View source for Leopardus

{{Short description|Genus of felines native to the Americas}}
{{Automatic taxobox
| name = ''Leopardus''
| fossil_range = [[Pleistocene]]-Recent<ref name=Berta1983/><br />~{{fossil range|2.5|0}}
| image = Leopardus_collage.png
| image_upright = 1.2
| image_caption = ''Leopardus'' species from top-left, clockwise: [[ocelot]] (''L. pardalis''), [[oncilla]] (''L. tigrinus''), [[Pampas cat]] (''L. colocola''), [[kodkod]] (''L. guigna''), [[margay]] (''L. wiedii''), [[Geoffroy's cat]] (''L. geoffroyi'')
| taxon = Leopardus
| authority = [[John Edward Gray|Gray]], 1842
| type_species = ''[[Leopardus pardalis|Leopardus griseus]]''<ref name=MSW3>{{MSW3 Wozencraft |pages=537–540 |id=14000080 |heading=Genus ''Leopardus''}}</ref>
| type_species_authority = Gray, 1842
| subdivision_ranks = Species
| range_map = Leopardus range.png
| range_map_caption = ''Leopardus'' distribution
| range_map_alt = Leopardus range map
}}

'''''Leopardus''''' is a [[genus]] comprising eight [[species]] of [[Felinae|small cat]]s native to the [[Americas]].<ref name=catsg>{{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 |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 |pages=46–58 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=47}}</ref> This genus is considered the oldest branch of a [[genetic lineage]] of small cats in the Americas whose [[common ancestor]] crossed the [[Bering land bridge]] from Asia to North America in the [[late Miocene]].<ref name=Johnson2006>{{cite journal |last1=Johnson|first1=W. E. |last2=Eizerik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy|first4=W. J. |last5=Antunes|first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=S. J. |name-list-style=amp |date=2006 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=Science |volume=311 |issue=5757 |pages=73–77 |doi=10.1126/science.1122277 |url=https://zenodo.org/record/1230866 |pmid=16400146 |bibcode=2006Sci...311...73J |s2cid=41672825}}</ref>

== Characteristics ==
''Leopardus'' species have spotted [[fur]], with ground colors ranging from pale [[Buff (colour)|buff]], [[ochre]],  [[fulvous]] and [[Tawny (color)|tawny]] to light gray.<ref>{{cite journal |author=Allen J. A. |year=1919 |title=Notes on the synonymy and nomenclature of the smaller spotted cats of tropical America |journal=Bulletin of the American Museum of Natural History |volume=41 |pages=341–419}}</ref> Their small [[ears]] are rounded and white-spotted; their [[rhinarium]] is prominent and naked above, and their [[nostril]]s are widely separated.<ref>{{cite journal |last1=Pocock |first1=R. I. |year=1917 |title=The classification of the existing Felidae |journal=Annals and Magazine of Natural History |series=Series 8 |volume=XX |issue=119 |pages=329–350 |url=https://archive.org/stream/annalsmagazineof8201917lond#page/344/mode/2up |doi=10.1080/00222931709487018}}</ref> They have 36 [[chromosome]]s, whereas other felids have 38.<ref name=Trigo_al2008>{{cite journal |author1=Trigo T. C. |name-list-style=amp |author2=Freitas T. R. O. |author3=Kunzler G. |author4=Cardoso L. |author5=Silva J. C. R. |author6=Johnson, W. E. |author7=O’Brien S. J. |author8=Bonatto S. L. |author9=Eizirik E. |year=2008 |title=Inter-species hybridization among Neotropical cats of the genus ''Leopardus'', and evidence for an introgressive hybrid zone between ''L. geoffroyi'' and ''L. tigrinus'' in southern Brazil |journal=Molecular Ecology |volume=17 |issue=19 |pages=4317–4333 |doi=10.1111/j.1365-294X.2008.03919.x |pmid=18785898 |pmc=6993176|bibcode=2008MolEc..17.4317T }}</ref>

