Macrauchenia

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Macrauchenia
Temporal range: Middle Pleistocene – Late Pleistocene
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Macrauchenia Holotype AMNH.jpg
Skeleton of M. patachonica (larger) and Phenacodus primaevus (smaller) at American Museum of Natural History
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order:† Litopterna
Family:† Macraucheniidae
Subfamily:† Macraucheniinae
Genus:† Macrauchenia
Owen, 1838
Type species
†Macrauchenia patachonica
Owen, 1838
Macrauchenia range map.svg
Map showing the distribution of Macrauchenia in red, and Xenorhinotherium in yellow, inferred from fossil finds

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 Pliocene [1] or Middle Pleistocene to the end of the Late Pleistocene. [2] 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. [3] The bodyform of Macrauchenia has been described as similar to a camel, [4] being one of the largest-known litopterns, with an estimated body mass of around 1 tonne. [3] 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, [5] which in Macrauchenia manifests as the nasal opening being on the top of the skull between the eye sockets. [6] This has historically been argued to correspond to the presence of a tapir-like proboscis, though recent authors suggest a moose-like prehensile lip [7] or a saiga antelope-like nose to filter dust are more likely.

Contents

Only one species is generally considered valid, [8] M. patachonica, which was described by Richard Owen based on remains discovered by Charles Darwin during the voyage of the Beagle. [9] 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 north as Southern Peru. Another genus of macraucheniid Xenorhinotherium was present in northeast Brazil and Venezuela during the Late Pleistocene. [6]

Macrauchenia is thought to have been a mixed feeder that both consumed woody vegetation and grass that lived in herds and probably engaged in seasonal migrations. Macrauchenia is suggested to have been a swift runner that was capable of moving at considerable speed.

Macrauchenia became extinct as part of the end-Pleistocene extinction event around 12,000 years ago, along with the vast majority of other large mammals native to the Americas. [3] This followed the arrival of humans to the Americas, and possible evidence of human interactions with Macrauchenia has been found at a number of sites with some authors suggesting human hunting may have played a role in its extinction.

Taxonomy

An outdated historical 1913 restoration by Robert Bruce Horsfall depicting M. patachonica with an elephantine trunk Macrauchenia.jpg
An outdated historical 1913 restoration by Robert Bruce Horsfall depicting M. patachonica with an elephantine trunk

Macrauchenia fossils were first collected on 9 February 1834 at Port St Julian in southern Patagonia in what is now Argentina by Charles Darwin, when HMS Beagle was surveying the port (the Argentine Confederation claimed the region but did not effectively control it at the time). [10] 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's 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. [11] In naming it, Owen noted the original Greek terms µακρος (makros, large or long) and αυχην (auchèn, neck), as used by Illiger as the basis of Auchenia as a generic name for the llama, Vicugna and so on. [12]

Macrauchenia patachonica is currently considered to be the only valid species of Macrauchenia. [13] Macrauchenia boliviensis from the probably early Miocene aged Kollukollu Formation of Bolivia described by Thomas Henry Huxley in 1860 is now considered to be an indeterminate member of Macraucheniidae. [14] The species Macrauchenia ensenadensis described by Florentino Ameghino in 1888 from the Early Pleistocene has been transferred to the closely related genus Macraucheniopsis . [15]

Evolution

Macrauchenia is part of the extinct ungulate order Litopterna, which is grouped with several other orders as part of the South American native ungulates (SANUs), which formed a conspicuous element of South America's Cenozoic mammal fauna beginning during the Paleocene, over 60 million years ago. [16] Litopterns generally have body forms similar to those of living ungulates. [16]

The relationships of litopterns (as well as other SANUs) to living mammals was historically uncertain. Sequences of mitochondrial DNA extracted from remains of M. patachonica found in a cave in southern Chile published in 2017 indicates that the closest living relatives of Macrauchenia (and by inference, Litopterna) are members of the extant ungulate order Perissodactyla (which includes the equids, rhinoceroses, and tapirs), with litopterns estimated to have genetically diverged from perissodactyls around 66 million years ago. [17] [18] Analysis of collagen sequences obtained from Macrauchenia and the rhinoceros-like Toxodon (which belongs to another SANU order, Notoungulata) in 2015 reached a similar conclusion and suggests that litopterns are more closely related to notoungulates than to perissodactyls. [19] [20]

