Editing Great white shark (section)

=== Fossil ancestry ===
{{cladogram|caption=Phylogenetic relationship between the Great white shark and other sharks based on molecular data conducted by Human ''et al.'' (2006)<ref name=Humanetal2005>{{cite journal |author1=Human, B. A. |author2=Owen, E. P. |author3=Compagno, L. J. V. |author4=Harley, E. H. |title=Testing morphologically based phylogenetic theories within the cartilaginous fishes with molecular data, with special reference to the catshark family (Chondrichthyes; Scyliorhinidae) and the interrelationships within them |journal=Molecular Phylogenetics and Evolution |volume=39 |issue=2006 |year=2006 |pages=384–391 |doi=10.1016/j.ympev.2005.09.009 |pmid=16293425|bibcode=2006MolPE..39..384H }}</ref>
|cladogram={{clade|style=font-size:85%;line-height:75%;width:250px;
|1={{clade
|1=''[[Sand tiger shark|Carcharias taurus]]''<span style="{{MirrorH}}">[[File:Sandtiger shark (Duane Raver).png|50 px]]</span>
|2={{clade
        |1=''[[Basking shark|Cetorhinus maximus]]''
        |2={{clade
            |1={{clade
                 |1=''[[Porbeagle|Lamna nasus]]'' <span style="{{MirrorH}}">[[File:Lamna nasus.jpg|50 px]]</span>
                 |2=''[[Salmon shark|Lamna ditropis]]''
                     }}
            |2={{clade
                |1=''Carcharodon carcharias'' <span style="{{MirrorH}}">[[File:White shark (Duane Raver).png|50 px]]</span>
                |2={{clade
                    |1=''[[Shortfin mako|Isurus oxyrinchus]]''<span style="{{MirrorH}}">[[File:Shortfin mako shark (Duane Raver).png|50 px]]</span>
                    |2=''[[Longfin mako|Isurus paucus]]''
                    }}
                }}
            }}
        }}
    }}
}}
}}
[[Molecular clock]] studies published between 1988 and 2002 determined the closest living relative of the great white to be the mako sharks of the genus ''Isurus'', which diverged some time between 60 and 43 million years ago.<ref name=Martin1996>{{cite book |author=Martin, A. P. |editor1=Klimley, A. P. |editor2=Ainley, D. G. |title=Great White Sharks: The Biology of ''Carcharodon carcharias'' |chapter=Systematics of the Lamnidae and the Origination Time of ''Carcharodon carcharias'' Inferred from the Comparative Analysis of Mitochondrial DNA Sequences |year=1996 |pages=49–53 |publisher=Academic Press |isbn=978-0-12-415031-7 |doi=10.1016/B978-0-12-415031-7.X5000-9}}</ref><ref name=Kent2018>{{cite book |author=Kent, B. W. |chapter=The Cartilaginous Fishes (Chimaeras, Sharks, and Rays) of Calvert Cliffs, Maryland, USA |title=The Geology and Vertebrate Paleontology of Calvert Cliffs, Maryland |series=Smithsonian Contributions to Paleobiology |editor=Godfrey, S. J. |pages=45–157 |year=2018 |volume=100 |issue=100 |publisher=The Smithsonian Institution |doi=10.5479/si.1943-6688.100 |s2cid=134274604 |url=https://smithsonian.figshare.com/articles/book/The_Geology_and_Vertebrate_Paleontology_of_Calvert_Cliffs_Maryland_USA/9761762 |issn=1943-6688 |access-date=9 November 2020 |archive-date=10 November 2020 |archive-url=https://web.archive.org/web/20201110051916/https://smithsonian.figshare.com/articles/book/The_Geology_and_Vertebrate_Paleontology_of_Calvert_Cliffs_Maryland_USA/9761762 |url-status=live}}</ref> Tracing this evolutionary relationship through [[fossil]] evidence, however, remains subject to further [[paleontology|paleontological]] study.<ref name=Kent2018 />

