Panthera onca mesembrina

Short description: Extinct subspecies of carnivore

Panthera onca mesembrina

Temporal range: Late Pleistocene
~0.016–0.012 Ma[1]

PreЄ

Lost holotype skull of P. onca mesembrina
Scientific classification
Kingdom:  | Animalia
Phylum:  | Chordata
Class:  | Mammalia
Order:  | Carnivora
Suborder:  | Feliformia
Family:  | Felidae
Subfamily:  | Pantherinae
Genus:  | Panthera
Species:  |

P. onca

Subspecies:  |

†P. o. mesembrina

Trinomial name
Panthera onca mesembrina

Cabrera, 1934

Synonyms

* Iemish listai Roth, 1899
* Felis listai Roth, 1904

Panthera onca mesembrina, also known as the Patagonian panther,[2] is an extinct subspecies of jaguar (Panthera onca) that was endemic to southern Patagonia during the late Pleistocene epoch. It is known from several fragmentary specimens, the first of which found was in 1899 at "Cueva del Milodon" in Chile. These fossils were referred to a new genus and species "Iemish listai" by naturalist Santiago Roth, who thought they might be the bones of the mythological iemisch of Tehuelche folklore. A later expedition recovered more bones, including the skull of a large male that was described in detail by Angel Cabrera in 1934. Cabrera created a new name for the giant felid remains, Panthera onca mesembrina, after realizing that its fossils were near-identical to modern jaguars'. P. onca mesembrina's validity is disputed, with some paleontologists suggesting that it is a synonym of Panthera atrox.

The bones of P. onca mesembrina are nearly double the size of the largest living jaguars (P. onca onca), and estimates place it at over 231 kilograms (509 lb) in weight. This would make it the largest jaguar and one of the heaviest known felids. Fragments of skin have been collected from "Cueva del Milodon", showing that it was dark red with lighter, yellowish stripes on its forelimbs. Cave paintings made by indigenous peoples possibly depicting the animal have been found in the El Ceibo, which features a red coat with stripes and spots. P. onca mesembrina was carnivorous and hunted a variety of large mammals, including the ground sloth Mylodon, horse Hippidion, and camelid Lama.

## Contents

* 1 History and taxonomy
* 2 Description
* 2.1 Skin and coloration
* 3 Paleobiology and paleoecology
* 4 See also
* 5 References

## History and taxonomy

The "Cueva del Milodon" in southern Chile, where the type fossils of Panthera onca mesembrina were collected

Near his farm in mountains in Última Esperanza Province, Chile, German explorer Hermann Eberhard came upon a cave bearing fossils of the ground sloth Mylodon, leading it to be dubbed "Cueva del Milodón". These fossils date to the late Pleistocene, dating to as recent as 12,000 BP, making them from the Lujanian SALMA.[2][1] In March of 1899, excavations were made by Erland and Otto Nordenskjöld, adventurers from Sweden, who found a radius and ulna that they referred to a jaguar.[3][4] Later that year, expeditions by German naturalist Rodolfo Hauthal (de) found several isolated postcranial elements coming from a giant felid, which he donated to the Museo de La Plata.[5][6]

There, the museum's director Santiago Roth described the fossils in 1899 as being from a novel genus and species of feline, which he designated "Iemish listai". This name references iemisch, a mythological monster of native Patagonian folklore.[7] Roth noted that the size of the fossils fit descriptions made by Tehuelche locals and by Argentine naturalist Florentino Ameghino in the 1880s. This led Roth to believe it could still be living, with a Tehuelche guide telling him that a living individual resided near Lake Buenos Aires. However, this name is now considered a nomen nudum as Roth based it on the legends and fossils, violating Article 72.5 of the ICZN, which stipulates that species can only be based on physical specimens or depictions.[8][9] The next year, English paleontologist Arthur Smith Woodward referred the material to Felis, noting the bones' similarities to those of extant jaguars.[10] Later in 1900, Hauthal collected additional felid elements from the cave including a partial skull, two mandibles, other postcranial remains, and isolated portions of skin. Roth did not describe these remains until 1904. Following Woodward's hypothesis, Roth abandoned the genus name "Iemish" and instead made "I. listai" a species of Felis, but argued that it was distinct from Felis onca due to its size.[11]