== Taxonomy ==
The [[Generic name (biology)|generic name]] ''Leopardus'' was proposed by [[John Edward Gray]] in 1842, when he described two spotted cat skins from Central America and two from [[India]] in the collection of the [[Natural History Museum, London]].<ref>{{cite journal |last1=Gray |first1=J. E. |year=1842 |title=Descriptions of some new genera and fifty unrecorded species of Mammalia |journal=Annals and Magazine of Natural History |volume=10 |issue=65 |pages=255–267 |url=https://archive.org/details/annalsmagazineof10lond/page/260 |doi=10.1080/03745484209445232}}</ref>
Several genera were proposed in the 19th and early 20th centuries for small spotted cats in the Americas, including:
*''Dendrailurus'', ''Lynchailurus'', ''Noctifelis'', ''Oncifelis'' and ''Oncoïdes'' by [[Nikolai Severtzov]] in 1858;<ref>{{cite journal |last1=Severtzow |first1=M. N. |year=1858 |title=Notice sur la classification multisériale des Carnivores, spécialement des Félidés, et les études de zoologie générale qui s'y rattachent |journal=Revue et Magasin de Zoologie Pure et Appliquée |volume=X |pages=385–396 |url=https://archive.org/stream/revueetmagasinde10soci#page/386/mode/2up}}</ref>
*''Margay'', ''Pajeros'', ''Pardalina'' and ''Pardalis'' by Gray in 1867;<ref>{{cite journal |last1=Gray |first1=J. E. |year=1867 |title=Notes on the skulls of the Cats (Felidae) |journal=Proceedings of the Zoological Society of London |issue=March |pages=258–277 |url=https://archive.org/details/proceedingsofzoo1867zool/page/266/mode/2up}}</ref>
*''Oncilla'' by [[Joel Asaph Allen]] in 1919;<ref>{{cite journal |author=Allen, J. A. |year=1919 |title=Severtzow's classification of the Felidae |journal=Bulletin of the American Museum of Natural History |volume=41 |issue=6 |pages=335–342 |url=http://digitallibrary.amnh.org/bitstream/handle/2246/1795//v2/dspace/ingest/pdfSource/bul/B041a06.pdf?sequence=1&isAllowed=y}}</ref>
*''Oreailurus'' by [[Ángel Cabrera (naturalist)|Ángel Cabrera]] in 1940;<ref>{{cite journal |author=Cabrera, Á. |year=1940 |title=Notas sobre Carnívoros sudamericanos |journal=Notas del Museo de la Plata |volume=V |issue=29 |pages=1–22 |url=http://naturalis.fcnym.unlp.edu.ar/repositorio/_documentos/sipcyt/bfa001212.pdf}}</ref>
*''Colocolo'' by [[Reginald Innes Pocock]] in 1941.<ref>{{cite journal |author=Pocock, R.I. |year=1941 |title=The Examples of the Colocolo and of the Pampas Cat in the British Museum |journal=The Annals and Magazine of Natural History |volume=7 |issue=39 |pages=257–274 |doi=10.1080/03745481.1941.9727931 }}</ref>
Analysis of skull [[Morphology (biology)|morphology]] of these [[taxa]] revealed close similarities in their [[base of skull]]s and [[nasal bone]]s, their [[masticatory muscles]], and dentition.<ref>{{cite book |author=Salles, L. O. |year=1992 |title=Felid phylogenetics: extant taxa and skull morphology (Felidae, Aeluroidea) |series=American Museum Novitates |volume=3047 |publisher=American Museum of Natural History |location=New York |url=http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/5011/N3047.pdf?sequence=1}}</ref>
[[Phylogenetic]] analysis of tissue samples of these taxa and their ability to [[Hybrid (biology)|hybridise]] support the notion that they are members of the same genus.<ref name=Johnson2006/><ref name=Trigo_al2008/>
The following eight extant ''Leopardus'' species have commonly been recognized as [[Valid name (zoology)|valid]] taxa since 2017:<ref name=catsg/>

