hdl-access=free |hdl=11336/17607 }}

A biomechanical analysis of Carcharodontosaurus' lifting capabilities was conducted by paleontologist Donald Henderson and paleoartist Robert Nicholls in 2015. The authors used 3D models of the animal as well as a subadult sauropod Limaysaurus, which although not found alongside Carcharodontosaurus, is similar to the rebbachisaurids of the Kem Kem Beds. The models included the size of the lungs and other pneumatic structures of the two, fostering an accurate weight simulation of the scenario. Henderson & Nicholls' study found that an adult C. saharicus could hold a maximum of {{cvt|424|kg}}, half the weight of an adult Limaysaurus. However, two C. saharicus adults could together lift as much as {{Convert|850|kg|lb}}.

File:Carcharodontosaurus teeth.jpg

The dentition of allosauroids is distinct, with carcharodontosaurid teeth bearing distinctly thin and blade-like teeth. However, these teeth are thin and likely could not sustain impact against hard surfaces like bone without potentially bending and snapping. This danger is exacerbated by the straight edges, slightly recurved tips, and sinusoidal shapes observed in their dentition. Despite these traits, the teeth are still much more robust than those of smaller theropods and due to their overall size could take more pressure. Carcharodontosaurus also had a high tooth replacement rate meaning that damaged teeth could be replaced easily in contrast to extant bone-crushing mammals who spend much of their energy maintaining their teeth.{{Cite journal |last=Van Valkenburgh |first=Blaire |date=1988 |title=Incidence of Tooth Breakage Among Large, Predatory Mammals |journal=The American Naturalist |volume=131 |issue=2 |pages=291–302 |doi=10.1086/284790 |s2cid=222330098 |jstor=2461849|bibcode=1988ANat..131..291V }}{{Cite journal |last=Van Valkenburgh |first=Blaire |year=2008 |title=Costs of carnivory: tooth fracture in Pleistocene and Recent carnivorans |journal=Biological Journal of the Linnean Society |volume=96 |issue=1 |pages=68–81 |doi=10.1111/j.1095-8312.2008.01108.x |s2cid=85623961 |doi-access=free }} Evidence of bone-crunching bites is observed in Allosaurus, which would engage in ritual face-biting with other individuals and bite into the pelves of Stegosaurus as shown by bite marks.{{Cite journal |last1=Hone |first1=David W. E. |last2=Rauhut |first2=Oliver W. M. |date=2010 |title=Feeding behaviour and bone utilization by theropod dinosaurs |journal=Lethaia |volume=43 |issue=2 |pages=232–244 |doi=10.1111/j.1502-3931.2009.00187.x |bibcode=2010Letha..43..232H |s2cid=86037076}}{{Cite journal |last1=Tanke |first1=Darren H.|last2=Currie |first2=Phillip J. |date=1998 |title=Head-biting behavior in theropod dinosaurs: Paleopathological evidence |journal=Gaia|issue=15|pages=167–184|doi=10.7939/R34T6FJ1P |s2cid=90552600 |doi-access=free}}{{Cite journal |last1=Drumheller |first1=Stephanie K. |last2=McHugh |first2=Julia B. |last3=Kane |first3=Miriam |last4=Riedel |first4=Anja |last5=D’Amore |first5=Domenic C. |date=May 27, 2020 |title=High frequencies of theropod bite marks provide evidence for feeding, scavenging, and possible cannibalism in a stressed Late Jurassic ecosystem |journal=PLOS ONE |volume=15 |issue=5 |pages=e0233115 |doi=10.1371/journal.pone.0233115 |pmid=32459808 |pmc=7252595 |bibcode=2020PLoSO..1533115D |doi-access=free }}