The original hypothesis of the great white shark's origin held that it is a descendant of a lineage of [[Carcharocles|mega-toothed sharks]], and is closely related to the prehistoric [[megalodon]].<ref name=Kent2018 /><ref name=Applegate>{{cite book |author1=Applegate, S. P. |author2=Espinosa-Arrubarrena, L. |chapter=The Fossil History of ''Carcharodon'' and Its Possible Ancestor, ''Cretolamna'': A Study in Tooth Identification |editor1=Klimley, A. P. |editor2=Ainley, D. G. |title=Great White Sharks: The Biology of Carcharodon carcharias |year=1996 |pages=19–36; 49–53 |publisher=Academic Press |isbn=978-0-12-415031-7 |doi=10.1016/B978-0-12-415031-7.X5000-9}}</ref> These sharks were considerably larger in size, with megalodon attaining an estimated length of up to {{cvt|14.2|-|20.3|m|ft}}.<ref name="Shimada2019">{{Cite journal |last=Shimada |first=K. |year=2019 |title=The size of the megatooth shark, ''Otodus megalodon'' (Lamniformes: Otodontidae), revisited |journal=Historical Biology |volume=33 |issue=7 |pages=1–8 |doi=10.1080/08912963.2019.1666840 |s2cid=208570844 |issn=0891-2963}}</ref><ref name="Cooper2020">{{cite journal |first1=J. A. |last1=Cooper |first2=C. |last2=Pimiento |first3=H. G. |last3=Ferrón |first4=M. J. |last4=Benton |year=2020 |title=Body dimensions of the extinct giant shark ''Otodus megalodon'': a 2D reconstruction |journal=Scientific Reports |volume=10 |issue=14596 |page=14596 |doi=10.1038/s41598-020-71387-y |pmid=32883981 |pmc=7471939 |bibcode=2020NatSR..1014596C|doi-access=free}}</ref><ref name="Perezetal2021">{{Cite journal |last1=Perez |first1=Victor |last2=Leder |first2=Ronny |last3=Badaut |first3=Teddy |date=2021 |title=Body length estimation of Neogene macrophagous lamniform sharks (Carcharodon and Otodus) derived from associated fossil dentitions |url=https://palaeo-electronica.org/content/2021/3284-estimating-lamniform-body-size |journal=Palaeontologia Electronica |volume=24 |issue=1 |pages=1–28 |doi=10.26879/1140 |doi-access=free |access-date=4 March 2023 |archive-date=7 March 2023 |archive-url=https://web.archive.org/web/20230307072346/https://palaeo-electronica.org/content/2021/3284-estimating-lamniform-body-size |url-status=live}}</ref> Similarities between the teeth of great white and mega-toothed sharks, such as large triangular shapes, serrated blades, and the presence of dental bands, led the primary evidence of a close evolutionary relationship. As a result, scientists classified the ancient forms under the genus ''Carcharodon''. Although weaknesses in the hypothesis existed, such as uncertainty over exactly which species evolved into the modern great white and multiple gaps in the fossil record, palaeontologists were able to chart the hypothetical lineage back to a 60-million-year-old shark known as ''[[Cretalamna]]'' as the common ancestor of all sharks within the Lamnidae.<ref name=Martin1996 /><ref name=Applegate />