{| class="wikitable sortable"
|-
! Name !! [[IUCN Red List]] status and distribution
|-- style="vertical-align: top;"
| [[Ocelot]] ''L. pardalis'' {{small|([[Carl Linnaeus|Linnaeus]], 1758)}}<ref>{{cite book |last1=Linnaeus |first1=C. |year=1758 |title=Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |location=Holmiae |publisher=Laurentius Salvius |page=42 |chapter=Felis pardalis |volume=I |edition=Tenth |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}}</ref><br /> [[File:Ocelot (Leopardus pardalis)-8.jpg|frameless]] 
||{{IUCN status|LC}}<ref>{{cite iucn |author1=Paviolo, A. |author2=Crawshaw, P. |author3=Caso, A. |author4=de Oliveira, T. |author5=Lopez-Gonzalez, C.A. |author6=Kelly, M. |author7=De Angelo, C. |author8=Payan, E. |name-list-style=amp |year=2015 |title=''Leopardus pardalis'' |errata=2016 |page=e.T11509A97212355}}</ref><br />[[File:Ocelot_distribution.jpg|frameless]]
|-- style="vertical-align: top;"
| [[Oncilla]] ''L. tigrinus'' {{small|([[Johann Christian Daniel von Schreber|Schreber]], 1775)}}<ref>{{cite book |author=Schreber, J. C. D. |year=1778 |chapter=Die Maragua |chapter-url=https://archive.org/stream/SaYugthiereAbbiIIISchr#page/396/mode/1up |pages=396–397 |title=Die Säugethiere in Abbildungen nach der Natur, mit Beschreibungen |location=Erlangen |publisher=Wolfgang Walther}}</ref><br />[[File:Leopardus tigrinus - Parc des Félins.jpg|frameless]]
||{{IUCN status|VU}}<ref>{{cite iucn |title=''Leopardus tigrinus'' |author1=Payan, E. |author2=de Oliveira, T. |name-list-style=amp |page=e.T54012637A50653881 |date=2016}}</ref><br />[[File:Oncilla distribution.jpg|frameless]]
|-- style="vertical-align: top;"
|[[Pampas cat]] ''L. colocola'' {{small|([[Juan Ignacio Molina|Molina]], 1782)}}<ref name="Molina" /><br />[[File:Leopardus pajeros 20101006.jpg|frameless]]
||{{IUCN status|NT}}<ref>{{Cite iucn |title=''Leopardus colocolo'' |author=Lucherini, M. |author2=Eizirik, E. |author3=de Oliveira, T. |author4=Pereira, J. |author5=Williams, R.S.R. |page=e.T15309A97204446 |date=2016}}</ref><br />[[File:PampasCat_distribution.jpg|frameless]]
Range includes multiple species
|-- style="vertical-align: top;"
|[[Kodkod]] ''L. guigna'' {{small|(Molina, 1782)}}<ref name="Molina">{{cite book |author=Molina, G. I. |year=1782 |title=Saggio sulla storia naturale del Chilli |location=Bologna |publisher=Stamperia di S. Tommaso d’Aquino |chapter=La Guigna ''Felis guigna'' |chapter-url=http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=192&Pagina=296 |page=295 |access-date=2019-03-25 |archive-date=2019-06-08 |archive-url=https://web.archive.org/web/20190608065804/http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=192&Pagina=296 |url-status=dead }}</ref><br />[[File:Leopardus guigna.jpeg|frameless]]
||{{IUCN status|VU}}<ref>{{Cite iucn |title=''Leopardus guigna'' |author=Napolitano, C. |author2=Gálvez, N. |author3=Bennett, M. |author4=Acosta-Jamett, G. |name-list-style=amp |author5=Sanderson, J. |page=e.T15311A50657245 |date=2015 |access-date=14 January 2018}}</ref><br />[[File:Guigna distribution.jpg|frameless]]
|-- style="vertical-align: top;"
