Editing Sand cat (section)

==Taxonomy==
[[File:Sandkat1 (Felis margarita).jpg|thumb|Sand cat at [[Ree Park]] zoo, Denmark]]
''Felis margarita'' was the [[scientific name]] proposed by [[Victor Loche]] in 1858 who first described a sand cat specimen found in the area of "Négonça" in the northern Algerian Sahara.<ref name=Loche1858>{{cite journal|last=Loche|first=V. |title=Description d'une nouvelle espèce de Chat |trans-title=Description of a new species of cat |journal=Revue et Magasin de Zoologie Pure et Appliquée |series=2 |volume=X |year=1858 |pages=49–50 |url=https://archive.org/stream/revueetmagasinde10soci#page/49/mode/1up}}</ref> This [[holotype]] specimen appears to have been lost.<ref name=Haltenorth53>{{cite journal|last1=Haltenorth|first1=T.|author-link=Theodor Haltenorth |title=Lebende arabische Sandkatze (''Felis margarita'' Loche, 1858) |trans-title=Live Arabian sand cat (''Felis margarita'' Loche, 1858) |journal=Säugetierkundliche Mitteilungen| year=1953| volume=1| pages=71–73}}</ref><ref name=Schauenberg>{{cite journal |author=Schauenberg, P. |year=1974 |title=Données nouvelles sur le Chat des sables ''Felis margarita'' Loche, 1858 |trans-title=New data on the Sand Cat ''Felis margarita'' Loche, 1858 |journal=[[Revue suisse de Zoologie]] |volume=81 |issue=4 |pages=949–969 |doi=10.5962/bhl.part.76053 |doi-access=free}}</ref> The species was named after the French General [[Jean Auguste Margueritte]].<ref name=Cole/>

In the 20th century, the following [[zoological specimen]]s of sand cats were described:
*''Eremaelurus thinobius'' was proposed as a [[species]] by [[Sergey Ognev]] in 1926. The specimen had been collected in the eastern Karakum Desert in [[Turkmenistan]].<ref name=Ognev>{{cite journal |last1=Ognev |first1=S. |title=A new genus and species of cat from the Transcaspian region |journal=Annuaire du Musée Zoologique Académie des Sciences USSR |year=1926 |volume=27 |pages=356–362}}</ref> In 1938, [[Reginald Innes Pocock]] also considered it a species, but subordinated it to the [[genus]] ''[[Felis]]'' using the scientific name ''Felis thinobius''.<ref name=Pocock1938>{{cite journal |last1=Pocock |first1=R. I. |title=The Algerian sand cat (''Felis margarita'' Loche) |journal=Proceedings of the Zoological Society of London |year=1938 |volume=B 108 |issue=1 |pages=41–46 |doi=10.1111/j.1096-3642.1938.tb00021.x}}</ref> Later he considered it a sand cat [[subspecies]],<ref name=Pocock1951>{{cite book |last1=Pocock |first1=R. I. |year=1951 |chapter=''Felis margarita'', Loche |pages=139–145 |title=Catalogue of the genus Felis |publisher=British Museum (Natural History) |location=London |chapter-url=https://archive.org/stream/catalogueofgenus00brit#page/140/mode/2up}}</ref> which to date is widely recognised.<ref name=Haltenorth53/><ref name=ems1966>{{cite book |last1=Ellerman |first1=J. R. |last2=Morrison-Scott |first2=T. C. S. |name-list-style=amp |year=1966 |title=Checklist of Palaearctic and Indian mammals 1758 to 1946 |edition=Second |location=London |publisher=British Museum of Natural History |chapter=''Felis margarita'' Loche, 1858 Sand Cat |pages=307–308 |chapter-url=https://archive.org/stream/checklistofindia00elle#page/306/mode/2up}}</ref><ref name=Schauenberg/><ref name=Sliwa2013/><ref name=Cole>{{cite journal |last1=Cole |first1=F. R. |last2=Wilson |first2=D. E. |name-list-style=amp |title=''Felis margarita'' (Carnivora: Felidae) |journal=[[Mammalian Species]] |year=2015 |volume=47 |issue=924 |pages=63–77 |doi=10.1093/mspecies/sev007|doi-access=free|pmc=7149465 }}</ref>
*''F. m. meinertzhageni'' proposed by Pocock in 1938 was a sand cat skin from the Algerian Sahara.<ref name=Pocock1938/>
*''F. m. aïrensis'' proposed by Pocock in 1938 was a female specimen collected in the [[Aïr Mountains]] in 1937.<ref>{{cite journal |last1=Pocock |first1=R. I. |year=1938 |title=A new race of the sand cat (''Felis margarita'') |journal=Annals and Magazine of Natural History |series=Series 11 |volume=1 |issue=4 |pages=472–476 |doi=10.1080/00222933808526790}}</ref>
* ''F. m. scheffeli'' proposed by German zoologist Helmut Hemmer in 1974 was described on the basis of seven sand cats that had been captured alive in [[Pakistan]]'s [[Nushki District|Nushki]] desert.<ref name=Hemmer1974>{{cite journal |last1=Hemmer |first1=H. |year=1974 |title=Studien zur Systematik und Biologie der Sandkatze (''Felis margarita'' Loche, 1858) |trans-title=Studies on the systematics and biology of the sand cat |journal=Zeitschrift des Kölner Zoo |volume=17 |issue=1 |pages=11–20}}</ref>
* ''F. m. harrisoni'' proposed by Hemmer, [[Peter Grubb (zoologist)|Grubb]] and [[Colin Groves|Groves]] in 1976 was described on the basis of a skin and [[skull]] of an adult male sand cat captured in 1967 in [[Umm al Samim]], [[Oman]].<ref name=Hemmer1976>{{cite journal |last1=Hemmer |first1=H. |last2=Grubb |first2=P. |last3=Groves |first3=C. P. |name-list-style=amp |year=1976 |title=Notes on the sand cat, ''Felis margarita'' Loche 1858 |journal=Zeitschrift für Säugetierkunde |issue=41 |pages=286–303 |url=https://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_41_0286-0303.pdf}}</ref>