Later analysis by Argentine paleontologist Angel Cabrera in 1934 led him to conclude that some of the bones were from Neomylodon and that the name "Iemish listai" was invalid, as it was used to refer to these erroneously assigned fossils and iemisch. Cabrera noted that the bones were near identical to those of jaguars, but their size was greater than that of any known jaguar. Due to "Iemish listai" being unusable, Cabrera proposed a new name, Panthera onca mesembrina, with the incomplete skull (MLP 10-90) as the holotype. The subspecies name mesembrina means "southern" after the discovery of its fossils in Patagonia. Unfortunately the skull MLP 10-90 was lost, but was illustrated by Cabrera and Roth. Cabrera stated that the jaguar may have been seen or hunted by native peoples, pointing out a pathology on the skull matching those characteristic of arrow wounds.[9] More fossils were later found at the cave, including feces, and described during the 20th century. Several other caves across Chilean and Argentine Patagonia produced additional fossils, including those of juveniles and elderly individuals. In a cave near Sofia Lake, many isolated specimens from juveniles were unearthed.[12]

A 2016 study of a mitochondrial DNA sequence obtained from bones attributed to P. onca mesembrina found that they represented those of a true jaguar, but belonged to a mitochondrial lineage highly divergent from those found in living jaguars. This supported its validity and independence from other P. onca subspecies.[1] However, in 2017 morphological research of the holotype and other remains by Chimento et al instead referred it to the lion relative Panthera atrox.[12] This conclusion has seen acceptance by some authors. In his 2020 thesis, researcher Nicholas Freymueller argued that this could be a case of Bergmann's rule, as traits correlated with larger body size were used to justify the synonymy in Chimento et al's study.[13] In 2024, it was suggested that the validity of subspecific assignments on both P. o. mesembrina and the North American P. o. augusta remains unresolved, since both fossil and living jaguars show a considerable variation in morphometry.[14]

## Description

Panthera onca mesembrina is the largest subspecies of P. onca, with a 2017 estimate placing its body mass at 231 kilograms (509 lb) based on the type material.[12] This makes it not only the heaviest known jaguar by as much as 90 kilograms (200 lb), but one of the largest known felids.[15][16] The trend of large body size among Pleistocene felids was likely due to a multitude of factors such as: prey size, environmental conditions of the epoch, and, for P. onca mesembrina specifically, Bergmann's Rule. A similar trend of gigantism is observed in the North American subspecies P. o. augusta, which was around 15-20% larger than modern jaguars at around 190 kilograms (420 lb).[12] P. onca mesembrina shared the robust, stocky build of P. onca onca but to an even greater degree.[9]

P. o. mesembrina and P. o. augusta share many similarities that are lacking in living jaguars, such as on the skull, with the presence of exposed foramina (small holes) near the nasal (nose bone) and an expanded nasal aperture (nose opening). While the skulls of extant P. onca onca individuals are short, heavy, and wider at the end of the snout, P. onca mesembrina's skull had a longer snout which tapered slightly at the premaxilla. The teeth of P. onca mesembrina were remarkably large compared to those of other Panthera species, a trait shared by living jaguars. However, few teeth are known from this subspecies. In the first molar, the paraconid (mesial cusp) is much shorter than the protoconid (distal cusp), a feature present in P. atrox and P. onca mesembrina but absent in most other pantherines.[12]

In the postcranial anatomy, P. onca mesembrina also stands out from other P. onca in its robusticity. The humerus (upper arm bone) is thicker overall and has an expanded epicondylar crest, similar to P. atrox. The dorsal vertebrae feature short, dorsally projected processes (bony projections), characteristics absent in most jaguars.[12]

### Skin and coloration

Cave painting of P. onca mesembrina in El Ceibo archaeological locality, Santa Cruz, Patagonia, Argentina.

