Leopardus

Leopardus species have spotted fur, with ground colors ranging from pale buff, ochre, fulvous and tawny to light gray.{{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}} Their small ears are rounded and white-spotted; their rhinarium is prominent and naked above, and their nostrils are widely separated.{{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}} They have 36 chromosomes, whereas other felids have 38.{{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 }}

The 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.{{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}}

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;{{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}}
• Margay, Pajeros, Pardalina and Pardalis by Gray in 1867;{{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}}
• Oncilla by Joel Asaph Allen in 1919;{{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}}
• Oreailurus by Ángel Cabrera in 1940;{{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}}
• Colocolo by Reginald Innes Pocock in 1941.{{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 }}

Analysis of skull morphology of these taxa revealed close similarities in their base of skulls and nasal bones, their masticatory muscles, and dentition.{{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}}

Phylogenetic analysis of tissue samples of these taxa and their ability to hybridise support the notion that they are members of the same genus.

The following eight extant Leopardus species have commonly been recognized as valid taxa since 2017:

Additionally, more recent genetic analyses in 2017 and 2021 proposed the recognition of a third tigrina-like species, Leopardus emiliae.{{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}}{{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=}}

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.{{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 }} 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.{{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 }}

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.{{Cite Q|Q121764991|doi-access=free}}

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.{{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 }}

An expanded list of Leopardus species would be:*{{cite mdd|genus=Leopardus|access-date=8 December 2023}}

• 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
• 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

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}}. Analysis of mitochondrial DNA of all Felidae species indicates a radiation at around {{mya|16.76|6.46}}.{{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}}

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. Analysis of their mitochondrial DNA indicates that their last common ancestor lived {{mya|14.04|6.83}}. Leopardus forms an evolutionary lineage that genetically diverged between {{mya|4.25|2.02}} and {{mya|5.19|0.98}}. It crossed the Isthmus of Panama probably during the Great American Biotic Interchange in the late Pliocene.

Leopardus vorohuensis is an extinct species of the genus, of which fossils 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.{{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}}

Within the genus, three distinct clades 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.{{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 }}{{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 }} 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:

|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:

|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}}

{{Reflist}}

• {{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: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