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''Panthera onca mesembrina'' is an extinct
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of the
jaguar The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus ''Panthera'' native to the Americas. With a body length of up to and a weight of up to , it is the largest cat species in the Americas and the th ...
that was endemic to southern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
during the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
epoch (1.8 mya–11,000 years ago). Its fossils have been excavated primarily in
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
and
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
, though few fossils are known. Genetic analysis in 2016 showed that ''P. onca mesembrina'' was in an extinct sister lineage to extant ''Panthera onca'' species based on genetic evidence, and is the largest known subspecies of jaguar.


History and taxonomy

In the 1890s in the “
Cueva del Milodon Cuevas or Cueva (Spanish for "''cave(s)''") may refer to: Places * Cueva de Ágreda, a municipality located in the province of Soria, Castile and León, Spain * Cuevas Bajas, a town and municipality in the province of Málaga, part of the autonom ...
” in southern Chile, fossil collector Rodolfo Hauthal collected a fragmentary postcranial skeleton of a large Felid that he sent to Santiago Roth who described them as a new genus and species of Felid, "Iemish listai", in 1899, though the name is considered a
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate desc ...
. 5 years later in 1904, Roth reassessed the phylogenetic affinities of “Iemish” and moved the species to ''Felis,'' referring several cranial and fragmentary postcranial elements to the taxon. Notably, 2 fragmentary mandibles, a partial skull, and pieces of skin were some of the specimens referred in 1904. 30 years later in 1934, ''Felis onca mesembrina'' was properly named by
Angel Cabrera In various theistic religious traditions an angel is a supernatural spiritual being who serves God. Abrahamic religions often depict angels as benevolent celestial intermediaries between God (or Heaven) and humanity. Other roles incl ...
based on that partial skull, likely from a male,Simpson, G. G. (1941). Large pleistocene felines of North America. ''American museum novitates'', ''1136'', 1-27. from “Cueva del Milodon” and the other material from the site was referred to it. Unfortunately, the skull ( MLP 10-90) was lost, and was only illustrated by Cabrera and Roth. Later, more material, including feces and mandibles, was referred to as ''F. onca mesembrina'' from the island of Tierra del Fuego, Argentina and other sites in southern Chile. One site in Última Esperanza, Chile was excavated in the 1980s and 90s and bore many fossils, including those of juveniles and old adults. All material from ''P. onca mesembrina'' dates to the late Pleistocene. In 2016, the subspecies was found to belong to ''Panthera onca'' in a genetic study, which supported its identity as a unique subspecies of jaguar. Later in 2017, the subspecies was synonymized with ''Panthera atrox'' based on morphological similarities of all material, although these similarities are unreliable and the subspecies is valid.


Fossil distribution

Fossils have been uncovered from southern Chile at "Cueva del Milodon", the southernmost site of any jaguar recorded, to possibly Formosa Province in Argentina, the northernmost record of the subspecies. In 2021, fossils from Cuvieri Cave in Minas Gerais, Brazil were referred to ''P. onca'' and possibly belong to ''P. onca mesembrina.'' The cave contains many fossils from several individuals of differing growth stages preserved alongside other taxa, such as '' Smilodon populator'' and ''
Catonyx cuvieri ''Catonyx'' is an extinct genus of ground sloth of the family Scelidotheriidae, endemic to South America during the Pliocene and Pleistocene epochs. It lived from 2.5 Ma to about 10,000 years ago, existing for approximately . The most recen ...
'', and are the only fossils known of the subspecies from outside Argentina and Chile.


Description and Paleobiology


Size

Of the many ''Panthera onca'' subspecies, none is as large as that of ''Panthera onca mesembrina'', with an estimated body mass of 231.23 kilograms in 2017 based on the material from the "Cueva del Milodon". This size not only places it as far larger than any other known ''Panthera onca'' by as much as 90 kg, but was among the largest known felids with the mass being in the range of felids as big as '' Smilodon populator'' and ''
Panthera atrox ''Panthera atrox'', better known as the American lion, also called the North American lion, or American cave lion, is an extinct pantherine cat that lived in North America during the Pleistocene epoch and the early Holocene epoch, about 340, ...
.'' The trend of large body size among Pleistocene felids was likely due to a multitude of factors, mostly prey size and the different environmental conditions of the epoch, with the North American '' P. onca augusta'' being 15-20% larger than modern ''P. onca augusta'' at around 190 kg. The focused prey size was also calculated in 2017, at 663.4 kilograms, the prey range is also one of the greatest known from any Felid, with evidence of bite marks on ''Mylodon'' and other large fossil mammals supporting this.


Skin and coloration

In 1904, Santiago Roth described pieces of skin from the face and limbs, along with a piece of leather, from the previously mentioned site at "Cueva del Milodon".Roth, S. (1904)
Nuevos restos de mamíferos de la Caverna Eberhardt en Última Esperanza.
''Revista del Museo de La Plata'', ''11'', 39-53.
The skin from the face shows a reddish brown color tone, and the skin from the limbs preserves a yellowish colored stripe. A cave painting from El Ceibo in the Santa Cruz Province of Argentina seems to confirm this, and reduce the possibility of confusion with other jaguars, as similar cave paintings accurately depict jaguars as yellow in color. The painting is depicted alongside those of
guanacos The guanaco (; ''Lama guanicoe'') is a camelid native to South America, closely related to the llama. Guanacos are one of two wild South American camelids, the other being the vicuña, which lives at higher elevations. Etymology The guanaco ...
and humans and is the likely only known depiction of the taxon by Pre-Columbians.


Behavior

Evidence of social groups has been found in the lost holotype of ''Panthera onca mesembrina'' in the form of a shallow irregular pit on the lateral surface of the maxilla-nasal suture posterior to the ascending process of the premaxilla; this represents a puncture by the canine made by other jaguars, healed during life. As George Simpson described in 1941, all extant collected specimens with these punctures were males, and no females with such scars have been found. The presence of a puncture on the lateral surface of the skull suggests that the specimen was a male based on this social behavior between male living jaguars. In the type site at “
Cueva del Milodon Cuevas or Cueva (Spanish for "''cave(s)''") may refer to: Places * Cueva de Ágreda, a municipality located in the province of Soria, Castile and León, Spain * Cuevas Bajas, a town and municipality in the province of Málaga, part of the autonom ...
”, fossils of '' Mylodon, Hippidion'', and many other Pleistocene herbivores have been found with tooth and scratch marks that match the teeth of ''P. onca mesembrina''.BORRERO, L. A. (1998). The extinction of the megafauna: a supra-regional approach. ''Anthropozoologica'', ''25'', 209-216. The majority of the fossils found were limb elements, the same circumstance happens with modern ''Panthera'' species' burrows and caves where food is taken. The fossils notably were of very large sizes, including bite marks on the skull of the 3-4 meter ground sloth ''Mylodon,'' the largest animal found in the cave. Coprolites containing many ''Mylodon'' dermal ossicles was found in the cave, the coprolite likely belonging to ''P. onca mesembrina.'' The majority of sites where ''P. onca mesembrina'' has been found in Argentina and Chile were very open and arid, contrasting to the lush rainforest habitats of modern jaguars.


See also

* '' Panthera onca augusta'' * '' Panthera gombaszoegensis'' * South American jaguar


References

{{Taxonbar, from=Q2710439 Panthera onca mesembrina onca mesembrina Pleistocene carnivorans Pleistocene first appearances Pleistocene extinctions Pleistocene mammals of South America Lujanian Pleistocene Brazil Fossils of Brazil Pleistocene Chile Fossils of Chile Fossil taxa described in 1934 Pleistocene Argentina