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Toxodon

Toxodon is an extinct genus of large ungulate native to South America from the Pliocene to the end of the Late Pleistocene. Toxodon is a member of Notoungulata, an order of extinct South American native ungulates distinct from the two living ungulate orders that had been indigenous to the continent for over 60 million years since the early Cenozoic, prior to the arrival of living ungulates into South America around 2.5 million years ago during the Great American Interchange. Toxodon is a member of the family Toxodontidae, which includes medium to large sized herbivores. Toxodon was one of the largest members of Toxodontidae and Notoungulata, with Toxodon platensis having an estimated body mass of 1,000–1,200 kilograms (2,200–2,600 lb).

Taxonomy and evolution
Charles Darwin, who was in South America as part of the second voyaging expedition of HMS Beagle, was one of the first to collect Toxodon fossils. In his book covering the expedition, The Voyage of the Beagle. Darwin wrote, "November 26th – I set out on my return in a direct line for Montevideo. Having heard of some giant's bones at a neighbouring farm-house on the Sarandis, a small stream entering the Rio Negro, I rode there accompanied by my host, and purchased for the value of eighteen pence the head of the Toxodon." The skull had been propped up against a fence and been used as target practice for throwing stones by local children, who had knocked out its teeth. Since Darwin discovered that the fossils of similar mammals of South America were different from those in Europe, he invoked many debates about the evolution and natural selection of animals. In his own words, Darwin wrote down in his journal, Toxodon and its type species, T. platensis, were described in 1837 by Richard Owen based on remains collected by Darwin, in a paper titled "A description of the cranium of the Toxodon platensis, a gigantic extinct mammiferous species, referrible by its dentition to the Rodentia, but with affinities to the Pachydermata and the herbivorous Cetacea", reflecting the many unusual characteristics of its anatomy. Evolution Toxodon is a member of Notoungulata, a group of South American native ungulates that had been part of the fauna of South America since the Paleocene, over 60 million years ago, and had evolved in isolation in South America, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the Great American Interchange. Notoungulates are morphologically diverse, including forms morphologically distant from Toxodon such as rodent and rabbit-like forms. This finding has been corroborated by an analysis of mitochondrial DNA extracted from a Macrauchenia fossil, which yielded a date of 66 million years ago for the time of the split from perissodactyls. Toxodon belongs to Toxodontidae, a large bodied group of notoungulates which first appeared in the Late Oligocene (Deseadan), ~28-23 million years ago, and underwent a great radiation during the Miocene epoch (~23-5.3 million years ago), when they reached their apex of diversity. The diversity of toxodontids, along with other notoungulates began to decline from around the Pliocene onwards, By the Late Pleistocene (Lujanian), the once great diversity of notoungulates had declined to only a few of species of toxodontids (belong to the genera Toxodon, Mixotoxodon, Trigodonops and Piauhytherium, the last possibly being a synonym of Trigodonops) with all other notoungulate families having become extinct. }} Species There has not been a recent taxonomic revision of the genus Toxodon, leaving the number of valid species uncertain. The species Toxodon chapalmalensis is known from the Pliocene (Montehermosan-Chapadmalalan) of Argentina, while Toxodon platensis, the type species, is known from the Pleistocene. The validity of other potential species like Toxodon darwini Burmeister, 1866, and Toxodon ensenadensis Ameghino, 1887 from the Early Pleistocene of Argentina is uncertain, and the species Toxodon gezi C. Ameghino, 1917 and Toxodon aguirrei Ameghino, 1917 have been considered junior synonyms of Toxodon platensis by recent authors. Some recent authors have argued that Toxodon gracilis Gervais and Ameghino, 1880, should be recognised as a distinct species from the Pleistocene of the Pampas significantly smaller than T. platensis, with these authors suggesting that T. platensis and T. gracilis represent the only valid species of Toxodon in the Pleistocene of the Pampas region. Other authors have argued that all Pleistocene Toxodon species should be considered synonymous with T. platensis. == Description ==
Description