The earliest known fossils of litopterns are from the early Paleocene, around 62.5 million years ago. [1] The family to which Macrauchenia belongs, Macraucheniidae, first appeared during the Late Eocene or Oligocene, around 39-30 million years ago, depending on what species are included. [1] [21] Members of the family are typically characterised by having three-toed feet and long necks. [21] The family reached its apex of diversity in the Late Miocene, around 10-6 million years ago, before declining to low diversity during the Pliocene and Pleistocene, [21] as part of a broader decline of SANU diversity during this period. [16] The cause of this diversity decline is uncertain, though it has been suggested to be due to climatic changes, [22] as well as possibly competiton/predation from immigrants from North America, who arrived following the formation of the Isthmus of Panama during the Pliocene as part of an event called the Great American Interchange. [16] [23] The earliest fossils attributed to Macrauchenia date to the late Pliocene, [5] though remains of Macrauchenia patagonica are primarily known from the Late Pleistocene. [15]


Cladogram of Macraucheniidae after Lobo, Gelfo & Azevedo (2024): [21]

Macraucheniidae

Description

Size of M. patachonica compared to a human Macrauchenia wiki.png
Size of M. patachonica compared to a human
Skeletal reconstruction of M. patachonica Macrauchenia patachonica Skeletal (cropped).png
Skeletal reconstruction of M. patachonica

Macrauchenia had a bodyform superficially like a camel, [24] with a long neck composed of camel or giraffe-like elongated cervical vertebrae. In most of the cervical vertebrae, the canal for the artery passes through the neural arch. [25] Macrauchenia was one of the largest macraucheniids and South American native ungulates, with an estimated body mass of around 1,000 kilograms (2,200 lb), [25] considerably larger than earlier macraucheniids, which generally only weighed around 80–120 kilograms (180–260 lb). [3]

Skull

Illustration of the skull of Macrauchenia Anales del Museo Publico de Buenos Aires (1864) (17980766940).jpg
Illustration of the skull of Macrauchenia

The skull of Macrauchenia is relatively elongate and has an eye socket (orbit) entirely enclosed by bone, which is situated behind the teeth. [25] Like other macraucheniids, there are a total of 44 teeth in the upper and lower jaws (the primitive number in placental mammals). [21] The teeth form a continuous row on both jaws without any diastema (gaps) and they are all brachydont (low crowned). [25] The most unusual feature of Macrauchenia's skull is its retracted nasal region, shared with other derived macraucheniines, which have the opening on the top of the skull roof between the eyesockets. [25] [26] Behind the nasal opening there is a substantially depressed region with numerous pits and ridges, which served as attachments for the nasal muscles. While historically this unusual nasal structure was taken as evidence for a tapir-like probiscis/trunk, recent authors have expressed doubts about this, alternatively suggesting that it may have instead formed a moose-like prehensile lip, or a saiga antelope-like nasal structure which served to filter dust (which was likely prevalent in the environment where Macrauchenia lived), [25] perhaps combining the function of dust filtering organ and a prehensile lip. [27] Behind the nasal opening, the top of the skull shows the development of extensive sinuses. [28]

Limbs

Limb bones of M. patachonica Machrauchenia patachonica, liptoptern, Museu de Ciencies Naturals de Valencia.JPG
Limb bones of M. patachonica

The humerus bone is very short and robust. The radius and ulna in the forelimbs and the tibia and fibula in the hindlimbs are fused to each other, with the combined radius-ulna bone being broad in front view, and the fibula is much more slender than the tibia. The femur has a well developed third trochanter, and is long relative to the length of the tibia. The forefeet and hindfeet each had three functional digits. [25] The development of a suprapatellar fossa (an indentation) on the knee joint has led to suggestions that this functioned analogously to the stay apparatus found in living horses, allowing the knees to be passively locked while standing. [29]

