{{Short description|Extinct genus of mackerel shark}} {{Use American English|date=April 2022}} {{Use mdy dates|date=April 2022}} {{Split|Carcharodon hastalis|Carcharodon planus|date=September 2025}} {{Automatic taxobox | fossil_range = Rupelian-Calabrian, {{Fossil range|30|1.07|ref=<ref name=Klug>{{citation|author1=Klug, S.|author2=Kriwet, J.|title=Skeletal remains of the Miocene lamniform shark, Cosmopolitodus hastalis, from Peru|journal=Erlanger Geologische Abhandlungen|volume=6|page=99|url=https://www.petit-fichier.fr/2013/01/27/klug-s-kriwet-j-2008-cosmopolitodus-hasta/klug-s-kriwet-j-2008-cosmopolitodus-hasta.pdf}}</ref><ref name=Ebersoleetal>{{cite journal|author1=Ebersole, J.A.|author2=Ebersole, S.M.|author3=Cicimurri, D.J.|title=The occurrence of early Pleistocene marine fish remains from the Gulf Coast of Mobile County, Alabama, USA|journal=Palaeodiversity|year=2017|volume=10|issue=1|pages=97–115|doi=10.18476/pale.v10.a6|s2cid=134476316|doi-access=free}}</ref>}} | image = Isurus hastalis teeth.jpg | image_caption = Tooth series of ''C. hastalis'' | taxon = Cosmopolitodus | authority = Glikman, 1964 | type_species = {{extinct}}''Cosmopolitodus hastalis'' | type_species_authority = Agassiz, 1843 | subdivision_ranks = Other species | subdivision = * {{extinct}}''C. xiphodon''? <small>(Agassiz, 1843)</small> * {{extinct}}'''''C. planus''''' <small>Agassiz, 1856</small> }}
'''''Cosmopolitodus''''' is an extinct genus of mackerel shark that lived between thirty and one million years ago during the late Oligocene to the Early Pleistocene epochs. Its type species is ''Cosmopolitodus hastalis''. In 2021, ''Isurus planus'' was reassigned to the genus, and thus became the second species ''C. planus''.<ref name = Yun>{{cite journal|author=Yun, C.|year=2021|title=A tooth of the extinct lamnid shark, ''Cosmopolitodus planus'' comb. nov. (Chondrichthyes, Elasmobranchii) from the Miocene of Pohang City, South Korea|journal=Acta Palaeontologica Romaniae|volume=18|issue=1|pages=9–16|doi=10.35463/j.apr.2022.01.02|s2cid=242113412|url=https://actapalrom.geo-paleontologica.org/Online_first/Chan_Cosmopolidus_planus.pdf}}</ref> However, some researchers still consider both species of ''Cosmopolitodus'' as species of ''Carcharodon''.<ref>{{Cite journal|last1=Zevallos-Valdivia |first1=L. |last2=Martínez-Pérez |first2=C. |last3=García-Flores |first3=V. |last4=Chávez-Valencia |first4=A. |last5=Botella |first5=H. |title=First record of palaeozoic vertebrates from Peru |year=2023 |journal=Spanish Journal of Palaeontology |volume=38 |issue=1 |pages=95–100 |doi=10.7203/sjp.25691 |s2cid=256585278 |doi-access=free }}</ref> A possible third species, ''C. xiphodon'' was proposed, though it is now placed as junior synonym of ''Carcharodon plicatilis''.
Its teeth can reach lengths up to 3.5 in (7.5 cm) and are found worldwide.<ref name = fossilworks>{{cite web|title=Carcharodon hastalis Agassiz 1843 (white shark)|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=265174|website=Fossilworks|access-date=17 December 2021}}</ref> It is believed to be an ancestor to the modern day great white shark, an argument supported by the transitional species ''Carcharodon hubbelli'',<ref>{{Cite news | url=https://www.bbc.com/news/science-environment-20318175 | title=Great whites 'not from megashark'| work=BBC News| date=2012-11-14| last1=Crumpton| first1=Nick}}</ref><ref>Dana J. Ehret; Bruce J. Macfadden; Douglas S. Jones; Thomas J. Devries; David A. Foster; Rodolfo Salas-Gismondi (2012). "Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru". Palaeontology. 55 (6): 1139–1153. https://doi.org/10.1111/j.1475-4983.2012.01201.x.</ref> but as of 2021, no phylogenetic analyses have been done for proof.<ref name=Yun></ref>
==Taxonomy== ===Etymology=== ''Cosmopolitodus'' is derived from the Ancient Greek κοσμοπολίτης "''kosmopolítēs''" meaning "citizen of the world" and ὀδών "''odṓn''" meaning "tooth". The specific name ''hastalis'' may be derived from the Latin word ''hasta'' meaning "spear". The disputed species ''xiphodon'' is derived from the Ancient Greek ξίφος "''xíphos''" meaning "sword" and ὀδών "''odṓn''" meaning "tooth".