Bite forces of Carcharodontosaurus as well as other giant theropods including Acrocanthosaurus and Tyrannosaurus have been analyzed. Studies reported that carcharodontosaurids had much lower bite forces than Tyrannosaurus despite being in the same size class. The anterior bite force of C. saharicus was estimated in a 2022 paper to be 11,312 newtons while the posterior bite force was 25,449 newtons. This is much lower than that of Tyrannosaurus, implying that it did not eat bones.{{Cite journal |last=Sakamoto |first=Manabu |date=July 12, 2022 |title=Estimating bite force in extinct dinosaurs using phylogenetically predicted physiological cross-sectional areas of jaw adductor muscles |journal=PeerJ |volume=10 |pages=e13731 |doi=10.7717/peerj.13731 |pmc=9285543 |pmid=35846881 |doi-access=free }}{{Cite journal |last1=Gignac |first1=Paul M. |last2=Erickson |first2=Gregory M. |date=May 17, 2017 |title=The Biomechanics Behind Extreme Osteophagy in Tyrannosaurus rex |journal=Scientific Reports |volume=7 |issue=1 |pages=2012 |doi=10.1038/s41598-017-02161-w |pmc=5435714 |pmid=28515439|bibcode=2017NatSR...7.2012G |doi-access=free }} Finite element accounts of the skulls of theropods have also been taken, which further supported the idea that Carcharodontosaurus ate softer food than tyrannosaurids. Great amounts of stress were recovered in the posterior part of the cranium near the quadrate in Carcharodontosaurus, Spinosaurus, and Acrocanthosaurus. The skulls of these theropods had higher relative stress quantities in opposition to that of smaller genera. This indicates that the crania of giant taxa (ex. Carcharodontosaurus) were unstable due to having large pneumatic structures to save weight instead of creating a firm build. However, Spinosaurus and Suchomimus experienced even greater values of stress meaning that they could only consume light, small prey instead of larger items, which the stronger skull of Carcharodontosaurus could bite while sustaining the stress.{{cite book |last1=Rayfield |first1=Emily J. |title=STUDIES ON FOSSIL TETRAPODS |date=2011 |publisher=Palaeontological Association |pages=241–253 |chapter=Structural performance of tetanuran theropod skulls, with emphasis on the Megalosauridae, Spinosauridae and Carcharodontosauridae |hdl=1983/aaaea1c8-8c3a-4f99-abd6-982b47664aac |isbn=978-1-4443-6189-6 }}

Isotopic analyses of the teeth of C. saharicus have found δ18O values that are higher than that of the contemporary Spinosaurus, suggesting the latter pursued semi-aquatic habits whereas Carcharodontosaurus was more terrestrial.{{Cite journal |last1=Goedert |first1=J. |last2=Amiot |first2=R. |last3=Boudad |first3=L. |last4=Buffetaut |first4=E. |author-link4=Éric Buffetaut |last5=Fourel |first5=F. |last6=Godefroit |first6=P. |last7=Kusuhashi |first7=N. |last8=Suteethorn |first8=V. |last9=Tong |first9=H. |last10=Watabe |first10=M. |last11=Lecuyer |first11=C. |date=2016 |title=Preliminary investigation of seasonal patterns recorded in the oxygen isotope compositions of theropod dinosaur tooth enamel. |journal=PALAIOS |volume=31 |issue=1 |pages=10–19|doi=10.2110/palo.2015.018 |bibcode=2016Palai..31...10G |s2cid=130878403 }} This is further supported by the taphonomy of C. saharicus teeth, which are more often found in land terrains than aquatic ones. Carcharodontosaurus was also a homeotherm with an endotherm-like thermophysiology as inferred by these isotopes meaning that most of its oxygen was accumulated by drinking water rather than being in it.{{Cite journal |last1=Amiot |first1=Romain |last2=Wang |first2=Xu |last3=Lécuyer |first3=Christophe |last4=Buffetaut |first4=Eric |last5=Boudad |first5=Larbi |last6=Cavin |first6=Lionel |last7=Ding |first7=Zhongli |last8=Fluteau |first8=Frédéric |last9=Kellner |first9=Alexander W.A. |last10=Tong |first10=Haiyan |last11=Zhang |first11=Fusong |date=2010 |title=Oxygen and carbon isotope compositions of middle Cretaceous vertebrates from North Africa and Brazil: Ecological and environmental significance |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |volume=297 |issue=2 |pages=439–451 |doi=10.1016/j.palaeo.2010.08.027 |bibcode=2010PPP...297..439A |s2cid=131607253 }}{{Cite journal |last1=Amiot |first1=Romain |last2=Lécuyer |first2=Christophe |last3=Buffetaut |first3=Eric |last4=Escarguel |first4=Gilles |last5=Fluteau |first5=Frédéric |last6=Martineau |first6=François |date=June 15, 2006 |title=Oxygen isotopes from biogenic apatites suggest widespread endothermy in Cretaceous dinosaurs |journal=Earth and Planetary Science Letters |volume=246 |issue=1 |pages=41–54 |doi=10.1016/j.epsl.2006.04.018 |bibcode=2006E&PSL.246...41A |s2cid=55100956 |jstor=41125672 }}