The bodyform of Toxodon and other toxodontids have been compared to those of hippopotamuses and rhinoceroses. Toxodon platensis is one of the largest known toxodontids and notoungulates, with an estimated body mass of approximately , and a body length of around . The skull of Toxodon is proportionally large, All of the teeth in the jaws are high-crowned (hypsodont). Like other toxodontids, the upper and lower first incisors (I1 and i1) are large and protrude, with the second upper incisors (I2) and lower third incisors (i3) being modified into evergrowing tusks. The upper incisors display an arched shape, while the lower incisors project horizontally forwards at the front of the lower jaw. Like other derived toxodontids, Toxodon had long, ever-growing (hypselodont) cheek (premolar and molar) teeth, with the name Toxodon deriving from the curved shape of the upper molars, which are bowed inwards towards the midline of the skull to fit in the upper jaw. Evergrowing cheek teeth are unknown in any living ungulates (though they occur in the extinct rhinoceros Elasmotherium), but are present in some other mammal groups like wombats and rodents. The surface of the cheek teeth is primarily composed of dentine. The thoracic vertebrae of Toxodon have elongate neural spines, which likely anchored muscles and ligaments which supported the large head. The hindlimb is considerably longer than the forelimb. Although Toxodon has been historically reconstructed in museum mounts with bent limbs, the ulna of the forelimb has a strongly backwardly projecting olecranon process similar to that of rhinos, suggesting that the front legs and likely the hind legs were held extended straight beneath the body when standing. There are three functional digits on each foot, == Distribution ==
Distribution
Toxodon had a widespread distribution in South America east of the Andes, ranging from northern Argentina and Bolivia to the western Amazon on the Peru-Brazil border, to Northeast Brazil. == Palaeobiology ==
Palaeobiology
(foreground right), the glyptodonts Glyptotherium and Panochthus, the pampathere Holmesina paulacoutoi (midground centre-left) and the armadillo Pachyarmatherium brasiliense'' (foreground left) Although some authors have suggested that Toxodon was semiaquatic based on the similarity of some aspects of its anatomy to hippopotamuses, this has been disputed by other authors, and analysis of oxygen isotope ratios (which differ between terrestrial and aquatic animals) suggests a terrestrial lifestyle for Toxodon. with its diet varying according to local conditions, with an almost totally C3 browsing diet in the Amazon rainforest, mixed feeding C3 in Bahia and the Pampas, and an almost completely C4 dominated grazing diet in the Chaco. Within the Brazilian Intertropical Region (BIR), T. platensis was a mixed feeder; seasonal variations in the BIR had little impact on the diet of T. platensis. Although Toxodon is thought to have inhabited open landscapes like steppe and savannah, in some areas like the southwestern Brazilian Amazon, it is suggested to have inhabited woodland. Like living animals of similar size, it has been suggested that Toxodon probably only gave birth to a single offspring at a time. T. platensis bones have been found displaying signs of disease like osteomyelitis and spondyloarthropathies. The teeth of Toxodon often display enamel hypoplasia (loss of tooth enamel) in the form of grooves and pits, which is likely due to their evergrowing nature and/or environmental stresses. Tracks probably attributable to Toxodon have been reported from eastern Pernambuco in Northeast Brazil. == Extinction ==
Extinction
Toxodon and the other remaining toxodontids became extinct at the end of the Late Pleistocene around 12,000 years as part of the end-Pleistocene extinction event alongside almost all other large animals in South America. These extinctions followed the first arrival of humans in the Americas, and it has been suggested human hunting may have been a casual factor in the extinctions. At the Paso Otero 5 site in the Pampas of northeast Argentina, burned bones of Toxodon alongside those of numerous other extinct megafauna species are associated with Fishtail points (a type of knapped stone spear point common across South America at the end of the Pleistocene, suggested to be used to hunt large mammals). The bones of the megafauna were probably deliberately burned as fuel. No cut marks are visible on the vast majority of bones at the site (with only one bone of a llama possibly displaying any butchery marks), which may be due to the burning degrading the bones. Various remains of Toxodon platensis in the collection of the ''Museum national d'Histoire naturelle'' collected from the Pampas region in the 19th century including a femur, an iliac fragment, a tibia, as well as a mandible (the latter of which has been radiocarbon dated to around 13,000 years ago), have been found to display cut marks indicative of butchery. == References ==
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