===Taxonomic history=== thumb|left|Fossil teeth of ''C. hastalis'' The taxonomy of ''C. hastalis'', especially the status of its genus, has long been subject to debate. The initial binomial name was first given as ''Oxyrhina hastalis'' and ''Oxyrhina xiphodon'' for the narrow and broad-form variations respectively by Swiss naturalist Louis Agassiz in his 1843 paper ''Recherches sur les poissons fossiles'', although some indications show that he had coined the taxon as early as 1838.<ref name = COLLARETA>{{cite journal|title=A Well Preserved Skeleton of the Fossil Shark Cosmopolitodus Hastalis from the Late Miocene of Peru, Featuring Fish Remains as Fossilized Stomach Contents|issue=1|url=https://riviste.unimi.it/index.php/RIPS/article/view/8005|journal=Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy)|volume=123|doi=10.13130/2039-4942/8005|year=2017|last3=Altamirano-Sierra|first3=Alí|last5=Chacaltana|first5=César|access-date=August 8, 2019|archive-date=June 1, 2022|archive-url=https://web.archive.org/web/20220601172448/https://riviste.unimi.it/index.php/RIPS/article/view/8005|url-status=dead}}</ref> Throughout the early and mid 20th century, different genera and species of other lamniformes would be clumped into the two taxa as the genus ''Oxyrhina'' began to be used as a wastebasket taxon. Sharks previously identified as variations of ''Oxyrhina hastalis'' or ''Oxyrhina xiphodon'' included two species of extinct ancestral makos, ''Isurus desori'' and ''Isurus retroflexus'', the serrated mako (''Carcharomodus escheri''), and the false-toothed mako (''Parotodus benedenii'').<ref name = Hungary>{{cite journal|title=Central Paratethyan shark fauna (Ipolytarnóc, Hungary)|pages=27–40|url=https://www.researchgate.net/publication/228346892|journal=Geologica Carpathica|volume=58|date=February 2007|last1=Kocsis|first1=László}}</ref> Eventually, later studies would begin to show that much of the sharks within the genus and two species are distinct from each other, discarding the genus ''Oxyrhina'' altogether and creating another issue on what new taxa the sharks should be placed in. As of now, the genus is still uncertain and debated.<ref name = COLLARETA></ref>
Shortly after the discarding of the genus ''Oxyrhina'', a review by Holec et al. (1995) placed the species ''hastalis'' and ''xiphodon'' as a species of mako under the genus ''Isurus'', citing the similarities between the teeth of the two and that of modern mako sharks.<ref name=Hungary /> In 2001, a study by Ward & Bonavi (2001) considered ''Isurus hastalis'' and ''Isurus xiphodon'' as conspecific and declared the latter a ''nomen dubium''.<ref name = Ward>{{cite web|title=Additions to and a review of the Miocene Shark and Ray fauna of Malta.|url=https://www.researchgate.net/publication/280316614|website=ResearchGate}}</ref> Although this conclusion is widely accepted, some scientists disagree, with a study by Whitenack and Gottfried (2010) demonstrating geometrically morphological differences between ''I. hastalis'' and ''I. xiphodon''.<ref name = Gottfreid>{{cite journal|title=A morphometric approach for addressing toothbased species delimitation in fossil mako sharks, Isurus (Elasmobranchii: Lamniformes)|journal=Journal of Vertebrate Paleontology|volume=30|pages=17–25|doi=10.1080/02724630903409055|year=2010|last1=Whitenack †|first1=Lisa B.|last2=Gottfried|first2=Michael D.|issue=1 |bibcode=2010JVPal..30...17W |s2cid=85821523|doi-access=free}}</ref> The scientific name ''Isurus hastalis'' is considered as the "traditional view" in the debate regarding the shark's taxonomy.