Editing Cheetah (section)

==Phylogeny and evolution==
{{cladogram|title=
|caption=The ''Puma'' lineage of the family [[Felidae]], depicted along with closely related genera<ref name="bcw2"/>
|1={{clade | style=font-size:90%;line-height:100%;width:475px;
|1={{clade
   |1={{clade
      |1={{clade
         |label1=''Lynx'' lineage
         |1=''[[Lynx]]''
         }}
      |2={{clade
         |1={{clade
            |label1=''Puma'' lineage
            |1={{clade
               |label1=''[[Acinonyx]]''
               |1='''Cheetah''' [[File:Acinonyx jubatus (white background).jpg|60px|alt=Cheetah (Acinonyx jubatus)]]
               |2={{clade
                  |label1=''[[Puma (genus)|Puma]]''
                  |1=[[Cougar]] ''P. concolor'' [[File:Felis concolor - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|50px|alt=Cougar (Puma concolor)]]
                  |label2=''[[Herpailurus]]''
                  |2=[[Jaguarundi]] ''H. yagouaroundi'' [[File:Lydekker_-_Eyra_White_background.jpg|50px|alt=Jaguarundi (Herpailurus yagouaroundi)]]
                  }}
               }}
            }}
         |2={{clade
            |label1=Domestic cat lineage
            |1=''[[Felis]]''
            |label2=Leopard cat lineage
            |2={{clade
               |1=''[[Otocolobus]]''
               |2=''[[Prionailurus]]''
               }}
            }}
         }}
      }}
   }}
}}
}}

The cheetah's closest relatives are the [[cougar]] (''Puma concolor'') and the [[jaguarundi]] (''Herpailurus yagouaroundi'').<ref name="Catsg2017">{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O'Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S. |year=2017 |name-list-style=amp |title=A revised taxonomy of the Felidae: the final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |issue=Special Issue 11 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y |pages=30–31 |access-date=13 May 2018 |archive-date=17 January 2020 |archive-url=https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y |url-status=live}}</ref> Together, these three species form the ''Puma'' lineage, one of the eight lineages of the extant [[felid]]s; the ''Puma'' lineage [[Genetic divergence|genetically diverged]] from the rest 6.7 [[Mya (unit)|mya]]. The [[sister group]] of the ''Puma'' lineage is a [[clade]] of smaller [[Old World]] cats that includes the genera ''[[Felis]]'', ''[[Otocolobus]]'' and ''[[Prionailurus]]''.<ref name="bcw2">{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |chapter=Phylogeny and evolution of cats (Felidae) |name-list-style=amp |year=2010 |pages=59–82 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |access-date=7 January 2020 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live}}</ref>