| [[Margay]] ''L. wiedii'' {{small|([[Heinrich Rudolf Schinz|Schinz]], 1821)}}<ref>{{cite book |last=Schinz |first=H. R. |year=1821 |title=Das Thierreich eingetheilt nach dem Bau der Thiere: als Grundlage ihrer Naturgeschichte und der vergleichenden Anatomie von dem Herrn Ritter von Cuvier. Säugethiere und Vögel, Volume 1 |publisher=Cotta |location=Stuttgart, Tübingen |chapter=Wiedische Katze ''Felis wiedii'' |pages=235–236 |chapter-url=https://books.google.com/books?id=D3o-AAAAcAAJ&pg=PA235}}</ref><br />[[File:Margay.jpg|frameless]] 
||{{IUCN status|NT}}<ref>{{cite iucn |title=''Leopardus wiedii'' |author=de Oliveira, T. |author2=Paviolo, A. |author3=Schipper, J. |author4=Bianchi, R. |author5=Payan, E. |author6=Carvajal, S.V. |name-list-style = amp |page=e.T11511A50654216 |year=2015}}</ref><br />[[File:Margay distribution.jpg|frameless]]
|-- style="vertical-align: top;"
|[[Geoffroy's cat]] ''L. geoffroyi'' {{small|([[Alcide d'Orbigny|d'Orbigny]] & [[Paul Gervais|Gervais]], 1844)}}<ref>{{Cite journal |last1=D'Orbigny |first1=A. |last2=Gervais |first2=P. |name-list-style=amp |date=1844 |title=Mammalogie: Nouvelle espèce de Felis |url=https://archive.org/details/extraitsdesproc79184244soci/page/40 |journal=Extraits des Procès-verbaux des Séances |volume=9 |pages=40–41}}</ref><br />[[File:Salzkatze.jpg|frameless]]
||{{IUCN status|LC}}<ref>{{Cite iucn |title=''Leopardus geoffroyi'' |author=Pereira, J. |author2=Lucherini, M. |author3=Trigo, T. |name-list-style=amp |page=e.T15310A50657011 |date=2015}}</ref><br />[[File:GeoffroysCat_distribution.jpg|frameless]]
|-- style="vertical-align: top;"
|[[Andean mountain cat]] ''L. jacobita'' {{small|([[Emilio Cornalia|Cornalia]], 1865)}}<ref>{{cite journal |last1=Cornalia |first1=E. |year=1865 |title=Descrizione di una nuova specie del genere Felis. ''Felis jacobita'' (Corn.) |journal=Memorie della Societá Italiana di Scienze Naturali |volume=1 |pages=3–9 |url=https://archive.org/details/memoriedella118651866mila/page/n11}}</ref><br />[[File:Andean cat 1 Jim Sanderson.jpg|frameless]]
||{{IUCN status|EN}}<ref>{{Cite iucn |title=''Leopardus jacobita'' |author=Villalba, L. |author2=Lucherini, M. |author3=Walker, S. |author4=Lagos, N. |author5=Cossios, D. |author6=Bennett, M. |author7=Huaranca, J. |name-list-style=amp |page=e.T15452A50657407 |date=2016}}</ref><br />[[File:AndeanCat_distribution.jpg|frameless]]
|-- style="vertical-align: top;"
|[[Southern tigrina]] ''L. guttulus'' {{small|(Hensel, 1872)}}<ref>{{cite journal |last1=Hensel |first1=R. |year=1872 |title=Beiträge zur Kenntniss der Säugethiere Süd-Brasiliens |journal=Physikalische Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin |pages=1–130  |issue=1873 |url=https://archive.org/details/abhandlungenderk1872deut/page/n152}}</ref><br />[[File:Gato-do-mato-pequeno.jpg|frameless]]
|{{IUCN status|VU}}<ref>{{Cite iucn |title=''Leopardus guttulus'' |author=de Oliveira, T. |author2=Trigo, T. |author3=Tortato, M. |author4=Paviolo, A. |author5=Bianchi, R. |author6=Leite-Pitman, M. R. P. |name-list-style=amp |page=e.T54010476A54010576 |date=2016}}</ref><br />[[File:SouthernTigerCat distribution.jpg|frameless]]
|-
|}