In his 1904 description, Roth assigned pieces of skin from the face and limbs as well as a piece of leather to F. "listai", now P. onca mesembrina. These fragments had been collected from Cueva del Milodon in 1900, though a piece of Mylodon fur had earlier been misidentified as belonging to the cryptid iemisch. Roth described these portions in detail, noting that the skin from the face was reddish with hints of "shiny chestnut" while the limbs are dark overall with yellowish striping. He later called it a "cat of beautiful colors".[11] Notably, a cave painting from El Ceibo archaeological locality in Santa Cruz Province bears an illustration matching this description. The cave, which also features art of guanacos and human handprints, features a 1.5 metres (4.9 ft) long depiction of a remarkably large, striped feline. The reasoning for the size of this painting compared to others may be due to native people revering the animal. The colors are also shown, with a reddish background, black spots, and lightly colored forelimbs. All of these characteristics coincide with the skin found at Cueva del Milodon, strongly implying that they come from the same animal.[17][18][19]

## Paleobiology and paleoecology

The holotype of Panthera onca mesembrina preserved a shallow irregular pit on the lateral surface on the maxilla; this represents a puncture made by the canine of another jaguar, which healed during life. As George Simpson pointed out in 1941, all extant collected specimens with these punctures were males and no females with such scars have been found. The presence of this wound suggests that the specimen was a male based on this common social behavior between living male jaguars.[12][20]

Life reconstruction of Mylodon, a prey item of P. onca mesembrina.

At Cueva del Milodon, fossils of the ground sloth Mylodon, horse Hippidion, and camelids have been found with tooth and scratch marks that match the teeth of P. onca mesembrina. Based on isotopic analyses, P. onca mesembrina preferred to hunt Hippidion and Lama, as well as juveniles of larger mammals like Mylodon. Evidence from dens of P. onca mesembrina backs this, as Lama and Hippidion fossils were more common than other herbivores.[21] The majority of the fossils found were limb elements, a circumstance that happens with living felines' burrows and cave where food is taken and consumed.[22] Though P. onca mesembrina is one of four known carnivores in the cave, its fossils are more common and better preserved, whereas species like Smilodon populator are represented by a handful of fossils.[23][2] Modern jaguars and cougars are able to drag their kills long distances, which P. onca mesembrina was likely capable of as well.[24][25] The discovery of isolated herbivore bones in largely carnivore-occupied dens in other caves supports this as they were likely kills taken to be eaten. Cueva del Milodon, however, was a frequent nest of Mylodons based on the occurrence of juvenile and newborn Mylodon individuals. It is possible that it was a recurring hunting spot for P. onca mesembrina due to this.[21] Some of the prey items reached great sizes, such as a skull from a 3–4 metres (9.8–13.1 ft) long Mylodon that bears several bite marks.[2] Coprolites containing Mylodon dermal ossicles were found there, which were likely defecated by P. onca mesembrina.[12][23]

The majority of sites where P. onca mesembrina has been found in Argentina and Chile are very open and arid, contrasting to the lush rainforest habitats of modern jaguars.[26][27] This conclusion is backed by their diets, with P. onca mesembrina hunting grazing animals[21] whereas living jaguars prefer browsers like tapirs, anteaters, and peccaries.[28] Fossils of P. onca mesembrina have been reported from eight sites according to Chimento et al (2017) and Paunero et al (2017), all of which are in southern Argentina and Chile.[29][12][30] Cueva del Milodon is the southernmost site known to be occupied by jaguars. This region is extremely cold, with temperatures dropping as low as −16 °C (3 °F).[31]

