Panthera balamoides Temporal range: | |
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Holotype humerus in anterior and posterior views. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Felidae |
Subfamily: | Pantherinae |
Genus: | Panthera |
Species: | †P. balamoides |
Binomial name | |
†Panthera balamoides Stinnesbeck et al., 2019 | |
Panthera balamoides ("similar to jaguar") 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 (Rancholabrean NALMA, dated to 13,000 BP) age cenote in the Yucatan Peninsula, Mexico. P. balamoides has only a single reported specimen, the distal end of a right humerus (upper arm bone), 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.
Fossils attributed to Panthera balamoides were first unearthed from a debris mount in El Pit cenote, a submerged sinkhole in a diving and snorkel park known as Dos Ojos on the Yucatán Peninsula. [1] This site is located near Tulum in Quintana Roo, Mexico, dating to the Rancholabrean stage of the Late Pleistocene of around 13,000 BP, [2] [3] [1] in which several other fossils were unearthed such as two human individuals and other felids like Smilodon and Panthera atrox. The fossils had been discovered and collected in 2012 via diving by Jerónimo Avilés Olguín, a paleontologist from the Museo del Desierto, from around 44 meters down in the cenote. The fossils later attributed to P. balamoides are fragmentary, consisting only of the distal end of a humerus (upper arm bone) that was later taken and dried in the Colección Paleontológica de Coahuila of the Museo del Desierto in Saltillo, Mexico. [1]
The specimen, desposited under specimen number CPC-2205, remained undescribed until 2018. That year, a team of researchers from Germany and Mexico led by Sarah R. Stinnesbeck described the remains in addition to clavicles from Panthera atrox and a distal humerus of Smilodon gracilis. [4] [1] CPC-2205 was made the holotype of a novel species named Panthera balamoides, the specific name coming from "balam", the Mayan word for "jaguar" due to the discovery of the remains in former Mayan territory, and "eidos" from the Greek root for "similarity". The name calls upon the supposed likeness of the distal humerus' morphology to that of the extant jaguar. [4] [1]
Depending on the phylogenetic position of the species, it have either been a very large and robust pantherine similar to jaguars or a ginormous omnivorous ursid like Arctotherium. If P. balamoides is a Panthera species, it may have been over 100 kilograms (220 lb) in weight despite its build more akin to smaller felids such as jaguars and ocelots. [1] This would make P. balamoides one of the largest known carnivorans, around the weight of modern Asiatic bears and jaguars. [5] [6] If a specimen of Arctotherium, it would be from the smaller species A. wingei based on the discovery of nearby remains. A. wingei has been estimated to have been closer to 83 kilograms (183 lb). [7] [8] [9]
The holotype consists purely of the distal third of the right humerus (upper arm bone), with a preserved length of 136 millimetres (5.4 in) and maximum width of 81 millimetres (3.2 in). The humeral shaft is gracile and straight, attached to a prominent lateral supracondylar ridge with a distal thickness and a laterally convex ridge runs along the distal half of the shaft. Here, the m. bracialis muscle would contact the bone next to the m. triceps brachii. There is a crescent-shaped concave depression proximal to the articlar surfaces that would be a muscle attachment. There is an offset distal articular surface (joint) that longs along the axis of the humeral shaft medially. The epticondylar foramen is prominent and visible from the front and backsides, a trait used by the authors to diagnose the fossil. The coronoid fossa (a small depression) is shallow and located on the anterior face of the epiphysis (end of a long bone) adjacent to the distal articular surface, with an abraded capitulum and trochlea, the parts of the humerus that articulate with the ulna and radius. [1]
The olecranon fossa (depression that articulates with the radius) is located proximal to the distal articlular surface, with a height and weight of around 30 millimetres (1.2 in) making a triangular shape. This fossa was under the attachment of the m. anconeus muscle, making a firm and robust accessory point. [1] These traits of the humerus suggests P. balamoides had robust and short, akin to that of extant jaguars. A combination of these characteristics were used to diagnose the species by Stinnesbeck et al. (2019), but have come into question by other authors who theorize the humerus is not distinct from that of Arctotherium. [9]