<ref name = COLLARETA /> <div class="noprint">{{cladogram|align=right|style=width:25%;font-size:100%;line-height:100% |caption=Relationship between ''C. hastalis'' and its close relatives. |cladogram= {{clade |1=''Isurus schoutedeni'' |2={{clade |1=''Isurus praecursor'' |2={{clade |1=''Isurus desori'' |2={{clade |1='''''Cosmopolitodus hastalis''''' |2={{clade |1={{clade |label1=Narrow form |1={{clade |1=''Cosmopolitodus planus'' |2=''Carcharomodus escheri'' }}}} |label2= Broad form |2={{clade |1=''Carcharodon hubbelli'' |2=''Carcharodon carcharias'' }}}}}}}}}}}}}} </div> Ward & Bonavi (2001) reexamined the teeth of ''I. hastalis'' and noted a strong morphological similarity between it and the extant great white shark. The study concluded that ''I. hastalis'' is a direct ancestor of the great white and is more related to it than other makos. They moved the species into the genus ''Cosmopolitodus'', which was a move that was first proposed by Glikman (1964) but was long rejected beforehand.<ref name=Ward /> Later discoveries supports Ward & Bonavi's conclusion of its ancestry to the great white. An analysis of a newly discovered ''Carcharodon hubbelli'' by Ehret et al. (2012) further cemented the theory of ''C. hastalis''<nowiki/>'s ancestry to the great white, however, they also proposed that ''Cosmopolitodus'' should be moved to ''Carcharodon'', remarking that the two genera were separated solely due to the lack of serrations in ''C. hastalis'' and the lack of lateral cusplets in ''C. carcharias'' and pointing out examples of Late Miocene ''C. hastalis'' teeth showing basal serrations.<ref name = Ehret /> Cione et al. (2012) also noted a possibility of moving all species in the genus ''Cosmopolitodus'' into the genus ''Carcharodon'' to avoid a possible paraphyly that would occur if one of them, which was traditionally identified as ''C. xiphodon'', is a putative sister species of ''C. carcharias''. However, the study also noted that this would only be done if the putative sister species is conspecific with ''C. hastalis''. The study concluded that there is indeed a putative sister species of ''C. carcharias'' distinct from ''C. hastalis'' and proposed the taxon ''Carcharodon plicatilis'' for it, resolving the paraphyly issue.<ref name = Coine>{{cite journal|title=Oldest record of the Great White Shark (Lamnidae, Carcharodon; Miocene) in the Southern Atlantic|journal=Geobios|volume=45|issue=2|pages=167–172|doi=10.1016/j.geobios.2011.06.002|year=2012|last1=Cione|first1=Alberto Luis|last2=Cabrera|first2=Daniel Alfredo|last3=Barla|first3=María Julia|bibcode=2012Geobi..45..167C |hdl=11336/139163|hdl-access=free}}</ref>
However, Yun argued that the tooth fossil remains of ''Cosmopolitodus'' and the great white "have been documented from the same deposits, hence the former cannot be a chronospecific ancestor of the latter." He also criticized the fact that the "''Cosmopolitodus'' morphotype has never been tested through phylogenetic analyses," and denoted that as of 2021, the argument that the ''Carcharodon'' lineage with narrow, serrated teeth evolved from ''Cosmopolitodus'' with broad, unserrated teeth is uncertain.<ref name=Yun></ref><ref>{{Cite journal |last=Yun |first=Chan-gyu |date=2020-08-07 |title=New example of Cosmopolitodus hastalis (Lamniformes, Lamnidae) from the Miocene of South Korea {{!}} Zoodiversity |url=http://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/92 |journal=Zoodiversity |language=en-US |volume=54 |issue=5 |pages=433–438 |doi=10.15407/zoo2020.05.433|doi-access=free }}</ref>