## See also

Note: This topic belongs to "Cats" portal

* Panthera gombaszogensis

## References

1. ↑ 1.0 1.1 1.2 Metcalf, Jessica L.; Turney, Chris; Barnett, Ross; Martin, Fabiana; Bray, Sarah C.; Vilstrup, Julia T.; Orlando, Ludovic; Salas-Gismondi, Rodolfo et al. (2016-06-03). "Synergistic roles of climate warming and human occupation in Patagonian megafaunal extinctions during the Last Deglaciation". Science Advances 2 (6). doi:10.1126/sciadv.1501682. ISSN 2375-2548. PMID 27386563. Bibcode: 2016SciA....2E1682M.
2. ↑ 2.0 2.1 2.2 2.3 Martin, Fabiana María (2018). "Cueva del Milodón. The hunting grounds of the Patagonian panther" (in en). Quaternary International 466: 212–222. doi:10.1016/j.quaint.2016.05.005. Bibcode: 2018QuInt.466..212M. https://linkinghub.elsevier.com/retrieve/pii/S1040618215302962.
3. ↑ Latorre, Claudio (1998). "Paleontología de mamíferos del Alero Tres Arroyos I, Tierra del Fuego, XII Región, Chile.". Anales Instituto Patagonia. Cs 26: 77–90. http://200.54.63.162/bitstream/handle/20.500.11893/1178/Latorre_Anales_1998_vol26_pp77-90.pdf?sequence=1&isAllowed=y.
4. ↑ Nordenskjöld, Erland (1900). "Iakttagelser och fynd Grottor vid Ultima Esperanza sydvestra Patagonien.". Kongl. Svenska Vetenskaps-Akademiens Handlingar 33: 1–24.
5. ↑ "Hauthal, Rudolf Johannes Friedrich (Rodolfo) (1854-1928) on JSTOR". https://plants.jstor.org/stable/10.5555/al.ap.person.bm000044947.
6. ↑ Roth, Santiago (1899). "Descripción de los restos encontrados en la caverna de Última Esperanza.". Revista del Museo de La Plata 9: 421–453. https://www.biodiversitylibrary.org/item/37352#page/524/mode/1up.
7. ↑ Jacoby, Charlie (November 20, 2000). "A Century On And The Adventure Continues In Hunt For The Giant Sloth". The Express.
8. ↑ "International Code of Zoological Nomenclature". https://code.iczn.org/types-in-the-species-group/article-72-general-provisions/?frame=1.
9. ↑ 9.0 9.1 9.2 Cabrera, Angel (1934). "Los yaguares vivientes y extinguidos.". Notas Preliminares del Museo de la Plata 2: 9–39. http://naturalis.fcnym.unlp.edu.ar/id/004038.
10. ↑ Woodward, A. Smith (1900). "On some Remains of Grypotherium (Neomylodori) listai and associated Mammals from a Cavern near Consuelo Cove, Last Hope Inlet, Patagonia." (in en). Proceedings of the Zoological Society of London 69 (1): 64–78. doi:10.1111/j.1096-3642.1890.tb01704.x. ISSN 0370-2774. https://onlinelibrary.wiley.com/doi/10.1111/j.1096-3642.1890.tb01704.x.
11. ↑ 11.0 11.1 Roth, Santiago (1904). "Nuevos restos de mamíferos de la Caverna Eberhardt en Última Esperanza.". Revista del Museo de La Plata 11: 39–53. https://publicaciones.fcnym.unlp.edu.ar/rmlp/article/view/1209.
12. ↑ 12.0 12.1 12.2 12.3 12.4 12.5 12.6 12.7 12.8 Chimento, Nicolás R.; Agnolin, Federico L. (2017-11-01). "The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications" (in en). Comptes Rendus Palevol 16 (8): 850–864. doi:10.1016/j.crpv.2017.06.009. ISSN 1631-0683. Bibcode: 2017CRPal..16..850C.
13. ↑ Freymueller, N. A. (2020). Niche dynamics of the felid guild following the Pleistocene Megafaunal Extinction (MSc thesis). The University of New Mexico.
14. ↑ Srigyan, M.; Schubert, B.W.; Bushell, M.; Santos, S.H.D.; Figueiró, H.V.; Sacco, S.; Eizirik, E.; Shapiro, B. (2024). "Mitogenomic analysis of a late Pleistocene jaguar from North America". Journal of Heredity 115 (4): 424–431. doi:10.1093/jhered/esad082. PMID 38150503.
15. ↑ Christiansen, Per; Harris, John M. (2005). "Body size of Smilodon (Mammalia: Felidae)" (in en). Journal of Morphology 266 (3): 369–384. doi:10.1002/jmor.10384. ISSN 0362-2525. PMID 16235255. https://onlinelibrary.wiley.com/doi/10.1002/jmor.10384.
