Macrauchenia

Macrauchenia ("long llama", based on the now-invalid llama genus, Auchenia, from Greek "big neck") is an extinct genus of large ungulate native to South America from the late Pliocene to the end of the Pleistocene. It is a member of the extinct order Litopterna, a group of South American native ungulates distinct from the two orders which contain all living ungulates which had been present in South America since the early Cenozoic, 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. The bodyform of Macrauchenia has been described as similar to a camel, being one of the largest-known litopterns, with an estimated body mass of around 1 tonne. The genus gives its name to its family, Macraucheniidae, which like Macrauchenia typically had long necks and three-toed feet, as well as a retracted nasal region, which in Macrauchenia manifests as the nasal opening being on the top of the skull behind the eye sockets. This has historically been argued to correspond to the presence of a tapir-like proboscis, but some recent authors suggest a moose-like prehensile lip or a saiga antelope-like nose to filter dust are more likely.

Only one species is generally considered valid, M. patachonica, which was described by Richard Owen based on remains discovered by Charles Darwin during the voyage of the Beagle. M. patachonica is primarily known from localities in the Pampas, but is known from remains found across the Southern Cone extending as far south as southernmost Patagonia, and as far northeast as Southern Peru. Another genus of macraucheniid Xenorhinotherium was present in northeast Brazil and Venezuela during the Late Pleistocene.

Macrauchenia became extinct as part of the end-Pleistocene extinctions around 12,000 years ago, along with the vast majority of other large mammals native to the Americas.

Taxonomy
Macrauchenia fossils were first collected on 9 February 1834 at Port St Julian in Patagonia (Argentina) by Charles Darwin, when HMS Beagle was surveying the port. As a non-expert he tentatively identified the leg bones and fragments of spine he found as "some large animal, I fancy a Mastodon". In 1837, soon after the Beagle return, the anatomist Richard Owen identified the bones, including vertebrae from the back and neck, as from a gigantic creature resembling a llama or camel, which Owen named Macrauchenia patachonica. In naming it, Owen noted the original Greek terms µακρος (, large or long) and αυχην (, neck), as used by Illiger as the basis of Auchenia as a generic name for the llama, Vicugna and so on. The find was one of the discoveries leading to the inception of Darwin's theory. Since then, more Macrauchenia fossils have been found, mainly in Patagonia, but also in Bolivia, Chile and Venezuela.

The related genus Cramauchenia was named by Florentino Ameghino as a deliberate anagram of Macrauchenia.

Evolution
It is likely that Macrauchenia evolved from earlier litopterns Theosodon, Cramauchenia or Promacrauchenia, or a similar species. Litopterna was one of the five (four in some classifications) ancient orders of endemic South American mammals collectively called meridiungulates. Their relationships with other mammal groups outside South America have been poorly understood, as their early evolutionary history would have been in Western Gondwana, and outside of South America this area is now Antarctica. When South America separated from Antarctica in the Eocene, meridungulate orders survived in South America in isolation. Most flourished in the Paleogene and then diminished. Formerly, North American paleontologists considered them inferior to Northern Hemisphere taxa and to have been outcompeted to extinction in the Great American Biotic Interchange after the establishment of the Central American land bridge. However, more recent evidence shows that three of the meridungulate orders declined long before, just as happened to early mammal groups elsewhere. Litopterns and notoungulates continued, evolving into a variety of more derived forms. While toxodontid notoungulates expanded into North America during the GABI, litopterns remained confined to South America. Macrauchenia was among the last surviving meridungulates, along with litopterns such as Macraucheniopsis, Neolicaphrium and Xenorhinotherium and the large notoungulates Piauhytherium, Trigodonops, Toxodon and Mixotoxodon. These last endemic South American hoofed animals died out at the end of the Lujanian (10,000–20,000 years ago). 

Sequencing of mitochondrial DNA extracted from an M. patachonica fossil from a cave in southern Chile indicates that Macrauchenia (and by inference, Litopterna) is the sister group to Perissodactyla, with an estimated divergence date of sixty-six million years ago. Analysis of collagen sequences obtained from Macrauchenia and Toxodon reached a similar conclusion and extended membership in the sister group clade to notoungulates.