In the initial description of P. balamoides, it was decided by the authors that the partial humerus of the holotype could not have been canid or ursid due to the presence of an entepicondylar foramen, which is absent in the two groups with the exception of tremarctine bears like Tremarctos. Other traits like the size and outline of the foramen as well as the distribution of Tremarctos led the authors to favor a placement in Felidae. [1] The humerus preserves a combination of pantherine and machairodontine features, such as the entepicondylar foramen's shape though it is over twice as large as that of Smilodon's. This foramen is missing from humeri of Smilodon populator, [10] [11] leading the authors to speculate that P. balamoides was a pantherine. This, in addition to characteristics like the olecranon fossa's outline and size of the supracondylar process, convinced Stinnesbeck et al. (2019) to put the species in Panthera. They did note, however, that additional material was required to make a solid phylogenetic assessment. [1]
However, a 2019 study on Yucatán carnivorans suggested that Panthera balamoides may actually be misidentified remains of Arctotherium (a tremarctine bear), whose remains have also been found in Yucatán. If so, this would explain the unusual robustness of the bone and render Panthera balamoides an invalid species. [9] [12] The idea of a felid Panthera balamoides has not been repeated in literature, with a study on Mexican jaguar fossils also considering P. balamoides to be an ursid based on morphological characteristics and mentioned by several other works. [13] [14] A 2023 study also agreed that P. balamoides has nothing in common with Panthera, and that it is morphologically similar to Arctotherium. [12]
Based on the thickness of the cortical bone in the humerus, it was hypothesized by Stinnesbeck et al. that P. balamoides had powerful forelimbs akin to Smilodon that could be used for wrestling its prey. [1] [15] This has been theorized to have reduced the range of humeral abduction and greatened the amount of lateral rotation, indicating an arboreal (climbing) lifestyle. [16] [1] The greater ability of movement in the forelimbs made activities like short-distance running, jumping, climbing, and wrestling possible, an unusual trait for such a large animal. [1] This is in contrast to other pantherines, which bear a thicker cortical bone for a more pursuit-predator lifestyle. [1] [15] The olecranon fossa is short and shallow, suggesting a larger degree of elbow and arm extension was capable, a trait beneficial to arboreal animals. Several other characteristics of the humerus support the idea of arboreal or scansorial habits, features that led Stinnesbeck et al. to suggest that P. balamoides was an accelerator and competent jumper with the ability to climb tall rocks. This is more similar to smaller felines like jaguars, ocelots, and jaguarundis instead of larger pantherines. The arboreal habits may have led P. balamoides to occupy more dense habitats rather than the open habitats which featured cursorial felids like Homotherium, Panthera atrox, and Smilodon. [1] [17] [18] [15]
P. balamoides is known from a single site in the Yucatán dating to the Rancholabrean NALMA (North American Land Mammal Age) from around 13,000 BP, this being at the end of the Pleistocene and reign of many megafauna groups in the Americas. [1] [9] The areas around El Pit cenote likely had large open steppes separating the more densely forested shrubs on the Yucatán savannahs, explaining why a mix of cursorial and arboreal animal fossils have been collected from the locality. [1] The El Pit cenote is one of several cenotes known from the Pleistocene Yucatán, with the Hoyo Negro system that is also submerged being very fossil productive. [14] [9] [4] [2] This includes fossils of humans, [19] the gomphothere proboscidean Cuvieronius, ground sloths Nohochichak [20] and two species of Xibalbaonyx , [4] [21] large canid Protocyon , other fossil felids Panthera atrox and Smilodon in addition to the extant Puma and Lynx , [9] [13] procyonid Nasua , tayassuid Pecari , tapir Tapirus bairdii , and the giant ursid Arctotherium. [8] [22] It was postulated by Stinnesbeck et al. (2019) that P. balamoides used the caves for water as the densely forested environments it inhabited were devoid of drinking spots. [1]
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 term "cat" refers both to felids in general and specifically to the domestic cat.
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.