Additionally, more recent genetic analyses in 2017 and 2021 proposed the recognition of a third tigrina-like species, ''[[Leopardus emiliae]]''.<ref>{{cite journal|author1=do Nascimento, F.O. |author2=Feijó, A. |title=Taxonomic revision of the tigrina ''Leopardus tigrinus'' (Schreber, 1775) species group (Carnivora, Felidae) |journal=Papéis Avulsos de Zoologia |year=2017 |volume=57 |issue=19 |pages=231–264|doi=10.11606/0031-1049.2017.57.19 |doi-access=free}}</ref><ref>{{cite journal |last1= Trindade|first1= Fernanda J.|last2= Rodrigues|first2= Maíra R.|last3= Figueiró|first3= Henrique V.|last4= Li|first4= Gang|last5= Murphy|first5= William J.|last6= Eizirik|first6= Eduardo|title= Genome-Wide SNPS Clarify a Complex Radiation and Support Recognition of an Additional Cat Species|url= |journal= Molecular Biology and Evolution|year= 2021|volume= 38|issue= 11|pages= 4987–4991|doi= 10.1093/molbev/msab222|pmid= 34320647|pmc= 8557425|access-date=}}</ref>

A 2021 analysis of 142 pampas cat museum specimen collected across South America showed significant morphological differences between them. Therefore, it was proposed to split the historically-contentious pampas cat [[species complex]] into five species: ''Leopardus colocolo'', ''[[Leopardus braccatus]]'', ''[[Leopardus garleppi]]'', ''Leopardus munoai'', and ''[[Leopardus pajeros]]''.<ref>{{cite journal |last1=Nascimento |first1=F.O.D. |last2=Cheng |first2=J. |last3=Feijó |first3=A. |name-list-style=amp |year=2021 |title=Taxonomic revision of the pampas cat ''Leopardus colocola'' complex (Carnivora: Felidae): an integrative approach |journal=Zoological Journal of the Linnean Society |volume=191 |issue=2 |pages=575–611 |doi=10.1093/zoolinnean/zlaa043 |url=https://www.researchgate.net/publication/341726517|doi-access=free }}</ref> Later that same year, it was noted that the oldest available name for pampas cats of the Uruguayan savannah region was ''[[Leopardus fasciatus]]'', not ''L. munoai''.<ref>{{Cite journal|title=The oldest available name for the pampas cat of the Uruguayan Savannah ecoregion is Leopardus fasciatus (Larrañaga 1923) |year=2022 |last1=Martínez-Lanfranco |first1=Juan Andrés |last2=González |first2=Enrique M. |journal=Therya |volume=13 |issue=3 |pages=259–264 |doi=10.12933/therya-22-1187 |s2cid=252649692 |doi-access=free }}</ref>

Another study in 2023 described another new species, ''[[Leopardus narinensis]]'', based on a single dried skin collected in 1989 on the [[Galeras Volcano]] in the Nariño Department of Colombia. They found it to be very different from all other ''Leopardus'' species both morphologically and genetically.<ref name=narinensis>{{Cite Q|Q121764991|doi-access=free}}</ref>