Theropods such as Carcharodontosaurus, Allosaurus, and Acrocanthosaurus have enlarged lacrimal crests, whose purpose is unknown. Paleontologist Daniel Chure hypothesized that these crests were used for "head-butting" between individuals, but how durable they are has not been studied.{{Cite journal |last=Chure |first=Daniel |date=2000 |title=On the orbit of theropod dinosaurs. |url=https://www.researchgate.net/publication/228550944 |journal=Gaia |volume=15 |pages=233–240}}

{{multiple image

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| image1 = Carcharodontosaurus saharicus theropod dinosaur (Kem Kem beds, Upper Cretaceous; Gara es Sbaa, Kem Kem region, southeastern Morocco) 3 (15375691822).jpg

| alt1 = Skull of C. saharicus

| image2 = Carcharodontosaurus saharicus theropod dinosaur (Kem Kem beds, Upper Cretaceous; Gara es Sbaa, Kem Kem region, southeastern Morocco) 2 (15352983706).jpg

| alt2 =

| footer = Skull of C. saharicus showing its elongated, thin rostrum and limited degree of binocular vision

}}

A 2006 study by biologist Kent Stevens analyzed the binocular vision capabilities of the allosauroids Carcharodontosaurus and Allosaurus as well as several coelurosaurs including Tyrannosaurus and Stenonychosaurus. By applying modified perimetry to models of these dinosaurs' heads, Stevens deduced that the binocular vision of Carcharodontosaurus was limited, a side effect of its large, elongated rostrum. Its greatest degree of binocular vision was at higher elevations, suggesting that Carcharodontosaurus may have habitually held its head at a downward 40° angle with its eyes facing up accordingly to achieve maximum binocular vision. The range of vision seen in these allosauroids is comparable to that of crocodiles, suggesting that they were ambush predators. They likely sensed prey via motion parallax between prey and background, with a narrow binocular field of vision helping predators judge prey distances and time attacks.{{Cite journal |last=Stevens |first=Kent A. |date=June 12, 2006 |title=Binocular vision in theropod dinosaurs |journal=Journal of Vertebrate Paleontology |volume=26 |issue=2 |pages=321–330 |doi=10.1671/0272-4634(2006)26[321:BVITD]2.0.CO;2 |s2cid=85694979 |jstor=4524572 }}

{{Main|Theropod paleopathology}}

The neotype skull of C. saharicus is one of many allosauroid individuals to preserve pathologies, with signs of biting, infection, and breaks observed in Allosaurus and Acrocanthosaurus among others.{{Cite journal |last1=Chinsamy |first1=Anusuya |last2=Tumarkin-Deratzian |first2=Allison |date=2009 |title=Pathologic Bone Tissues in a Turkey Vulture and a Nonavian Dinosaur: Implications for Interpreting Endosteal Bone and Radial Fibrolamellar Bone in Fossil Dinosaurs |journal=The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology |volume=292 |issue=9 |pages=1478–1484 |doi=10.1002/ar.20991 |pmid=19711479 |s2cid=41596233 |doi-access=free }}{{cite book |last1=Molnar |first1=R. E.|editor1-last=Carpenter |editor1-first=Kenneth |editor2-last=Skrepnick |editor2-first=Michael William |editor3-last=Tanke |editor3-first=Darren H. |title=Mesozoic Vertebrate Life |date=2001 |publisher=Indiana Univ. Press |isbn=978-0-253-33907-2 |pages=337–363 |chapter=Theropod Paleopathology: A Literature Survey |oclc=248649755 }} This skull bears a circular puncture wound in the nasal and "an abnormal projection of bone on the antorbital rim". A later study theorized that this was the result of craniofacial bites.{{cite book |last1=Rothschild |first1=Bruce |title=The Carnivorous Dinosaurs |last2=Tanke |first2=Darren |publisher=Indiana University Press |year=2005 |isbn=978-0-253-34539-4 |edition=1st |location=Indianapolis |page=351 |chapter=Theropod paleopathology: state-of-the-art review }}