Smilodon is a genus of felids belonging to the extinct subfamily Machairodontinae. 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. 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.
The 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 Yucatán Peninsula is a large peninsula in southeast Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north and west of the peninsula from the Caribbean Sea to the east. The Yucatán Channel, between the northeastern corner of the peninsula and Cuba, connects the two bodies of water.
Thylacosmilus is an extinct genus of saber-toothed metatherian mammals that inhabited South America from the Late Miocene to Pliocene epochs. Though Thylacosmilus looks similar to the "saber-toothed cats", it was not a felid, like the well-known North American Smilodon, but a sparassodont, a group closely related to marsupials, and only superficially resembled other saber-toothed mammals due to convergent evolution. A 2005 study found that the bite forces of Thylacosmilus and Smilodon were low, which indicates the killing-techniques of saber-toothed animals differed from those of extant species. Remains of Thylacosmilus have been found primarily in Catamarca, Entre Ríos, and La Pampa Provinces in northern Argentina.
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.
Sistema Ox Bel Ha is a cave system in Quintana Roo, Mexico. It is the longest explored underwater cave in the world and ranks second including dry caves. As of January 2023 the surveyed length is 435.8 kilometers (270.8 mi) of underwater passages. There are more than 150 cenotes in the system.
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.
The Tremarctinae or short-faced bears is a subfamily of Ursidae that contains one living representative, the spectacled bear of South America, and several extinct species from four genera: the Florida spectacled bear, the North American giant short-faced bears Arctodus, the South American giant short-faced bear Arctotherium as well as Plionarctos(P. edensis and P. harroldorum), which is thought to be ancestral to the other three genera. Of these, the giant short-faced bears may have been the largest ever carnivorans in the Americas. The group is thought to have originated in eastern North America, and then invaded South America as part of the Great American Interchange. Most short-faced bears became extinct at the end of the Pleistocene.
Eve of Naharon is the skeleton of a 20– to 25-year-old human female found in the Naharon section of the underwater cave Sistema Naranjal in Mexico near the town of Tulum, around 80 miles (130 km) south west of Cancún. The Naranjal subsystem is a part of the larger Sistema Ox Bel Ha. The skeleton is carbon dated to 13,600 years ago, which makes it one of the oldest documented human finds in the Americas.
Arctodus is an extinct genus of short-faced bear that inhabited North America during the Pleistocene. There are two recognized species: the lesser short-faced bear and the giant short-faced bear. Of these species, A. simus was larger, is known from more complete remains, and is considered one of the most charismatic of North America's megafauna. A. pristinus was largely restricted to the Early Pleistocene of the eastern United States, whereas A. simus had a broader range, with most finds being from the Late Pleistocene of the United States, Mexico and Canada. A. simus evolved from A. pristinus, but both species likely overlapped in the Middle Pleistocene. Both species are relatively rare in the fossil record.
Panthera onca mesembrina, also known as the Patagonian panther, is an extinct subspecies of jaguar 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.
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.
Panthera zdanskyi is an extinct pantherine species, the fossils of which were excavated in Gansu Province, northwestern China. Due to its close relationship with the modern tiger, it is called the Longdan tiger.
Ahytherium is an extinct genus of megalonychid sloth that lived during the Pleistocene of what is now Brazil. It contains a single species, A. aureum.
Chan Hol, part of the Toh ha cave system, is a cenote and submerged cave system in Quintana Roo, Mexico, of interest to paleoanthropologists. The remains of three prehistoric human fossils were discovered within the cave system. Along with Eve of Naharon, Naia, the Man of El Templo and the Woman of Las Palmas, the three fossils at Chan Hol are among several ancient Paleo-Indian skeletons found in the submerged cave systems of the Yucatán Peninsula around Tulum, Quintana Roo.
Xibalbaonyx is an extinct genus of megalonychid ground sloth known from the Late Pleistocene of Mexico. Three species are known: X. oviceps and X. exiniferis from the Yucatan peninsula and X. microcaninus from Jalisco. The genus is named after Xibalba, the underworld in Maya mythology.
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