Editing Cheetah (section)

==Genetics==
The [[Diploidy|diploid]] number of [[chromosome]]s in the cheetah is 38, the same as in most other felids.<ref name=Geptner1972/> The cheetah was the first felid observed to have unusually low genetic variability among individuals,<ref name="bcw4">{{cite book |first1=M. |last1=Culver |first2=C. |last2=Driscoll |first3=E. |last3=Eizirik |first4=G. |last4=Spong |name-list-style=amp |chapter=Genetic applications in wild felids |pages=107–123 |year=2010 |chapter-url=https://www.researchgate.net/publication/308022395 |url=https://books.google.com/books?id=Z6USDAAAQBAJ&pg=PA115 |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=28 March 2023 |archive-url=https://web.archive.org/web/20230328013747/https://books.google.com/books?id=Z6USDAAAQBAJ&pg=PA115 |url-status=live}}</ref> which has led to poor breeding in captivity, increased [[spermatozoa]]l defects, high juvenile mortality and increased susceptibility to diseases and infections.<ref name="o'brien1985">{{cite journal |last1=O'Brien |first1=S. J. |last2=Roelke |first2=M. |last3=Marker |first3=L. |last4=Newman |first4=A. |last5=Winkler |first5=C. |last6=Meltzer |first6=D. |last7=Colly |first7=L. |last8=Evermann |first8=J. |last9=Bush |first9=M. |last10=Wildt |first10=D. E. |title=Genetic basis for species vulnerability in the cheetah|name-list-style=amp |journal=Science |year=1985 |volume=227 |issue=4693 |pages=1428–1434 |doi=10.1126/science.2983425 |pmid=2983425 |bibcode=1985Sci...227.1428O}}</ref><ref name=obrien2017>{{cite journal |last1=O'Brien |first1=S. J |last2=Johnson |first2=W. E |last3=Driscoll |first3=C. A |last4=Dobrynin |first4=P. |last5=Marker |first5=L. |name-list-style=amp |title=Conservation genetics of the cheetah: lessons learned and new opportunities |journal=[[Journal of Heredity]] |date=2017 |volume=108 |issue=6 |pages=671–677 |doi=10.1093/jhered/esx047 |pmid=28821181 |pmc=5892392}}</ref> A prominent instance was the deadly [[feline coronavirus]] outbreak in the cheetah breeding facility at [[Wildlife Safari]] in [[Winston, Oregon]] in 1983 which had a mortality rate of 60%, higher than that recorded for previous [[epizootic]]s of [[feline infectious peritonitis]] in any felid.<ref>{{cite journal |last1=Heeney |first1=J. L. |last2=Evermann |first2=J. F. |last3=McKeirnan |first3=A. J. |last4=Marker-Kraus |first4=L. |last5=Roelke |first5=M. E. |last6=Bush |first6=M. |last7=Wildt |first7=D. E. |last8=Meltzer |first8=D. G. |last9=Colly |first9=L. |last10=Lukas |first10=J. |name-list-style=amp |title=Prevalence and implications of feline coronavirus infections of captive and free-ranging cheetahs (''Acinonyx jubatus'') |journal=[[Journal of Virology]] |date=1990 |volume=64 |issue=5 |pages=1964–1972 |doi=10.1128/JVI.64.5.1964-1972.1990 |pmid=2157864 |pmc=249350}}</ref> The remarkable homogeneity in cheetah genes has been demonstrated by experiments involving the [[major histocompatibility complex]] (MHC); unless the MHC genes are highly homogeneous in a population, [[Skin grafting|skin grafts]] exchanged between a pair of unrelated individuals would be rejected. Skin grafts exchanged between unrelated cheetahs are accepted well and heal, as if their genetic makeup were the same.<ref name=yuhki>{{cite journal |last1=Yuhki |first1=N. |last2=O'Brien |first2=S. J. |name-list-style=amp |title=DNA variation of the mammalian major histocompatibility complex reflects genomic diversity and population history |journal=PNAS |year=1990 |volume=87 |issue=2 |pages=836–840 |pmid=1967831 |pmc=53361 |doi=10.1073/pnas.87.2.836 |bibcode=1990PNAS...87..836Y|doi-access=free}}</ref><ref name=tears>{{cite book |last1=O'Brien |first1=S. J. |title=Tears of the Cheetah: the Genetic Secrets of our Animal Ancestors |date=2003 |publisher=Thomas Dunne Books |location=New York |isbn=978-0-312-33900-5 |pages=15–34 |chapter=Tears of the cheetah |chapter-url=https://books.google.com/books?id=S9iLCgAAQBAJ&pg=PA15 |access-date=30 April 2020 |archive-date=28 March 2023 |archive-url=https://web.archive.org/web/20230328013712/https://books.google.com/books?id=S9iLCgAAQBAJ&pg=PA15 |url-status=live}}</ref>