Fossils of Carcharodontosaurus are known from several Cretaceous-age sites across North Africa, similar to the ranges of Spinosaurus and Deltadromeus. North Africa during this period bordered the Tethys Sea, which transformed the region into a mangrove-dominated coastal environment filled with vast tidal flats and waterways.{{Cite journal |last1=Hamed |first1=Younes |last2=Al-Gamal |first2=Samir Anwar |last3=Ali |first3=Wassim |last4=Nahid |first4=Abederazzak |last5=Dhia |first5=Hamed Ben |date=March 1, 2014 |title=Palaeoenvironments of the Continental Intercalaire fossil from the Late Cretaceous (Barremian-Albian) in North Africa: a case study of southern Tunisia |journal=Arabian Journal of Geosciences |volume=7 |issue=3 |pages=1165–1177 |doi=10.1007/s12517-012-0804-2 |bibcode=2014ArJG....7.1165H |s2cid=128755145 }}{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=185}} Isotopes from Carcharodontosaurus and Spinosaurus fossils suggest that the Kem Kem Beds witnessed a temporary monsoon season rather than constant rainfall, similar to modern conditions present in sub-tropical and tropical environments in Southeast Asia and Sub-Saharan Africa.{{cite journal |last1=Amiot |first1=Romain |last2=Buffetaut |first2=Eric |last3=Lécuyer |first3=Christophe |last4=Wang |first4=Xu |last5=Boudad |first5=Larbi |last6=Ding |first6=Zhongli |last7=Fourel |first7=François |last8=Hutt |first8=Steven |last9=Martineau |first9=François |last10=Medeiros |first10=Manuel Alfredo |last11=Mo |first11=Jinyou |last12=Simon |first12=Laurent |last13=Suteethorn |first13=Varavudh |last14=Sweetman |first14=Steven |last15=Tong |first15=Haiyan |last16=Zhang |first16=Fusong |last17=Zhou |first17=Zhonghe |title=Oxygen isotope evidence for semi-aquatic habits among spinosaurid theropods |journal=Geology |date=February 2010 |volume=38 |issue=2 |pages=139–142 |doi=10.1130/G30402.1 |bibcode=2010Geo....38..139A }} These riverine deposits bore large fishes, including the sawskate Onchopristis, coelacanth Mawsonia, and bichir Bawitius.{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=184}} This led to an abundance of piscivorous crocodyliformes evolving in response, such as the giant stomatosuchid Stomatosuchus in Egypt and the genera Elosuchus, Laganosuchus, and Aegisuchus from Morocco.{{Cite journal |last1=Holliday |first1=Casey M. |last2=Gardner |first2=Nicholas M. |date=January 31, 2012 |title=A New Eusuchian Crocodyliform with Novel Cranial Integument and Its Significance for the Origin and Evolution of Crocodylia |journal=PLOS ONE |volume=7 |issue=1 |pages=e30471 |doi=10.1371/journal.pone.0030471 |pmc=3269432 |pmid=22303441 |bibcode=2012PLoSO...730471H |doi-access=free }}{{sfn|Ibrahim|Sereno|Varricchio|Martill|2020|p=180, 189}} Morocco also bore an abundance of pterosaurs like Siroccopteryx and Nicorhynchus.{{Cite journal |last1=Ibrahim |first1=Nizar |last2=Unwin |first2=David M. |last3=Martill |first3=David M. |last4=Baidder |first4=Lahssen |last5=Zouhri |first5=Samir |date=May 26, 2010 |title=A New Pterosaur (Pterodactyloidea: Azhdarchidae) from the Upper Cretaceous of Morocco |journal=PLOS ONE |volume=5 |issue=5 |pages=e10875 |doi=10.1371/journal.pone.0010875 |pmc=2877115 |pmid=20520782 |bibcode=2010PLoSO...510875I |doi-access=free }}{{cite journal |author1=Borja Holgado |author2=Rodrigo V. Pêgas |year=2020 |title=A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae |journal=Acta Palaeontologica Polonica |volume=65 |issue=4 |pages=743–761 |doi=10.4202/app.00751.2020 |s2cid=222075296|doi-access=free }}