Distribution

Fossils of Macrauchenia are known from across the Southern Cone, ranging from northern Chile, southern Peru, southern Bolivia, and the Pampas in Uruguay, northern Argentina and southern Brazil, southwards to extreme southernmost part of Patagonia in southern Chile and Argentina. Macrauchenia is thought to have primarily inhabited arid, open environments with only scattered woody vegetation. A closely related genus, Xenorhinotherium , inhabited more tropical environments in eastern Brazil and Venezuela. [6]

Paleobiology

Restoration of a running Macrauchenia with a saiga antelope-like nose Macrauchenia patachonica Life Reconstruction.png
Restoration of a running Macrauchenia with a saiga antelope-like nose

Analysis of dental calculus extracted from the teeth of an individual of Macrauchenia suggests that it was a mixed feeder (engaging in both browsing and grazing), with this individual having a diet predominantly consisting of C3 grasses. [30] Dental microwear analysis of another individual also supports grazing being an important part of the diet for Macrauchenia. [6] Like living perissodactyls, litopterns including Macrauchenia were probably hindgut fermenters. [31] A 2022 study suggested that based on the anatomy of the cervical vertebrae Macrauchenia likely held its neck in an erect posture when at rest and browsing, similar to that of a llama, though the neck was highly flexible and able to adopt many postures including being lowered to the ground for feeding, as well as being able to flex side to side. [27] Its elongated neck likely allowed it to efficiently browse vegetation without wasting energy. [25] It has been speculated that Macrauchenia may have sometimes reared up onto its hind legs like a gerenuk when feeding. [27]

Macrauchenia is thought to have probably lived in herds, as evidenced by the finding of at least 3 individuals preserved together at the Kamac Mayu site in Chile, with herding individuals probably moving in coordination. Macrauchenia is suggested to have concentrated on foraging in small areas before swiftly moving on to other feeding areas. It has been suggested that Macrauchenia engaged in long distance seasonal migrations in search of food. [25] Like living animals of similar size, it has been suggested that Macrauchenia probably only gave birth to a single offspring at a time. [32]

A 2020 study suggested that Macrauchenia was a capable and fast runner with fossilised footprints suggesting that its feet were held in a digitigrade stance, with the neck probably being held horizontally when running. The running style of Macrauchenia has been suggested to be similar to that of a saiga antelope or spotted hyenas, with a lack of flexing in the spine. [25] A 2005 study suggested that Macrauchenia may have been adapted to swerving as a strategy of avoiding predators, based on the strength of the limb bones. [33] The morphology of its hindlimbs suggests that they were adapted to rapidly accelerating, which may have been useful for both efficient locomotion and escaping predators. [25] Isotopic analysis suggests that Macrauchenia was regularly consumed as prey by the large sabertooth cat Smilodon populator . [34]

Relationship with humans and extinction

Macrauchenia became extinct as part of the end-Pleistocene extinction event at the end of the Late Pleistocene, around 12-10,000 years ago, along with most large (megafaunal) mammals native to the Americas. The extinctions followed the arrival of humans in the Americas, [35] which in South America occurred at least 14,500 years ago (as evidenced by Monte Verde II in Chile). [36] The causes of the extinction have long been controversial with human hunting and climatic change widely considered to be the most probable causes. [37]

Several potential instances of human interaction with Macrauchenia have been recorded. A left mandible collected from somewhere in the Pampas region in the 19th century in the collections of the Museum national d’Histoire naturelle in France has been suggested to display cut marks caused by human butchery, probably to extract the tongue. [38] At Arroyo Seco 2 near Tres Arroyos in the Pampas in Argentina, bones of Macrauchenia amongst those of other megafauna were found associated with human artifacts dating to approximately 14,782–11,142 calibrated years Before Present. While some megafauna remains at the site show clear evidence of exploitation, those of Macrauchenia do not, perhaps because post-depositional degradation of the bones may have erased cut marks. [39] [40] At the El Guanaco site in the Argentinean Pampas, remains of Macrauchenia, alongside those of the glyptodont Doedicurus , horses and rhea eggshells are associated with stone tools. [41]