In 1974, ''F. m. margarita'', ''F. m. thinobia'' and ''F. m. scheffeli'' were temporarily recognized as [[Valid name (zoology)|valid]] [[Taxon|taxa]]. At the time, it was considered possible that sand cats eventually recorded in Afghanistan and [[Iran]] might constitute distinct subspecies.<ref name=Schauenberg/>
In 2005, ''F. m. margarita'', ''F. m. thinobia'', ''F. m. scheffeli'' and ''F. m. harrisoni'' were recognised as valid taxa by [[W. Chris Wozencraft]], who considered ''F. m. meinertzhageni'' and ''F. m. aïrensis'' [[Synonym (taxonomy)|synonyms]] of the [[nominate subspecies]] ''F. m. margarita''.<ref name=msw3/>
The Cat Classification Task Force of the Cat Specialist Group reviewed the existing information and in 2017 recognized only two subspecies based on [[Morphology (biology)|morphological]] differences, namely:<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. |name-list-style=amp |year=2017 |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 |volume=Special Issue 11 |pages=14–15 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y}}</ref>
[[File:Felis margarita harrisoni - Sandkatze.jpg|thumb|An Arabian sand cat at Osnabrück Zoo, Germany]]
* ''Felis margarita margarita'', also called the Saharan sand cat, occurs in North Africa. It is smaller in size with brighter, more yellow fur, with more pronounced markings and 2–6 rings on the tail.<ref name=CatSG2017/>
* ''Felis margarita thinobia'', also called the Turkestan sand cat, Arabian sand cat, and Pakistan sand cat, it occurs in West and Central Asia. It is larger in size with less pronounced markings, a darker, more greyish coat, and only 2–3 rings on the tail.<ref name=CatSG2017/><ref name=Howard-McCombe2019/>

Analysis of [[mitochondrial DNA]] of 47 individuals from across the sand cat's range showed that their [[haplotype]]s differed only by one to three [[base pair]] [[mutation]]s. This low degree of genetic differentiation between the African and Asian sand cats indicates that the [[Sinai Peninsula]] may have been a barrier to [[gene flow]].<ref name=Howard-McCombe2019>{{cite journal |author1=Howard-McCombe, J. |name-list-style=amp |author2=Banfield, L. |author3=Kitchener, A.C. |author4=Al Qahtani, H. |author5=Toosy, A. |author6=Al Qarqas, M. |author7=Craig, M. |author8=Abramov, A.V. |author9=Veron, G. |author10=Brito, J.C. |author11=Azizi, S. |author12=Ghazali, M. |author13=Breton, G. |author14=Sliwa, A. |author15=Kaltwaßer, K. |author16=Hochkirch, A. |author17=Senn, H. |year=2019 |title=A Mitochondrial Phylogeny of the Sand Cat (''Felis margarita'' Loche, 1858) |journal=Journal of Mammalian Evolution |volume=2019 |issue=3 |pages=525–534 |doi=10.1007/s10914-019-09473-w |s2cid=198190254}}</ref>

