Toxodon

Toxodon; Temporal range: Pliocene–Holocene PreꞒ Ꞓ O S D C P T J K Pg N
	Skeleton of Toxodon in Buenos Aires
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	†Notoungulata
Family:	†Toxodontidae
Subfamily:	†Toxodontinae
Genus:	†Toxodon; Owen, 1837
Type species
	†Toxodon platensis; Owen, 1837
Other species
	†T. burmeisteri Giebel, 1866; †T. chapalmalensis Ameghino, 1908; †T. ensenadensis Ameghino, 1887; †T. expansidens Cope, 1886; †T. gracilis Gervais and Ameghino, 1880;
Synonyms
	Genus-level Dilobodon Ameghino, 1886; Chapalmalodon Pascual, 1957; Chapadmalodon Tonni et al., 1992 (lapsus calami); T. platensis T. angustidens Owen, 1846; T. owenii Burmeister, 1866; T. gervaisii Gervais & Ameghino, 1880; T. aguirrei Ameghino, 1917; T. gezi Ameghino, 1917; T. burmeisteri T. paradoxus Ameghino, 1882; T. protoburmeisteri Ameghino, 1887; T. bilobidens Ameghino, 1887; T. chapalmalensis Chapalmalodon chapalmalensis Pascual, 1957; T. chapadmalensis Cione & Tonni, 1995 (lapsus calami); T. chapalmalalensis Oliva & Cerdeno, 2007 (lapsus calami); T. ensenadensis T. giganteus Moreno, 1888; T. elongatus Roth, 1898; T. gracilis T. voghti Moreno, 1888;

Toxodon (meaning "bow tooth" in reference to the curvature of the teeth) is an extinct genus of large ungulate native to South America from the Pliocene to the end of the Late Pleistocene.^[1]^[2] 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.^[3] Toxodon is a member of the family Toxodontidae, which includes medium to large sized herbivores.^[4] 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).^[5]

Toxodon has been found across much of South America, excluding southern Patagonia, the Andes and northeastern-most region of the continent.^[6] Evidence suggests that Toxodon was ecologically plastic and able to adapt its diet to local conditions.^[7]

Toxodon became extinct as part of the end-Pleistocene extinctions around 12,000 years ago, along with most large mammals across the Americas. The extinctions followed the arrival of humans to South America, who may have been a contributory factor in the extinctions.^[3]

Taxonomy

Charles Darwin was one of the first to collect Toxodon fossils, after paying 18 pence for a T. platensis skull from a farmer in Uruguay.^[8] In 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."^[9] 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,

Lastly, the Toxodon, perhaps one of the strangest animals ever discovered: In size it equaled an elephant or megatherium, but the structure of its teeth, as Mr. Owen states, proves indisputably that it was intimately related to the Gnawers, the order which, at the present day, includes most of the smallest quadrupeds: In many details it is allied to the Pachydermata: Judging from the position of its eyes, ears, and nostrils, it was probably aquatic, like the Dugong and Manatee, to which it is also allied. How wonderfully are the different Orders, at the present time so well separated, blended together in different points of the structure of the Toxodon!

Toxodon was described in 1837 by Richard Owen based on remains collected by Darwin.^[10]

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, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the Great American Interchange.^[3] Notoungulata represents the most diverse group of indigenous South American ungulates, with over 150 described genera in 13 different families.^[11] Notoungulates are morphologically diverse, including forms morphologically distant from Toxodon such as rodent and rabbit-like forms. Toxodon was a member of Toxodontidae, a large bodied group including vaguely rhinoceros like forms.^[3]

Analysis of collagen sequences obtained from Toxodon as well as from the litoptern (another group of indigenous South American ungulates) Macrauchenia found that notoungulates and litopterns were closely related to each other, and form a sister group to perissodactyls (which contains equids, rhinoceroses and tapirs) as part of the clade Panperissodactyla, making them true ungulates.^[12]^[13] 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.^[14]

Evolution

There has not been a recent taxonomic revision of the genus Toxodon, leaving the number of valid species uncertain.^[15]

The species Toxodon chapalmalensis is known from the Pliocene (Montehermosan-Chapadmalalan) of Argentina,^[16] 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.^[17] Some recent authors have argued that 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.^[15]

In 2014, a study identifying a new species of toxodontid resolved the phylogenetic relations of the toxodontids, including to Toxodon. The below cladogram was found by the study:^[18]

†Notoungulata

†Pampahippus arenalesi

†Rhynchippus spp.

	†Scarrittia canquelensis

	†Leontinia gaudri

†Toxodontidae

†Proadinotherium leptognathum

†Adinotherium spp.

	†Nesodon taweretus

	†Nesodon imbricatus

†Palyeidodon obtusum

†Hyperoxotodon speciosus

	†Nonotherium henningi

	†Xotodon spp.

†Andinotoxodon bolivariensis

	†Dinotoxodon paranensis

	†Toxodon platensis

†Gyrinodon quassus

	†Ocnerotherium intermedium

	†Hoffstetterius imperator

†Posnanskytherium desaguaderoi

†Pisanodon nazari

†Pericotoxodon platignathus

	†Calchaquitherium mixtum

	†Mixotoxodon larensis

	†Paratrigodon euguii

	†Trigodon gaudri

Description

Toxodon was about 2.7 m (8 ft 10 in) in body length, with an estimated weight up to 1,415 kg (3,120 lb)^[19] and about 1.5 m (4 ft 11 in) high at the shoulder and resembled a heavy rhinoceros, with a short and vaguely hippopotamus-like head.^[20] Because of the position of its nasal openings, it is believed that Toxodon had a well-developed snout.^{[citation needed]} Toxodon possessed a large, barrel shaped body. It had short stout legs with plantigrade feet with three functional relatively short toes. The hind limbs are longer and raised higher than the front limbs, giving a sloped appearance to the body.^[21] Like horses, it had a stay apparatus allowing the knees to be passively locked while standing.^[22]

The vertebrae were equipped with high apophyses, which most likely supported the massive weight and muscles as well as its powerful head.^{[citation needed]} Toxodon had broad jaws which were filled with bow shaped teeth and incisors.^[3] The molar teeth of Toxodon have no roots and are ever-growing (euhypsodont), like the incisors of rodents and lagomorphs, and often exhibit enamel hypoplasia.^[23]

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.^[24]

Palaeobiology

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.^[25]

Toxodon is suggested to have been capable of running at considerable speed.^[26] Toxodon is believed to have been ecologically plastic and have had a wide niche breadth,^[7] with its diet varying according to local conditions,^[27] with an almost totally C₃ browsing diet in the Amazon rainforest, mixed feeding C₃ in Bahia and the Pampas to almost completely C₄ dominated grazing diet in the Chaco.^[24] Within the Brazilian Intertropical Region, local climate had little impact on the diet of T. platensis.^[28] Like living animals of similar size, it has been suggested that Toxodon probably only gave birth to a single offspring at a time.^[29]

T. platensis specimens have been found displaying osteological signs of osteomyelitis and spondyloarthropathies.^[30]

Extinction

Toxodon 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. Previous mid-Holocene dates are now thought to be in error.^[31] These extinctions following the first arrival of humans in the Americas. Several sites record apparent interactions between Toxodon and humans. Remains from the Arroyo Seco 2 site in the Pampas are associated with butchered megafauna, but it is unclear if the Toxodon itself was actually butchered or the remains were naturally transported to the site.^[32] 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^[33]). 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.^[34] 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.^[35]