[[File:Evolution of the great white shark.jpg|thumb|upright=1.8|left|Illustrated evolution from ''C. hastalis'' to ''C. carcharias'']]
However, it is now understood that the great white shark holds closer ties to the mako sharks and is descended from a separate lineage as a [[chronospecies]] unrelated to the mega-toothed sharks.<ref name=Kent2018 /> This was proven with the discovery of a [[transitional species]] that connected the great white to an unserrated shark known as ''[[Carcharodon hastalis]]''.<ref name=Ehret2012>{{cite journal |author1=Ehret, D. A. |author2=MacFadden, B. J. |author3=Jones, D. |author4=DeVries, T. J. |author5=Foster, D. A. |author6=Salas-Gismondi, R. |title=Origin of the White Shark ''Carcharodon'' (Lamniformes: Lamnidae), Based on Recalibration of the Late Neogene, Pisco Formation of Peru |journal=Palaeontology |volume=55 |issue=6 |pages=1139–1153 |year=2012 |doi=10.1111/j.1475-4983.2012.01201.x |s2cid=128666594 |doi-access=free}}</ref><ref name=Nyberg>{{cite journal |author1=Nyberg, K. G. |author2=Ciampaglio, C. N. |author3=Wray, G. A. |title=Tracing the Ancestry of the Great White Shark, ''Carcharodon carcharias'', Using Morphometric Analyses of Fossil Teeth |journal=Journal of Vertebrate Paleontology |volume=26 |issue=4 |year=2006 |doi=10.1671/0272-4634(2006)26[806:ttaotg]2.0.co;2 |pages=806–814 |s2cid=53640614}}</ref> This transitional species, which was named ''[[Carcharodon hubbelli]]'' in 2012, demonstrated a mosaic of evolutionary transitions between the great white and ''C. hastalis'', namely the gradual appearance of serrations,<ref name=Ehret2012 /> in a span of between 8 and 5 million years ago.<ref name=Boesse2019>{{cite journal |author1=Boessenecker, R. W. |author2=Ehret, D. J. |author3=Long, D. J. |author4=Churchill, M. |author5=Martin, E. |author6=Boessenecker, S. J. |year=2019 |title=The Early Pliocene extinction of the mega-toothed shark ''Otodus megalodon'': a view from the eastern North Pacific |journal=PeerJ |volume=7 |pages=e6088 |doi=10.7717/peerj.6088 |pmid=30783558 |pmc=6377595 |doi-access=free}}</ref> The progression of ''C. hubbelli'' characterized shifting diets and niches; by 6.5 million years ago, the serrations were developed enough for ''C. hubbelli'' to handle marine mammals.<ref name=Ehret2012 /> Although both the great white and ''C. hastalis'' were known worldwide,<ref name=Kent2018 /> ''C. hubbelli'' is primarily found in [[California]], [[Peru]], [[Chile]], and surrounding coastal deposits,<ref name=Long2014>{{citation |author1=Long, D. J. |author2=Boessenecker, R. W. |author3=Ehret, D. J. |title=Timing of evolution in the ''Carcharodon'' lineage: Rapid morphological change creates a major shift in a predator's trophic niche |year=2014 |url=https://www.researchgate.net/publication/272745162 |access-date=9 November 2020 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505222759/https://www.researchgate.net/publication/272745162_Timing_of_evolution_in_the_Carcharodon_lineage_Rapid_morphological_change_creates_a_major_shift_in_a_predator's_trophic_niche |url-status=live}}</ref> indicating that the great white had [[Pacific Ocean|Pacific]] origins.<ref name=Ehret2012 /> ''C. hastalis'' continued to thrive alongside the great white until its last appearance around one million years ago<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 |journal=Palaeodiversity |year=2017 |volume=10 |issue=1 |pages=97–115 |doi=10.18476/pale.v10.a6 |s2cid=134476316 |doi-access=free}}</ref> and is believed to have possibly sired a number of additional species, including ''[[Carcharomodus escheri|Carcharodon subserratus]]''<ref name=Kent2018 /><ref name=Ehret2012 /> and ''[[Carcharodon plicatilis]]''.<ref name=Kent2018 />

However, Yun argued that the tooth fossil remains of ''C. hastalis'' and Great White Shark "have been documented from the same deposits, hence the former cannot be a chronospecific ancestor of the latter." He also criticized that the ''C. hastalis'' "morphotype has never been tested through phylogenetic analyses," and denoted that as of 2021, the argument that the modern Carcharodon lineage with narrow, serrated teeth evolved from ''C. hastalis'' with a broad, unserrated teeth is uncertain.<ref name = Yun>{{cite journal |author=Yun, C. |year=2021 |title=A tooth of the extinct lamnid shark, ''Cosmopolitodus planus'' comb. noc. (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 |archive-url=https://web.archive.org/web/20210712005302/https://actapalrom.geo-paleontologica.org/Online_first/Chan_Cosmopolidus_planus.pdf |archive-date=12 July 2021 |url-status=live}}</ref>

Tracing beyond ''C. hastalis'', another prevailing hypothesis proposes that the great white and mako lineages shared a common ancestor in a primitive mako-like species.<ref name=ElasmoFossil>{{cite web |author=Martin, R. A. |title=Fossil History of the White Shark |website=Elasmo Research |url=http://www.elasmo-research.org/education/white_shark/carcharodon.htm |access-date=9 November 2020 |archive-date=27 June 2020 |archive-url=https://web.archive.org/web/20200627223434/http://www.elasmo-research.org/education/white_shark/carcharodon.htm |url-status=live}}</ref> The identity of this ancestor is still debated, but a potential species includes ''[[Isurolamna]] inflata'', which lived between 65 and 55 million years ago. It is hypothesized that the great white and mako lineages split with the rise of two separate descendants, the one representing the great white shark lineage being ''[[Macrorhizodus]] praecursor''.<ref name=ElasmoFossil /><ref name=Trif>{{cite journal |author1=Trif, N. |author2=Ciobanu, R. |author3=Codrea, V. |year=2016 |title=The first record of the giant shark ''Otodus megalodon'' (Agassiz, 1835) from Romania |journal=Brukenthal, Acta Musei |volume=11 |issue=3 |pages=507–526 |url=https://www.researchgate.net/publication/309615167 |access-date=9 November 2020 |archive-date=22 February 2017 |archive-url=https://web.archive.org/web/20170222105957/https://www.researchgate.net/publication/309615167 |url-status=live}}</ref>