=== Phylogeny ===
[[Phylogenetic]] analysis of the [[nuclear DNA]] in tissue samples from all Felidae species revealed that the [[evolutionary radiation]] of the Felidae began in Asia in the [[Miocene]] around {{mya|14.45|8.38}}.<ref name=Johnson2006>{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. | author5=Antunes, A. |author6=Teeling, E. |author7=O'Brien, S. J. |name-list-style=amp |year=2006 |doi=10.1126/science.1122277 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |pages=73–77 |pmid=16400146 |issue=5757|url=https://zenodo.org/record/1230866 |bibcode=2006Sci...311...73J|s2cid=41672825 }}</ref><ref name=Werdelin2010>{{cite book |last1=Werdelin |first1=L. |author1-link=Lars Werdelin |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |name-list-style=amp |chapter=Phylogeny and evolution of cats (Felidae) |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}}</ref> Analysis of mitochondrial DNA of all Felidae species indicates a radiation at around {{mya|16.76|6.46}}.<ref name=Li_al2016>{{cite journal |author=Li, G. |author2=Davis, B. W. |author3=Eizirik, E. |name-list-style=amp |author4=Murphy, W. J. |year=2016 |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |volume=26 |issue=1 |pages=1–11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}</ref>
The sand cat is part of an [[evolution]]ary [[Lineage (evolution)|lineage]] that is estimated to have [[Genetic divergence|genetically diverged]] from the [[common ancestor]] of the ''Felis'' species around {{mya|4.44|2.16}}, based on analysis of their nuclear DNA.<ref name=Johnson2006/><ref name=Werdelin2010/> Analysis of their mitochondrial DNA indicates a genetic divergence of the ''Felis'' species at around {{mya|6.52|1.03}}.<ref name=Li_al2016/> Both models agree that the [[jungle cat]] (''F. chaus'') was the first ''Felis'' species that diverged, followed by the [[black-footed cat]] (''F. nigripes'') and then the sand cat.<ref name=Johnson2006/><ref name=Li_al2016/>
It migrated into Africa, possibly during [[Pleistocene glaciation]] events.<ref name=Johnson2006/> [[Migration (ecology)|Migration]] was likely facilitated by extended periods of low sea levels between continents.<ref name=Li_al2016/>

A [[fossil]] jaw and other [[Skeleton|skelet]]al remains of a sand cat were [[Excavation (archaeology)|excavated]] in a [[Late Pleistocene]] cave in El Harhoura located near [[Temara]] in Morocco.<ref>{{cite journal |author=Aouraghe, H. |year=2000 |title=Les carnivores fossiles d'El Harhoura 1, Temara, Maroc |journal=L'Anthropologie |volume=104 |pages=147–171 |doi=10.1016/S0003-5521(00)90007-4 |url=https://www.researchgate.net/publication/248592769}}</ref>

{{clade gallery |align=left |width=500px; |main-caption=Phylogenetic relationships of the sand cat as derived through analysis of
|header1=nuclear DNA:<ref name=Johnson2006/>
|cladogram1={{clade |label1=[[Felidae]]
 |1={{clade
   |label1=[[Felinae]]
   |1={{clade
    |1={{clade
     |1={{clade |label1=''Felis''
       |1={{clade
        |1={{clade
         |1={{clade
           |1={{clade
             |1={{clade
              |1={{clade
               |1=[[Domestic cat]] (''F. catus'')
               |2=[[European wildcat]] (''F. silvestris'') }}
              |2={{clade
               |1=[[African wildcat]] (''F. lybica'')
               |2=[[Chinese mountain cat]] (''F. bieti'') }} }}
             |2='''Sand cat''' }}
          |2=Black-footed cat }}
         |2=Jungle cat }} }}
      |2=other Felinae [[Lineage (genetic)|lineages]] }} }} }}
  |2=[[Pantherinae]] }} }}
|header2=mitochondrial DNA:<ref name=Li_al2016/>
|cladogram2={{clade |label1=Felidae
|1={{clade |label1=Felinae
   |1={{clade
      |1={{clade
         |1={{clade |label1=''Felis''
            |1={{clade
               |1={{clade
                  |1={{clade
                     |1={{clade
                        |1={{clade
                           |1={{clade |1=Domestic cat |2=Chinese mountain cat }}
                           |2={{clade |1=African wildcat |2=European wildcat }} }}
                        |2='''Sand cat''' }}
                  |2=Black-footed cat }}
               |2=Jungle cat }} }} }}
      |2=other Felinae lineages }} }}
   |2=Pantherinae }} }} }}
{{clear}}