The oldest cheetah [[fossil]]s, excavated in eastern and southern Africa, date to 3.5–3 mya; the earliest known specimen from South Africa is from the lowermost deposits of the Silberberg Grotto ([[Sterkfontein]]).<ref name=mammal/><ref name=skinner/> Though incomplete, these fossils indicate forms larger but less [[cursorial]] than the modern cheetah.<ref name="marker2">{{cite book |author1=Van Valkenburgh, B. |title=Cheetahs: Biology and Conservation |author2=Pang, B. |author3=Cherin, M. |author4=Rook, L. |date=2018 |publisher=Academic Press |isbn=978-0-12-804088-1 |editor1=Marker, L. |location=London |chapter=The cheetah: evolutionary history and paleoecology |editor2=Boast, L. K. |editor3=Schmidt-Kuentzel, A. |chapter-url=https://books.google.com/books?id=H3rXDgAAQBAJ&pg=PA25 |name-list-style=amp |access-date=26 March 2023 |archive-date=28 March 2023 |archive-url=https://web.archive.org/web/20230328013452/https://books.google.com/books?id=H3rXDgAAQBAJ&pg=PA25 |url-status=live}}</ref> The first occurrence of the modern species ''A. jubatus'' in Africa may come from Cooper's D, a site in South Africa dating back to 1.5 to 1.4 Ma, during the [[Calabrian (stage)|Calabrian]] [[stage (geology)|stage]].<ref>{{Cite journal |last1=O'Regan |first1=Hannah J. |last2=Steininger |first2=Christine |date=30 June 2017 |title=Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora) |url=https://bioone.org/journals/Geodiversitas/volume-39/issue-2/g2017n2a8/Felidae-from-Coopers-Cave-South-Africa-Mammalia-Carnivora/10.5252/g2017n2a8.short |journal=[[Geodiversitas]] |volume=39 |issue=2 |pages=315–332 |doi=10.5252/g2017n2a8 |bibcode=2017Geodv..39..315O |s2cid=53959454 |access-date=28 January 2024 |archive-date=29 January 2024 |archive-url=https://web.archive.org/web/20240129032419/https://bioone.org/journals/Geodiversitas/volume-39/issue-2/g2017n2a8/Felidae-from-Coopers-Cave-South-Africa-Mammalia-Carnivora/10.5252/g2017n2a8.short |url-status=live}}</ref> Fossil remains from Europe are limited to a few [[Middle Pleistocene]] specimens from [[Hundsheim]] (Austria) and Mosbach Sands (Germany).<ref name="hemmer">{{cite journal |last1=Hemmer |first1=H. |last2=Kahlke |first2=R.-D. |last3=Keller |first3=T. |name-list-style=amp |title=Cheetahs in the Middle Pleistocene of Europe: ''Acinonyx pardinensis'' (sensu lato) ''intermedius'' (Thenius, 1954) from the Mosbach Sands (Wiesbaden, Hesse, Germany) |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |date=2008 |volume=249 |issue=3 |pages=345–356 |doi=10.1127/0077-7749/2008/0249-0345|bibcode=2008NJGPA.249..345H }}</ref> Cheetah-like cats are known from as late as 10,000 years ago from the Old World. The [[giant cheetah]] (''A. pardinensis''), significantly larger and slower compared to the modern cheetah, occurred in Eurasia and eastern and southern Africa in the [[Villafranchian]] period roughly 3.8–1.9 mya.<ref name=caro1994/><ref>{{cite journal |last1=Cherin |first1=M. |last2=Iurino |first2=D. A. |last3=Sardella |first3=R. |last4=Rook |first4=L. |name-list-style=amp |title=''Acinonyx pardinensis'' (Carnivora, Felidae) from the Early Pleistocene of Pantalla (Italy): predatory behavior and ecological role of the giant Plio–Pleistocene cheetah |journal=[[Quaternary Science Reviews]] |year=2014 |volume=87 |pages=82–97 |doi=10.1016/j.quascirev.2014.01.004 |bibcode=2014QSRv...87...82C}}</ref> In the Middle Pleistocene a smaller cheetah, ''A. intermedius'', ranged from Europe to China.<ref name=mammal>{{cite journal |last1=Krausman |first1=P. R. |last2=Morales |first2=S. M. |name-list-style=amp |year=2005 |title=''Acinonyx jubatus'' |journal=[[Mammalian Species]] |volume=771 |pages=1–6 |url=http://www.science.smith.edu/departments/biology/VHAYSSEN/msi/pdf/i1545-1410-771-1-1.pdf |doi=10.1644/1545-1410(2005)771[0001:aj]2.0.co;2 |s2cid=198969000 |archive-url=https://web.archive.org/web/20160304045916/http://www.science.smith.edu/departments/biology/VHAYSSEN/msi/pdf/i1545-1410-771-1-1.pdf |archive-date=4 March 2016}}</ref> The modern cheetah appeared in Africa around 1.9 mya; its fossil record is restricted to Africa.<ref name=marker2/>

Extinct North American cheetah-like cats had historically been classified in ''Felis'', ''Puma'' or ''Acinonyx''; two such species, ''F. studeri'' and ''F. trumani'', were considered to be closer to the puma than the cheetah, despite their close similarities to the latter. Noting this, palaeontologist Daniel Adams proposed ''[[Miracinonyx]]'', a new subgenus under ''Acinonyx'', in 1979 for the North American cheetah-like cats;<ref name="adams"/> this was later elevated to genus rank.<ref name="Valkenburgh1990" /> Adams pointed out that North American and Old World cheetah-like cats may have had a common ancestor, and ''Acinonyx'' might have originated in North America instead of Eurasia.<ref name="adams">{{cite journal |last1=Adams |first1=D. B. |s2cid=17951039 |title=The cheetah: native American |journal=[[Science (journal)|Science]] |year=1979 |volume=205 |issue=4411 |pages=1155–1158 |doi=10.1126/science.205.4411.1155 |pmid=17735054 |bibcode=1979Sci...205.1155A}}</ref> However, subsequent research has shown that ''Miracinonyx'' is phylogenetically closer to the cougar than the cheetah;<ref name=sabre>{{cite journal |last1=Barnett |first1=R. |last2=Barnes |first2=I. |last3=Phillips |first3=M. J. |last4=Martin |first4=L. D. |last5=Harington |first5=C. R. |last6=Leonard |first6=J. A. |last7=Cooper |first7=A. |title=Evolution of the extinct sabretooths and the American cheetah-like cat |journal=[[Current Biology]] |year=2005 |volume=15 |issue=15 |pages=R589–R590 |doi=10.1016/j.cub.2005.07.052 |pmid=16085477 |s2cid=17665121 |name-list-style=amp |doi-access=free |bibcode=2005CBio...15.R589B}}</ref> the similarities to cheetahs have been attributed to [[parallel evolution]].<ref name="bcw2"/>