A different study in 2024 did a detailed analysis of both the morphology and genetics of specimens assigned to ''Leopardus tigrinus'', ''Leopardus guttulus'', and ''Leopardus emiliae''. It suggested ''L. t. pardinoides'' should be elevated to species status as ''Leopardus pardinoides'' due to significant differences in morphology, genetics, and ecology. The study also assigned the subspecies ''L. t. oncilla'' to be a subspecies of ''L. pardinoides'' as ''L. p. oncilla''. Additionally, genetic analysis suggested that ''Leopardus emiliae'' was not genetically distinct from ''L. tigrinus'', and thus may be invalid. The study recommended the common names savannah tiger-cat for ''L. tigrinus'', Atlantic Forest tiger-cat for ''L. guttulus'', and clouded tiger-cat for ''L. pardinoides''.<ref>{{Cite journal|title=Ecological modeling, biogeography, and phenotypic analyses setting the tiger cats' hyperdimensional niches reveal a new species |year=2024 |last1=de Oliveira |first1=Tadeu G. |last2=Fox-Rosales |first2=Lester A. |last3=Ramírez-Fernández |first3=José D. |journal=Scientific Reports |volume=14 |issue=1 |pages=2395 |doi=10.1038/s41598-024-52379-8 |pmid= 38287072|pmc=10825201 |bibcode=2024NatSR..14.2395D }}</ref>

An expanded list of ''Leopardus'' species would be:<ref name=mdd-Leopardus>*{{cite mdd|genus=Leopardus|access-date=8 December 2023}}</ref>
*''Leopardus braccatus'', the [[Pantanal cat]] or Brazilian pampas cat
*''Leopardus colocola'', the colocolo or Central Chilean pampas cat
*''Leopardus emiliae'', the eastern tigrina or Snethlage's tigrina
*''Leopardus fasciatus'', Muñoa’s pampas cat or Uruguayan pampas cat
*''Leopardus garleppi'', the northern pampas cat or Garlepp's pampas cat
*''Leopardus geoffroyi'', Geoffroy's cat
*''Leopardus guigna'', the kodkod
*''Leopardus guttulus'', the southern tigrina or Atlantic Forest tiger-cat
*''Leopardus jacobita'', the Andean mountain cat
*''Leopardus narinensis'', the Nariño cat, Galeras cat, or red tigrina<ref name=narinensis/>
*''Leopardus pajeros'', the southern pampas cat
*''Leopardus pardalis'', the ocelot
*''Leopardus pardinoides'', the clouded tiger-cat
*''Leopardus tigrinus'', the oncilla, northern tigrina, or savannah tiger-cat
*''Leopardus weidii'', the margay

=== 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/> 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 last [[common ancestor]] of ''Leopardus'', ''Puma'' and ''Lynx'' is estimated to have lived {{mya|10.95|6.30}}, based on analysis of nuclear DNA of cat species.<ref name=Johnson2006/> Analysis of their mitochondrial DNA indicates that their last common ancestor lived {{mya|14.04|6.83}}.<ref name=Li_al2016/> ''Leopardus'' forms an [[evolution]]ary [[Lineage (evolution)|lineage]] that [[Genetic divergence|genetically diverged]] between {{mya|4.25|2.02}}<ref name=Johnson2006/> and {{mya|5.19|0.98}}.<ref name=Li_al2016/> It crossed the [[Isthmus of Panama]] probably during the [[Great American Biotic Interchange]] in the [[late Pliocene]].<ref name=Johnson2006/>
''[[Leopardus vorohuensis]]'' is an extinct species of the genus, of which [[fossil]]s were found in the Argentinian [[Vorohué Formation]] dated to the [[early Pleistocene]]; its [[supraorbital foramen]] and shape of teeth resemble those of the pampas cat.<ref name=Berta1983>{{cite journal |last1=Berta |first1=A. |year=1983 |title=A new species of small cat (Felidae) from the late Pliocene – early Pleistocene (Uquian) of Argentina |journal=Journal of Mammalogy |volume=64 |issue=4 |pages=720–725 |doi=10.2307/1380541 |jstor=1380541}}</ref>