At the Paso Otero 5 site in the Pampas of northeast Argentina, burned bones of Macrauchenia 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 [42] ). 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. [43]

Related Research Articles

<span class="mw-page-title-main">Litopterna</span> Extinct order of hoofed mammals

Litopterna is an extinct order of South American native ungulates that lived from the Paleocene to the end of the Pleistocene-early Holocene around 62.5 million-12,000 years ago, and were also present in Antarctica during the Eocene. They represent the second most diverse group of South American ungulates after Notoungulata. It is divided into nine families, with Proterotheriidae and Macraucheniidae being the most diverse and last surviving families.

<i>Toxodon</i> Extinct genus of notoungulates

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).

<span class="mw-page-title-main">Notoungulata</span> Extinct order of hoofed mammals

Notoungulata is an extinct order of ungulates that inhabited South America from the early Paleocene to the end of the Pleistocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resembling animals as disparate as rabbits and rhinoceroses. Notoungulata are the largest group of South American native ungulates, with over 150 genera in 14 families having been described, divided into two major subgroupings, Typotheria and Toxodontia. Notoungulates first diversified during the Eocene. Their diversity declined from the late Neogene onwards, with only the large toxodontids persisting until the end of the Pleistocene, perishing as part of the Late Pleistocene megafauna extinctions along with most other large mammals across the Americas. Collagen sequence analysis suggests that notoungulates are closely related to litopterns, another group of South American ungulates, and their closest living relatives being perissodactyls, including rhinoceroses, tapirs and equines as part of the clade Panperissodactyla. However their relationships to other South American ungulates are uncertain. Several groups of notoungulates separately evolved ever-growing cheek teeth.

<span class="mw-page-title-main">South American native ungulates</span> Extinct clade of mammals

South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals that were indigenous to South America from the Paleocene until the end of the Late Pleistocene. They represented a dominant element of South America's Cenozoic terrestrial mammal fauna prior to the arrival of living unguate groups in South America during the Pliocene and Pleistocene as part of the Great American Interchange. They comprise five major groups conventionally ranked as orders—Astrapotheria, Litopterna, Notoungulata, Pyrotheria, and Xenungulata—as well as the primitive "condylarth" groups Didolodontidae and Kollpaniinae. It has been proposed that some or all of the members of this group form a clade, named Meridiungulata, though the relationships of South American ungulates remain largely unresolved. The two largest groups of South American ungulates, the notoungulates and the litopterns, were the only groups to persist beyond the mid Miocene. Only a few species of notoungulates and litopterns survived until the end-Pleistocene extinction event around 12,000 years ago where they became extinct with most other large mammals in the Americas, shortly after the first arrival of humans into the region.

<span class="mw-page-title-main">Macraucheniidae</span> Family in the extinct South American ungulate order Litopterna, that resembled various camelids

Macraucheniidae is a family in the extinct South American ungulate order Litopterna, that resembled camelids. They had three functional digits on the fore and hind feet, as well as elongate necks. The family is generally divided up into two subfamilies, Cramaucheniinae and Macraucheniinae. The family shows retraction of the nasal region, most extremely to the top of the skull in derived macraucheniine taxa like Macrauchenia. which has been interpreted to have supported a probsocis, perhaps like that of a saiga antelope to filter dust, or a moose-like prehensile lip. The earliest unambiguous members of the family date to the late Oligocene around 30 million years ago. Polymorphis from the Eocene has historically been placed as a macraucheniid, but this has been doubted. Most early representatives had a body masses in the range of 80–120 kilograms (180–260 lb), though some like Llullataruca were as small as 35–55 kilograms (77–121 lb), and the last representatives of the family from the Pleistocene like Macrauchenia were over 1,000 kilograms (2,200 lb). The family reached its apex of diversity during the late Miocene around 10-6 million years ago, before declining to only a few species belong to the genera Macrauchenia and Xenorhinotherium by the Late Pleistocene.

<i>Theosodon</i> Extinct genus of litopterns

Theosodon is an extinct genus of litoptern mammal from the Early to Middle Miocene of South America.