The three species of the ''Puma'' lineage may have had a common ancestor during the [[Miocene]] (roughly 8.25 mya).<ref name=adams/><ref name=johnson>{{cite journal |pmid=9071018 |year=1997 |last1=Johnson |first1=W. E. |last2=O'Brien |first2=S. J. |name-list-style=amp |title=Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes |volume=44 |issue=S1 |pages=S98–S116 |journal=Journal of Molecular Evolution |doi=10.1007/PL00000060 |bibcode=1997JMolE..44S..98J |s2cid=40185850}}</ref> Some suggest that North American cheetahs possibly migrated to Asia via the [[Bering Strait]], then dispersed southward to Africa through Eurasia at least 100,000 years ago;<ref>{{cite journal |last1=Johnson |first1=W. E. |title=The Late Miocene radiation of modern Felidae: a genetic assessment |journal=Science |date=2006 |volume=311 |issue=5757 |pages=73–77 |doi=10.1126/science.1122277 |pmid=16400146 |bibcode=2006Sci...311...73J |s2cid=41672825}}</ref><ref name="dobrynin">{{cite journal |last1=Dobrynin |first1=P. |last2=Liu |first2=S. |last3=Tamazian |first3=G. |last4=Xiong |first4=Z. |last5=Yurchenko |first5=A. A. |last6=Krasheninnikova |first6=K. |last7=Kliver |first7=S. |last8=Schmidt-Küntzel |first8=A. |name-list-style=amp |year=2015 |title=Genomic legacy of the African cheetah, ''Acinonyx jubatus'' |journal=[[Genome Biology]] |volume=16 |page=277 |doi=10.1186/s13059-015-0837-4 |pmc=4676127 |pmid=26653294 |doi-access=free}}</ref><ref>{{cite journal |last1=O'Brien |first1=S. J. |last2=Johnson |first2=W. E. |name-list-style=amp |title=The evolution of cats |journal=[[Scientific American]] |date=2007 |volume=297 |issue=1 |pages=68–75 |doi=10.1038/scientificamerican0707-68 |bibcode=2007SciAm.297a..68O}}</ref> some authors have expressed doubt over the occurrence of cheetah-like cats in North America, and instead suppose the modern cheetah to have evolved from Asian populations that eventually spread to Africa.<ref name=sabre/><ref>{{cite journal |last1=Faurby |first1=S. |last2=Werdelin |first2=L. |last3=Svenning |first3=J. C. |name-list-style=amp |title=The difference between trivial and scientific names: there were never any true cheetahs in North America |journal=Genome Biology |year=2016 |volume=17 |issue=1 |page=89 |doi=10.1186/s13059-016-0943-y |pmid=27150269 |pmc=4858926 |doi-access=free}}</ref> The cheetah is thought to have experienced two [[population bottleneck]]s that greatly decreased the [[genetic variability]] in populations; one occurred about 100,000 years ago that has been correlated to migration from North America to Asia, and the second 10,000–12,000 years ago in Africa, possibly as part of the [[Quaternary extinction event|Late Pleistocene extinction event]].<ref name=dobrynin/><ref name="o'brien1987">{{cite journal |last1=O'Brien |first1=S. J. |last2=Wildt |first2=D. E. |last3=Bush |first3=M. |last4=Caro |first4=T. M. |author4-link=Tim Caro |last5=FitzGibbon |first5=C. |last6=Aggundey |first6=I. |last7=Leakey |first7=R. E. |name-list-style=amp |title=East African cheetahs: evidence for two population bottlenecks? |journal=[[PNAS]] |year=1987 |volume=84 |issue=2 |pages=508–511 |pmid=3467370 |pmc=304238 |doi=10.1073/pnas.84.2.508 |bibcode=1987PNAS...84..508O |doi-access=free}}</ref><ref name="o'brien1993">{{cite journal |last1=Menotti-Raymond |first1=M. |last2=O'Brien |first2=S. J. |name-list-style=amp |title=Dating the genetic bottleneck of the African cheetah |journal=PNAS |year=1993 |volume=90 |issue=8 |pages=3172–3176 |pmid=8475057 |doi=10.1073/pnas.90.8.3172 |pmc=46261 |bibcode=1993PNAS...90.3172M|doi-access=free}}</ref>