The composition of the dinosaur fauna of these sites is an anomaly, as there are fewer herbivorous dinosaur species relative to carnivorous dinosaurs than usual. This indicates that there was niche partitioning between the different theropod clades, with spinosaurids consuming fish while other groups hunted herbivorous dinosaurs.{{Cite journal |last1=Ibrahim |first1=N |last2=Dal Sasso |first2=C |last3=Maganuco |first3=S |last4=Fabbri |first4=M |last5=Martill |first5=D |last6=Gorscak |first6=E |last7=Lamanna |first7=M |date=2016 |title=Evidence of a derived titanosaurian (Dinosauria, Sauropoda) in the 'Kem Kem beds' of Morocco, with comments on sauropod paleoecology in the Cretaceous of Africa |url=https://econtent.unm.edu/digital/collection/bulletins/id/5964 |journal=Cretaceous Period: Biotic Diversity and Biogeography. New Mexico Museum of Natural History and Science Bulletin |volume=71 |pages=149–159 }} Isotopic evidence supports this, which found greater quantities of sizable, terrestrial animals in the diets of carcharodontosaurids and ceratosaurs from both the Kem Kem Beds and Elrhaz Formation.{{Cite journal |last1=Hassler |first1=A. |last2=Martin |first2=J. E. |last3=Amiot |first3=R. |last4=Tacail |first4=T. |last5=Godet |first5=F. Arnaud |last6=Allain |first6=R. |last7=Balter |first7=V. |date=April 11, 2018 |title=Calcium isotopes offer clues on resource partitioning among Cretaceous predatory dinosaurs |journal=Proceedings of the Royal Society B: Biological Sciences |volume=285 |issue=1876 |pages=20180197 |doi=10.1098/rspb.2018.0197 |pmc=5904318 |pmid=29643213 }} It also coexisted with the sauropod Rebbachisaurus which is found in the Kem Kem Beds.{{Cite journal |last1=Wilson |first1=Jeffrey A. |last2=Allain |first2=Ronan |date=July 4, 2015 |title=Osteology of Rebbachisaurus garasbae Lavocat, 1954, a diplodocoid (Dinosauria, Sauropoda) from the early Late Cretaceous–aged Kem Kem beds of southeastern Morocco |journal=Journal of Vertebrate Paleontology |volume=35 |issue=4 |pages=e1000701 |doi=10.1080/02724634.2014.1000701 |bibcode=2015JVPal..35E0701W |s2cid=129846042 }} Carcharodontosaurids are represented by C. saharicus and Sauroniops in the Kem Kem Beds, Tameryraptor in the Bahariya Formation, Eocarcharia and potentially Carcharodontosaurus in the Elrhaz Formation, and C. iguidensis in the Echkar Formation.

{{reflist}}

{{refbegin|40em}}

• {{cite journal|last=Stromer|first=Ernst|year=1934|title=Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 13. Dinosauria|journal=Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung|series=Neue Folge|language=de|volume=22|pages=1–79|url=https://www.zobodat.at/pdf/Abhandlungen-Akademie-Bayern_NF_22_0001-0079.pdf}}
• {{cite book |last1=Taquet |first1=Philippe |title=Géologie et paléontologie du gisement de Gadoufaoua (aptien du Niger) |date=1976 |publisher=Éditions du Centre national de la recherche scientifique |isbn=978-2-222-02018-9 |language=fr }}
• {{Cite book|first1=William|last1=Nothdurft|first2=Josh|last2=Smith|title=The Lost Dinosaurs of Egypt|year=2002|publisher=Random House Publishing Group|isbn=978-1-58836-117-2|place=New York|url=https://books.google.com/books?id=Axu-z-FXyboC&pg=PT28}}
• {{Cite journal|last1=Ibrahim|first1=Nizar|last2=Sereno|first2=Paul C.|last3=Varricchio|first3=David J.|last4=Martill|first4=David M.|last5=Dutheil|first5=Didier B.|last6=Unwin|first6=David M.|last7=Baidder|first7=Lahssen|last8=Larsson|first8=Hans C. E.|last9=Zouhri|first9=Samir|last10=Kaoukaya|first10=Abdelhadi|year=2020|title=Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco|journal=ZooKeys|issue=928|pages=1–216|doi=10.3897/zookeys.928.47517 |pmc=7188693|pmid=32362741|bibcode=2020ZooK..928....1I|doi-access=free}}

{{refend}}

{{Theropoda|A.}}

{{Taxonbar|from1=Q14431|from2=Q2313178|from3=Q12899173}}

Category:Carcharodontosauridae

Category:Dinosaur genera

Category:Cenomanian dinosaurs

Category:Fossil taxa described in 1931

Category:Taxa named by Ernst Stromer

Category:Dinosaurs of Morocco

Category:Taxa with lost type specimens