The low genetic diversity is thought to have been created by two [[population bottleneck]]s from about 100,000 years and about 12,000 years ago, respectively.<ref name="dobrynin" /> The resultant level of genetic variation is around 0.1–4% of average living species, lower than that of [[Tasmanian devil]]s, [[Virunga gorilla]]s, [[Amur tiger]]s, and even highly inbred domestic cats and dogs.<ref name=obrien2017/>

Selective retention of gene variants ([[Gene duplication|duplication]]) has been found in 10 gene candidates to explain energetics and anabolism related to muscle specialization in cheetahs:
* Regulation of muscle contraction (Five genes: ''ADORA1'', ''ADRA1B'', ''CACNA1C'', ''RGS2'', ''SCN5A'').
* Physiological stress response (Two genes: ''ADORA1'', ''TAOK2''). 
* Negative regulation of catabolic process (Four genes: ''APOC3'', ''SUFU'', ''DDIT4'', ''PPARA'').<ref name="dobrynin" /><ref name="obrien2017" />

Potentially harmful mutations has been found in a gene related to spermatogenesis (''AKAP4''). This could explain the high proportion of abnormal sperma in male cheetahs and poor reproductive success in the species.<ref name="dobrynin" /><ref name="obrien2017" />

===King cheetah===
[[File:King cheetah, De Wildt Cheetah Research Centre (South Africa).jpg|thumb|upright|alt=A king cheetah in profile|King cheetah]]

The king cheetah is a variety of cheetah with a rare [[mutation]] for cream-coloured fur marked with large, blotchy spots and three dark, wide stripes extending from the neck to the tail.<ref name=thompson>{{cite book |last1=Thompson |first1=S. E. |title=Built for Speed: The Extraordinary, Enigmatic Cheetah |year=1998 |publisher=Lerner Publications Co |location=Minneapolis |isbn=978-0-8225-2854-8 |chapter=Cheetahs in a bottleneck |chapter-url={{Google Books |plainurl=yes |page=61 |id=wNV5xsM1GVYC}} |pages=61–75 |url=https://archive.org/details/builtforspeedext00thom/page/61}}</ref> In [[Manicaland]], Zimbabwe, it was known as ''nsuifisi'' and thought to be a [[Hybrid (biology)|cross]] between a leopard and a [[hyena]].<ref name=bottriell/> In 1926, Major A.&nbsp;Cooper wrote about a cheetah-like animal he had shot near modern-day [[Harare]], with fur as thick as that of a [[snow leopard]] and spots that merged to form stripes. He suggested it could be a cross between a leopard and a cheetah. As more such individuals were observed it was seen that they had non-retractable claws like the cheetah.<ref name=pocock/><ref name=heuvelmans>{{cite book |last1=Heuvelmans |first1=B. |title=On the Track of Unknown Animals |year=1995 |publisher=[[Routledge]] |location=Abingdon |isbn=978-1-315-82885-5 |pages=495–502 |edition=3rd, revised |chapter=Mngwa, the strange one |chapter-url=https://books.google.com/books?id=u64ABAAAQBAJ&pg=PA495 |access-date=20 December 2019 |archive-date=28 March 2023 |archive-url=https://web.archive.org/web/20230328013747/https://books.google.com/books?id=u64ABAAAQBAJ&pg=PA495 |url-status=live}}</ref>