16. ↑ DeSantis, Larisa R. G.; Schubert, Blaine W.; Scott, Jessica R.; Ungar, Peter S. (2012-12-26). "Implications of Diet for the Extinction of Saber-Toothed Cats and American Lions" (in en). PLOS ONE 7 (12). doi:10.1371/journal.pone.0052453. ISSN 1932-6203. PMID 23300674. Bibcode: 2012PLoSO...752453D.
17. ↑ Borrero, L. A. (2001). "El poblamiento de la Patagonia: toldos, milodones y volcanes". Emecé editores.
18. ↑ Cardich, A. R. (1979). "Un motivo sobresaliente de las pinturas rupestres de" El Ceibo"(Santa Cruz)". Relaciones de la Sociedad Argentina de Antropología 13.
19. ↑ David, Bruno; McNiven, Ian J. (2018-10-17) (in en). The Oxford Handbook of the Archaeology and Anthropology of Rock Art. Oxford University Press. ISBN 978-0-19-084495-0. https://books.google.com/books?id=tXFyDwAAQBAJ&dq=Panthera+onca+mesembrina&pg=PA290.
20. ↑ Simpson, G. G. (1941). "Large pleistocene felines of North America". American Museum Novitates (1136): 1–27.
21. ↑ 21.0 21.1 21.2 Prevosti, Francisco J.; Martin, Fabiana M. (2013). "Paleoecology of the mammalian predator guild of Southern Patagonia during the latest Pleistocene: Ecomorphology, stable isotopes, and taphonomy" (in en). Quaternary International 305: 74–84. doi:10.1016/j.quaint.2012.12.039. Bibcode: 2013QuInt.305...74P. https://linkinghub.elsevier.com/retrieve/pii/S1040618213000049.
22. ↑ BORRERO, L. A. (1998). "The extinction of the megafauna: a supra-regional approach". Anthropozoologica 25: 209–216.
23. ↑ 23.0 23.1 Martin, F. M. (2008). Tafonomía y paleoecología de la transición Pleistoceno-Holoceno en Fuego-Patagonia. Doctoral Dissertation, Universidad Nacional de la Plata (Tesis). doi:10.35537/10915/4376.
24. ↑ Bank, M. S.; Franklin, W. L. (1998). "Puma (Puma concolor patagonica) feeding observations and attacks on guanacos (Lama guanicoe)". Mammalia (Paris) 62 (4): 599–605.
25. ↑ Anyonge, William (1993). "Body mass in large extant and extinct carnivores" (in en). Journal of Zoology 231 (2): 339–350. doi:10.1111/j.1469-7998.1993.tb01922.x. ISSN 0952-8369. https://onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.1993.tb01922.x.
26. ↑ Carman, R. L. (1984). "Límite austral de la distribución del tigre o yaguareté (Leo onca) en los siglos XVIII y XIX". Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" e Instituto Nacional de Investigación de las Ciencias Naturales 13 (29): 293–296.
27. ↑ Diaz, Norma I. (2010-01-01). "New historical records of the jaguar (Panthera onca) in Patagonia" (in es). Revista Mexicana de Mastozoología (Nueva Época) 14 (1): 23–35. doi:10.22201/ie.20074484e.2010.14.1.25. ISSN 2007-4484. http://www.revmexmastozoologia.unam.mx/ojs/index.php/rmm/article/view/25.
28. ↑ Hayward, Matt W.; Kamler, Jan F.; Montgomery, Robert A.; Newlove, Anna; Rostro-García, Susana; Sales, Lilian P.; Van Valkenburgh, Blaire (2016). "Prey Preferences of the Jaguar Panthera onca Reflect the Post-Pleistocene Demise of Large Prey". Frontiers in Ecology and Evolution 3. doi:10.3389/fevo.2015.00148. ISSN 2296-701X.
29. ↑ Paunero, Rafael; Davis, Catalina Valiza; Rindel, Diego; Tessone, Augusto (December 2017). "THE PLEISTOCENE FAUNA: ZOOARCHAEOLOGICAL EVIDENCE IN THE CENTRAL PLATEAU OF SANTA CRUZ, THE SITES OF LA MARÍA". Magallania (Punta Arenas) 45 (2): 181–198. doi:10.4067/S0718-22442017000200181. ISSN 0718-2244. http://www.scielo.cl/scielo.php?script=sci_abstract&pid=S0718-22442017000200181&lng=en&nrm=iso&tlng=en.
30. ↑ Moreno, A. K. M.; Lima-Ribeiro, M. S. (2015-12-31). "Ecological niche models, fossil record and the multi-temporal calibration for Panthera onca (Linnaeus, 1758) (Mammalia: Felidae)". Brazilian Journal of Biological Sciences 2 (4): 309–319. ISSN 2358-2731. http://revista.rebibio.net/v2n4/v02n04a14a.html.
31. ↑ "DESCRIPCIÓN CLIMATOLÓGICA" (in es). Dirección General de Aeronáutica Civil. http://www.meteochile.cl/climas/climas_duodecima_region.html.

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