==Description== thumb|left|''C. hastalis'' jaws ''C. hastalis'' teeth can grow up to {{convert|8.9|cm|in|abbr=on}} in length, suggesting a very large shark. Its body was probably very similar to that of modern great whites. It is also believed to have a cosmopolitan distribution, with ''C. hastalis'' teeth being found worldwide.<ref name=Fossilguy>{{cite web | url=http://www.fossilguy.com/gallery/vert/fish-shark/carcharodon/index.htm | title=Great White Sharks and Prehistoric White Sharks}}</ref> The species is divided into two forms based on tooth morphology, each with a unique evolutionary line. The maximum adult length is estimated between {{convert|5|and|7|m|ft|abbr=on}}.<ref>Patrick L. Jambura; Julia Türtscher; Faviel A. López-Romero; Catalina Pimiento; Jürgen Kriwet (2019). [https://www.researchgate.net/publication/336567772_On_the_origin_of_the_great_white_shark_Carcharodon_carcharias On the origin of the great white shark Carcharodon carcharias.]</ref><ref>{{cite journal|last1=Pimiento|first1=C.|last2=Cantalapiedra|first2=J.L.|last3=Shimada|first3=K.|last4=Field|first4=D.J.|last5=Smaers|first5=J.B.|year=2019|title=Evolutionary pathways toward gigantism in sharks and rays|journal=Evolution|volume=73|issue=3|pages=588–599|doi=10.1111/evo.13680|pmid=30675721|bibcode=2019Evolu..73..588P |s2cid=59224442|issn=1558-5646|url=https://cronfa.swan.ac.uk/Record/cronfa48696/Download/0048696-05022019162842.pdf }}</ref> Smaller individuals were about {{convert|2.6|-|4.5|m|ft}} long.<ref>{{Cite journal|last1=McCormack|first1=Jeremy|last2=Griffiths|first2=Michael L.|last3=Kim|first3=Sora L.|last4=Shimada|first4=Kenshu|last5=Karnes|first5=Molly|last6=Maisch|first6=Harry|last7=Pederzani|first7=Sarah|last8=Bourgon |first8=Nicolas|last9=Jaouen|first9=Klervia|last10=Becker|first10=Martin A.|last11=Jöns|first11=Niels|date=31 May 2022|title=Trophic position of Otodus megalodon and great white sharks through time revealed by zinc isotopes|journal=Nature Communications|language=en|volume=13|issue=1|pages=2980|doi=10.1038/s41467-022-30528-9|pmid=35641494|pmc=9156768 |bibcode=2022NatCo..13.2980M |s2cid=249235478|issn=2041-1723}}</ref>
===Teeth=== ''C. planus'' teeth are somewhat similar to those of other mackerel sharks, especially the extinct species ''C. hastalis'' and ''Isurus desori''. Adult upper teeth are generally 2–4 cm long with an unserrated cutting edge and no lateral cusplets. True to its name, each tooth is "hooked", the point of which is shifted away from the middle axis, in the direction of the corners of the mouth. In this, the teeth of ''C. planus'' and ''I. desori'' differ from ''C. hastalis''. ''I. desori'' teeth are much more heterodontic than ''C. planus'' and their roots are more uneven in thickness and shape, with more narrow builds and rounder lobes, somewhat making the appearance of a heart. But the crowns of the lower teeth of this species are described to be more straight and identical to the lower teeth of ''C. hastalis''.<ref>{{cite book |last=Cocke |first=Joe |date=2002 |title=Fossil Shark Teeth of the World: A collector's guide}}</ref><ref name = ciampaglioricketts>Ciampaglio, C., Ricketts,D. Miocene Madness: Fossil Finder ID and Information Guide. Bakersfield, California. Buena Vista Museum of Natural History.</ref> The hooked teeth, which probably would be efficient for gripping prey, would suggest a diet of smaller and medium-sized animals.