In 1927, Pocock described these individuals as a new species by the name of ''Acinonyx rex'' ("king cheetah").<ref name="pocock">{{cite journal |last1=Pocock |first1=R. I. |author-link=Reginald Innes Pocock |title=Description of a new species of cheetah (''Acinonyx'') |journal=[[Proceedings of the Zoological Society of London]] |year=1927 |volume=97 |issue=1 |pages=245–252 |doi=10.1111/j.1096-3642.1927.tb02258.x}}</ref> However, in the absence of proof to support his claim, he withdrew his proposal in 1939. [[Abel Chapman]] considered it a [[colour morph]] of the normally spotted cheetah.<ref name=catsg>{{cite web |title=Cheetah—guépard—duma—''Acinonyx jubatus'' |url=http://www.catsg.org/cheetah/01_information/1_2_species-information/species-information.htm#Phylogenetic%20history |publisher=IUCN/SSC Cat Specialist Group |access-date = 6 May 2014 |archive-date = 21 July 2017 |archive-url = https://web.archive.org/web/20170721022535/http://www.catsg.org/cheetah/01_information/1_2_species-information/species-information.htm#Phylogenetic%20history |url-status = live}}</ref> Since 1927, the king cheetah has been reported five more times in the wild in Zimbabwe, Botswana and northern [[Transvaal Province|Transvaal]]; one was photographed in 1975.<ref name="bottriell">{{cite book |last1=Bottriell |first1=L. G. |title=King Cheetah: The Story of the Quest |year=1987 |url=https://books.google.com/books?id=nNcUAAAAIAAJ&pg=frontcover |pages=26; 83–96 |publisher=[[Brill Publishers]] |location=Leiden |isbn=978-90-04-08588-6 |access-date=22 May 2020 |archive-date=28 March 2023 |archive-url=https://web.archive.org/web/20230328013736/https://books.google.com/books?id=nNcUAAAAIAAJ&pg=frontcover |url-status=live}}</ref>

In 1981, two female cheetahs that had mated with a wild male from Transvaal at the [[De Wildt Cheetah and Wildlife Centre]] (South Africa) gave birth to one king cheetah each; subsequently, more king cheetahs were born at the centre.<ref name = catsg/> In 2012, the cause of this coat pattern was found to be a mutation in the gene for [[Transmembrane protein|transmembrane]] [[aminopeptidase]] (Taqpep), the same gene responsible for the striped "mackerel" versus blotchy "classic" pattern seen in [[Cat coat genetics|tabby cats]].<ref>{{cite journal |last1=Aarde |first1=R. J. van |last2=Dyk |first2=A. van |name-list-style=amp |title=Inheritance of the king coat colour pattern in cheetahs ''Acinonyx jubatus'' |journal=Journal of Zoology |date=1986 |volume=209 |issue=4 |pages=573–578 |doi=10.1111/j.1469-7998.1986.tb03612.x}}</ref> The appearance is caused by reinforcement of a [[recessive (genetics)|recessive allele]]; hence if two mating cheetahs are [[heterozygosity|heterozygous]] [[genetic carrier|carriers]] of the mutated allele, a quarter of their offspring can be expected to be king cheetahs.<ref name="wcw">{{cite book |last1=Sunquist |first1=F. |last2=Sunquist |first2=M. |name-list-style=amp |title=Wild Cats of the World |date=2002 |publisher=[[The University of Chicago Press]] |location=Chicago |isbn=978-0-226-77999-7 |pages=19–36 |chapter=Cheetah ''Acinonyx jubatus'' (Schreber, 1776) |chapter-url={{Google Books |plainurl=yes |id=hFbJWMh9-OAC |page=19}} }}</ref>