Panthera onca mesembrina | |
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Lost holotype skull of P. onca mesembrina | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Felidae |
Subfamily: | Pantherinae |
Genus: | Panthera |
Species: | |
Subspecies: | †P. o. mesembrina |
Trinomial name | |
†Panthera onca mesembrina Cabrera, 1934 | |
Synonyms | |
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Panthera onca mesembrina, also known as the Patagonian panther, [1] 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.
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. [1] [2] 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 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. [2] 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. augusta and P. o. mesembrina remains unresolved, since both fossil and living jaguars show a considerable variation in morphometry. [14]
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. 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 Panthera onca 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]
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]
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]
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] [1] 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. [1] 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]
Felidae is the family of mammals in the order Carnivora colloquially referred to as cats. A member of this family is also called a felid.
The jaguar is a large cat species and the only living member of the genus Panthera that is native to the Americas. With a body length of up to 1.85 m and a weight of up to 158 kg (348 lb), it is the biggest cat species in the Americas and the third largest in the world. Its distinctively marked coat features pale yellow to tan colored fur covered by spots that transition to rosettes on the sides, although a melanistic black coat appears in some individuals. The jaguar's powerful bite allows it to pierce the carapaces of turtles and tortoises, and to employ an unusual killing method: it bites directly through the skull of mammalian prey between the ears to deliver a fatal blow to the brain.
The leopard is one of the five extant species in the genus Panthera. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of 92–183 cm (36–72 in) with a 66–102 cm (26–40 in) long tail and a shoulder height of 60–70 cm (24–28 in). Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).
Panthera is a genus within the family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, snow leopard and tiger, as well as a number of extinct species, including the cave lion and American lion.
Mylodon is an extinct genus of ground sloth belonging to the family Mylodontidae, known from southern South America. With a total length of 3 to 4 m and a body mass of 1-2 tonnes, it is one of the largest mylodontids.
Smilodon is an extinct genus of felids. It is one of the best known saber-toothed predators and prehistoric mammals. Although commonly known as the saber-toothed tiger, it was not closely related to the tiger or other modern cats, belonging to the extinct subfamily Machairodontinae, with an estimated date of divergence from the ancestor of living cats around 20 million years ago. Smilodon was one of the last surviving machairodonts alongside the distantly related Homotherium. Smilodon lived in the Americas during the Pleistocene epoch. The genus was named in 1842 based on fossils from Brazil; the generic name means "scalpel" or "two-edged knife" combined with "tooth". Three species are recognized today: S. gracilis, S. fatalis, and S. populator. The two latter species were probably descended from S. gracilis, which itself probably evolved from Megantereon. The hundreds of specimens obtained from the La Brea Tar Pits in Los Angeles constitute the largest collection of Smilodon fossils.
Panthera spelaea, commonly known as the cave lion, is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion, with the genetic divergence between the two species estimated at around 500,000 years ago. The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago. It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.
The American lion, with the species name meaning "savage" or "cruel", also called the North American lion) is an extinct pantherine cat native to North America during the Late Pleistocene from around 130,000 to 12,800 years ago. Genetic evidence suggests that its closest living relative is the lion, with the American lion representing an offshoot from the lineage of the largely Eurasian cave lion, from which it is suggested to have split around 165,000 years ago. Its fossils have been found across North America, from Canada to Mexico. It was about 25% larger than the modern lion, making it one of the largest known felids to ever exist, and an important apex predator.
The Pantherinae is a subfamily of the Felidae; it was named and first described by Reginald Innes Pocock in 1917 as only including the Panthera species, but later also came to include the clouded leopards. The Pantherinae genetically diverged from a common ancestor between 9.32 to 4.47 million years ago and 10.67 to 3.76 million years ago.
Panthera gombaszoegensis, also known as the European jaguar, is a Panthera species that lived from about 2.0 to 0.35 million years ago in Europe. The first fossils were excavated in 1938 in Gombasek Cave, Slovakia. Some records were also reported from Africa and Asia. P. gombaszoegensis was a medium-large sized species that formed an important part of the European carnivore guild for a period of over a million years. Many authors have posited that it is the ancestor of the American jaguar, with some authors considering it the subspecies Panthera onca gombaszoegensis, though the close relationship between the two species has been questioned.
Panthera fossilis is an extinct species of cat belonging to the genus Panthera, known from remains found in Eurasia spanning the Middle Pleistocene and possibly into the Early Pleistocene.
Cave lions are large extinct carnivorous felids that are classified either as subspecies of the lion, or as distinct but closely related species, depending on the authority.
Cueva del Milodón Natural Monument is a Natural Monument located in the Chilean Patagonia, 24 km (15 mi) northwest of Puerto Natales and 270 km (168 mi) north of Punta Arenas.
Paramylodon is an extinct genus of ground sloth of the family Mylodontidae endemic to North America during the Pliocene through Pleistocene epochs, living from around ~4.9 Mya–12,000 years ago.
Cueva Fell or Fell'sCave is a natural cave and archaeological site in southern Patagonia. Cueva Fell is in proximity to the Pali Aike Crater, another significant archaeological site. Cueva Fell combined with the nearby Pali Aike site have been submitted to UNESCO as a possible World Heritage Site.
Panthera onca augusta is an extinct subspecies of the jaguar that was endemic to North America during the Last Glacial Maximum of the Pleistocene epoch.
Arctotherium is an extinct genus of the Pleistocene short-faced bears endemic to Central and South America. Arctotherium migrated from North America to South America during the Great American Interchange, following the formation of the Isthmus of Panama during the late Pliocene. The genus consists of one early giant form, A. angustidens, and several succeeding smaller species, which were within the size range of modern bears. Arctotherium was adapted to open and mixed habitat. They are genetically closer to the spectacled bear, than to Arctodus of North America, implying the two extinct forms evolved large size in a convergent manner.
Piedra Museo is an archaeological site in Santa Cruz Province, Argentina, and one of the earliest known archaeological remains in the Americas.
Panthera pardus spelaea, also known as the European Ice Age leopard or the cave leopard, is a fossil leopard subspecies which roamed Europe in the Late Pleistocene and possibly the Holocene.
Panthera balamoides is a possibly dubious species described as an extinct species of the big cat genus Panthera that is known from a single fossil found in a Late Pleistocene age cenote in the Yucatan Peninsula, Mexico. P. balamoides has only a single reported specimen, the distal end of a right humerus, that is notably of exceptional size for a felid. It was unearthed in 2012 from an underwater cave and described in 2019 by an international group of paleontologists from Mexico and Germany led by Sarah R. Stinnesbeck. However, several authors have since proposed the humerus represents that of a bear, possibly the extinct Arctotherium, and not a cat.