====Broad-form==== {{multiple image |align=right |perrow=2 |total_width=300
|image1=Isurus hastalis labiale.JPG |caption1=A broad form ''C. hastalis'' tooth
|image2=Cosmopolitodus hastalis tooth.jpg |caption2=A narrow form ''C. hastalis'' tooth
}} The broad form is characterized by broad-shaped teeth often described as being identical to that of modern great whites besides the lack of serrations.<ref name=Fossilguy /> Fossil evidence shows that the broad-form is the direct ancestor of the genus ''Carcharodon'', and specimens from the Pisco Formation in Peru show an evolutionary mosaic between them.<ref name = Ehret>{{cite journal|title=Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru|journal=Palaeontology|volume=55|issue=6|pages=1139–1153|doi=10.1111/j.1475-4983.2012.01201.x|year=2012|last1=Ehret|first1=Dana J.|last2=MacFadden|first2=Bruce J.|last3=Jones|first3=Douglas S.|last4=Devries|first4=Thomas J.|last5=Foster|first5=David A.|last6=Salas-Gismondi|first6=Rodolfo|bibcode=2012Palgy..55.1139E |s2cid=128666594|doi-access=free}}</ref>
This form is also commonly labeled by its species synonym ''xiphodon'' to reflect its difference from the narrow-form, although this is scientifically invalid.<ref name=Elasmo>{{cite web|url=http://www.elasmo.com/genera/slides/gw_evo/gw_evo.html?gwa |title=Great White Lineage |publisher=elasmo.com |date= |accessdate=2022-04-20}}</ref>
====Narrow-form==== The narrow form has more slender teeth than its broad-form counterpart, which resemble broadened ''Isurus oxyrhinchus'' teeth. Unlike the broad-form, the narrow form is believed to be the ancestor of two extinct sharks, the hooked-tooth "mako" (''Cosmopolitodus/Isurus planus''), and the serrated "mako" (''Carcharomodus escheri'')<ref name=Fossilguy />
==Paleobiology== ===Hunting behavior=== thumb|Specimen CPI-7899, featuring aggregate of skeletal and dermal fish remains ''C. hastalis'' was a confirmed hunter of marine mammals.<ref>{{Cite journal |last1=Godfrey |first1=Stephen J. |last2=Perez |first2=Victor J. |last3=Jones |first3=Marcus |last4=Chapman |first4=Phillip F. |last5=Spencer |first5=Nathan |last6=Osborne |first6=Jason E. |date=3 July 2025 |title=New light on the trophic ecology of Carcharodon hastalis from teeth embedded in Miocene cetacean vertebrae from Calvert Cliffs in Maryland, USA |url=https://www.researchgate.net/publication/393367603 |journal=Acta Palaeontologica Polonica |volume=70 |issue=2 |pages=329–337 |doi=10.4202/app.01241.2025 |access-date=19 August 2025 |via=ResearchGate}}</ref> It most likely would have been one of the major predators in its ecosystem, preying upon small whales and other mammals. Trace fossils in the form of tooth marks on the bones of a Pliocene dolphin of the species ''Astadelphis gastaldii'' reveal that ''C. hastalis'' attacked its prey from below and behind, much like the modern great white shark does. The deepest bite marks on the dolphin's ribs indicate the shark aimed for the abdomen of its prey to inflict a fatal bite quickly and incapacitate its prey, and that when the dolphin was attacked a second time, it was bitten near the dorsal fin, suggesting that the dolphin rolled over while injured. The size of the bites indicates further that the shark responsible was estimated to be {{convert|4|m|ft|abbr=on}} long.<ref>{{cite web | url=https://www.sciencedaily.com/releases/2010/03/100316142519.htm | title=Jaws -- 4 million BC: How an extinct shark attacked its prey}}</ref>
A well-preserved skeleton of a juvenile ''C. hastalis'' from Peru described in 2017 showcases that the shark also included fish in its diet. Stomach contents preserved with the fossil show this shark also fed on pilchards such as ancient sardines (genus ''Sardinops''), suggesting bony fish featured prominently in the shark's diet while it was young.<ref>{{cite journal | url=https://repositorio.ingemmet.gob.pe/handle/20.500.12544/723 | title=A well preserved skeleton of the fossil shark Cosmopolitodus hastalis from the late Miocene of Peru, featuring fish remains as fossilized stomach contents | journal=Repositorio Institucional Ingemmet | date=March 2017 | last1=Collareta | first1=Alberto | last2=Landini | first2=Walter | last3=Chacaltana Budiel | first3=César Augusto | last4=Valdivia Vera | first4=Waldir | last5=Altamirano Sierra | first5=Ali J. | last6=Urbina Schmitt | first6=Mario | last7=Bianucci | first7=Giovanni }}</ref>
==See also== {{Portal|Sharks}} * List of sharks * List of prehistoric cartilaginous fish
==References== {{Reflist}}
{{Taxonbar|from=Q20686304}}
Category:Lamnidae Category:Prehistoric shark genera Category:Fossil taxa described in 1964 Category:Miocene first appearances Category:Miocene sharks Category:Pliocene sharks