Within the genus, three distinct [[clade]]s were identified: one comprising the ocelot and the margay, a second the Andean mountain cat and Pampas cat, and the third the kodkod, oncilla and Geoffroy's cat.<ref>{{cite journal |author1=Johnson, W.E. |name-list-style=amp |author2=Dratch, P.A. |author3=Martenson, J.S. |author4=O'Brien, S.J. |year=1996 |title=Resolution of recent radiations within three evolutionary lineages of Felidae using mitochondrial restriction fragment length polymorphism variation |journal=Journal of Mammalian Evolution |volume=3 |issue=2 |pages=297–120 |doi= 10.1007/BF01454358|s2cid=38348868 |doi-access=free }}</ref><ref>{{cite journal |doi=10.1093/jhered/89.3.227 |last1=Johnson |first1=W. E. |last2=Culver |first2=M. |last3=Iriarte |first3=J. A. |last4=Eizirik |first4=E. |last5=Seymour |first5=K. L. |last6=O'Brien |first6=S. J. |name-list-style=amp |year=1998 |title=Tracking the evolution of the elusive Andean mountain cat (''Oreailurus jacobitus'') from mitochondrial DNA |journal=Journal of Heredity |volume=89 |issue=3 |url=http://jhered.oxfordjournals.org/cgi/reprint/89/3/227.pdf |pages=227–232 |pmid=9656464|doi-access=free }}</ref> The following [[cladogram]] shows estimated [[divergence times]] in million years ago (mya).

{{clade gallery |align=left |width=650px; |main-caption=Phylogenetic relationships of living ''Leopardus'' species as derived through analysis of
|header1=nuclear DNA:<ref name=Johnson2006/>
|cladogram1={{clade |label1=[[Felidae]]
 |1={{clade |label1=[[Felinae]]
   |1={{clade |label1='''''Leopardus''''' |sublabel1=4.25–2.02 mya
     |1={{clade |sublabel1=1.48–0.56 mya |sublabel2=3.56–1.68 mya
      |1={{clade |sublabel1=1.21–0.41 mya
         |1={{clade |1=Kodkod |2=Geoffroy's cat }}
         |2={{clade |1=Oncilla }} }}
      |2={{clade |sublabel1=2.41–1.01 mya |sublabel2=2.70–1.18 mya
          |2={{clade |1=Pampas cat |2=Andean mountain cat }}
          |1={{clade |1=Margay |2=Ocelot }} }} }}
    |2=other Felinae lineages }}
  |2=[[Pantherinae]] }} }}
|header2=mitochondrial DNA:<ref name=Li_al2016/>
|cladogram2=
{{clade |style=font-size:90%;line-height:100% |label1=Felidae
|1={{clade |label1=Felinae
  |1={{clade |label1='''''Leopardus''''' |sublabel1=5.19–0.98 mya
    |1={{clade |sublabel1=4.91–0.64 mya |sublabel2=4.85–0.35 mya
      |1={{clade |sublabel1=4.71–0.26 mya |sublabel2=4.76–0.05 mya
       |1={{clade |sublabel1=4.17–0.02 mya
         |1={{clade |1=South American oncilla |2=Pampas cat }}
         |2=Andean mountain cat }}
       |2={{clade |1=Ocelot |2=Margay }} }}
      |2={{clade |sublabel1=4.64–0.04 mya
        |1={{clade |1=Geoffroy's cat |2=Kodkod }}
        |2=Central American oncilla }} }}
    |2=other Felinae lineages }}
  |2=Pantherinae }} }} }}
{{clear|left}}

== References ==
{{Reflist}}

== External links ==
*{{BioRef|asm|id=1005994 |title=''Leopardus'' |version=1.5 |access-date=26 August 2021}}

{{Carnivora|Fe.}}
{{Feliformia|Fel.|state=collapsed}}
{{Taxonbar|from=Q318414}}
{{Authority control}}

[[Category:Leopardus| ]]
[[Category:Carnivoran genera]]
[[Category:Extant Pleistocene first appearances]]
[[Category:Felids of Central America]]
[[Category:Felids of South America]]
[[Category:Felids of North America]]
[[Category:Mammals described in 1842]]
[[Category:Taxa named by John Edward Gray]]
[[Category:Felines]]