Description
Macrauchenia had a somewhat camel-like body, with sturdy legs, a long neck and a relatively small head. Its feet, however, more closely resembled those of a modern rhinoceros, with one central toe and two side toes on each foot. It was a large animal measuring around 2.5 - 3 m long, 1.8 m tall at the shoulder and weighing up to 909 - 1000 kg, about the size of a black rhinoceros.

One striking characteristic of Macrauchenia is the openings for the nostrils on top of the head, above and between the eyes. Increasingly retracted nostrils are an evolutionary trend in later litopterns. Because mammals with trunks show the nostrils in a similar position, a popular hypothesis is that Macrauchenia had a trunk similar to a tapir or an inflated snout like that of the saiga antelope, perhaps to keep dust out of the nostrils. However, a 2018 study comparing the skulls of tapirs and various other herbivorous extant and extinct mammal species instead saw similarities with the skulls of moose, suggesting that Macrauchenia and other macraucheniids, such as Huayqueriana did not possess trunks. However, pictographs depicting various extinct megafauna dated to around 12,600 to 11,800 years ago from the Serranía de La Lindosa rock formation of Guaviare, Colombia showed what appears to be a possible trunked macraucheniid, presumably Xenorhinotherium.

The snout of Macrauchenia is completely enclosed by bone, and the animal has an elongated neck that allowed it to reach upward; no living mammal with a proboscis has these features. An alternative hypothesis is that these litopterns were high browsers on tough and thorny vegetation, and retracted nostrils allowed them to reach leaves without being impaled in the nose. Sauropods (reconstructed as high browsers on conifer needles and cycads) have similar snouts, and living giraffes and gerenuks, high browsers on thorny vegetation, have more retracted nostrils than related taxa with other feeding habits.

One insight into Macrauchenia habits is that its ankle joints and shin bones may indicate that it was adapted to have unusually good mobility, being able to rapidly change direction when it ran at high speed.

Macrauchenia is known, like its relative Theosodon, to have had a full set of 44 teeth.

Paleobiology
Macrauchenia was a herbivore, likely living on leaves from trees or grasses. Carbon isotope analysis of M. patachonica tooth enamel, as well as analysis of its hypsodonty index (low in this case; i.e., it was brachydont), body size and relative muzzle width suggests that it was a mixed feeder, combining browsing on C3 foliage with grazing on C4 grasses. A dental microwear, occusal enamel, and carbon isotope analysis of Macrauchenia and Xenorhinotherium found that both were grazers on C3 grasses. Evidence from dental calculus has further confirmed the status of M. patachonica as a C3 grazer.

The genus was widespread, found in environments that ranged from dry to humid, from southern Chile to northeastern Brazil and the coast of Venezuela. Fossils of M. ullomensis have been found in Bolivia at altitudes up to 4,000 metres. Habits and diet may have varied depending on the environment, but in plant feeders an elongated neck is usually an adaptation to allow high browsing on trees and shrubs. As the genus was not confined to forest, it was probably able to exploit more marginal environments by mixing high browsing with low browsing and grazing. A site in northern Chile preserved the remains of five subadults associated together, which suggests Macrauchenia may have lived in large herds or family groups.

When Macrauchenia first arose, it could have been preyed upon by the largest of native South American mammal predators, the sabertoothed sparassodont Thylacosmilus. The largest phorusrhacid birds may also have been able to prey on juveniles. After the GABI, the primary predator on adults would have been the very large sabertoothed cat Smilodon populator and giant short-faced bears. Dire wolves and jaguars may also have hunted Macrauchenia, particularly juveniles.

It is presumed that Macrauchenia dealt with its predators primarily by outrunning them, or, failing that, kicking them with its long, powerful legs. The large size of adults would have limited their vulnerability to most predators. Its potential ability to twist and turn at high speed could have enabled it to evade pursuers; both Thylacosmilus and S. populator were ambush hunters likely unable to run down prey over distance if the prey evaded the first attack.

Distribution
Fossils of Macrauchenia have been found in:


 * Miocene
 * Epecuén Formation, Argentina


 * Pliocene
 * Argentina


 * Pleistocene


 * Arroyo Seco and Luján Formations, Argentina
 * Ulloma, Charana, Umala, Tarija and Ñuapua Formations, Bolivia
 * Touro Passo Formation, Brazil
 * Cueva del Milodón, Chile
 * Paraguay
 * San Sebastián Formation, Peru
 * Sopas and Dolores Formations, Uruguay
 * Taima-Taima, Venezuela