astrapotheria

Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, ."The uruguaytheriine Astrapotheriidae from the rich middle Miocene Honda Group of the upper Magdalena River valley in Colombia (...) are the youngest securely dated remains of that order in South America." Astrapotheres were large and rhinoceros-like animals and have been called one of the most bizarre orders of mammals with an enigmatic evolutionary history.

This taxonomy of this order is not clear, but it may belong to Meridiungulata (along with Notoungulata, Litopterna, Pyrotheria and Xenungulata). In turn, Meridungulata is believed to belong to the extant superorder Laurasiatheria. Some scientists have regarded the astrapotheres (and sometimes the Meridiungulata as a whole) as members of the clade Atlantogenata. However, collagen and mitochondrial DNA sequence data analysed in 2015 places at least the notoungulates and litopterns firmly within Laurasiatheria, as a sister group to the perissodactyls.

Description

Their lophodont molars and tusk-like canines became extremely large and ever-growing in later astrapotheres. The upper molars lack an ectocingulum and are dominated by well-developed ectoloph and protoloph. Additional lophs formed in some derived taxa. They had lower molars with two cross-lophs, including a high protocristid, and eventually became almost selenodont. As a result, their dentition is similar to notoungulates, but it seems to have evolved independently. The cheek teeth are similar to rhinocerotoids, including similar microstructure, which indicate they had the same function.

Postcranially, astrapotheres are relatively robust and more or less graviportal but have slender long bones, most notably in the hindlegs, suggesting they were amphibious. In order to support their proboscises and large heads they had relatively long and massive necks in relation to the rest of the vertebral column. Their feet are pentadactyl with short and stout podial and metapodial bones. Most characteristic for the order are the flat astragalus, equipped with a short neck and a flat head, articulating with both the navicular and cuboid bones; and their calcaneus with its enlarged peroneal tubercle.

Three families are recognized: Eoastrapostylopidae from the late Paleocene, Trigonostylopidae from the Paleocene-Eocene, and Astrapotheriidae from the Eocene-Miocene. The Brazilian, Itaboraian ''Tetragonostylops'' and the Argentinian, Riochican '' Eoastrapostylops'' are the oldest astrapotheres. The latter, with its low-crowned and lophoselenodont cheek teeth, is considered the most primitive astrapothere. Trigonostylopids are distinct from other astrapotheres in their ear anatomy but are included in the order because of otherwise similar characters. '' Antarctodon'' is one of few eutherian mammals, as well as one of the last known terrestrial vertebrates, found in Antarctica.

The most famous member of the order is undoubtedly '' Astrapotherium'', a long elephant-like animal that had lost its upper incisors and developed ever-growing canine tusks. They had lost their anterior premolars, resulting in a gap between their tusks and the hypsodont cheek teeth. The short and retracted nasal bones indicate a moderately developed proboscis. The small Eocene '' Trigonostylops'' lacked such retracted nasals and probably also a proboscis. Other astrapotheriids, such as the Casamayoran '' Scaglia'' and ''Albertogaudrya