<i>Xenorhinotherium</i> Extinct genus of litopterns

Xenorhinotherium is an extinct genus of macraucheniine macraucheniids, native to northern South America during the Pleistocene epoch, closely related to Macrauchenia of Patagonia. The type species is X. bahiense.

<i>Cramauchenia</i> Extinct genus of litoptern South American ungulate

Cramauchenia is an extinct genus of litoptern South American ungulate. Cramauchenia was named by Florentino Ameghino. The name has no literal translation. Instead, it is an anagram of the name of a related genus Macrauchenia. This genus was initially discovered in the Sarmiento Formation in the Chubut Province, in Argentina, and later it was found in the Chichinales Formation in the Río Negro Province and the Cerro Bandera Formation in Neuquén, also in Argentina, in sediments assigned to the SALMA Colhuehuapian, as well as the Agua de la Piedra Formation in Mendoza, in sediments dated to the Deseadan. In 1981 Soria made C. insolita a junior synonym of C. normalis. A specimen of C. normalis was described in 2010 from Cabeza Blanca in the Sarmiento Formation, in sediments assigned to the Deseadan SALMA.

<i>Scalabrinitherium</i> Extinct genus of litopterns

Scalabrinitherium is an extinct genus of mammals of the family Macraucheniidae. Fossils of this animal were found among the fossils of prehistoric xenarthrans in the Ituzaingó Formation of Argentina.

<span class="mw-page-title-main">Astrapotheria</span> Extinct order of mammals

Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, 59 to 11.8 million years ago. Astrapotheres were large, rhinoceros-like animals and have been called one of the most bizarre orders of mammals with an enigmatic evolutionary history.

<span class="mw-page-title-main">Toxodontidae</span> Extinct family of notoungulate mammals

Toxodontidae is an extinct family of notoungulate mammals, known from the Oligocene to the Holocene of South America, with one genus, Mixotoxodon, also known from the Pleistocene of Central America and southern North America. Member of the family were medium to large-sized, ranging from around 350–400 kilograms (770–880 lb) in Nesodon to 1,000–1,200 kilograms (2,200–2,600 lb) in Toxodon, and had medium to high-crowned dentition, which in derived members of the group evolved into ever-growing cheek teeth. Isotopic analyses have led to the conclusion that Pleistocene members of the family were flexible mixed feeders.

<span class="mw-page-title-main">Proterotheriidae</span> Extinct family of litopterns

Proterotheriidae is an extinct family of litoptern ungulates known from the Eocene-Late Pleistocene of South America. Members of the group were small-medium sized cursorial herbivores with brachydont teeth, with their toes showing progressive reduction, with later members of the group bearing weight on a single large toe similar to living horses.

<i>Neolicaphrium</i> Extinct genus of ungulate mammal

Neolicaphrium is an extinct genus of ungulate mammal belonging to the extinct order Litopterna. This animal lived from the Late Pliocene (Chapadmalalan) to the Late Pleistocene (Lujanian) in southern South America, being the last survivor of the family Proterotheriidae.

Cullinia is an extinct genus of litoptern, an order of South American native ungulates that included horse-like and camel-like animals such as Macrauchenia. It is only known from fragmentary remains. Cullinia levis is known from Chasicoan remains found in the Arroyo Chasicó Formation of Argentina, and remains from the Brazilian state of Acre and the Huayquerian Ituzaingó Formation have been assigned to Cullinia sp..

Promacrauchenia is an extinct genus of macraucheniids that lived during the Late Miocene to Late Pliocene epochs of what is now Argentina and Bolivia. It belongs to the subfamily Macraucheniinae, which also includes Huayqueriana, Macrauchenia, and Xenorhinotherium. Fossils of this genus have been found in the Ituzaingó, Andalhuala, and Cerro Azul Formations of Argentina.

Llullataruca is an extinct genus of macraucheniid litoptern. It lived during the Middle Miocene of what is now Bolivia.

Sparnotheriodontidae is an enigmatic extinct family of litopterns known primarily from teeth. Sparnotheriodontids are one of two South American native ungulate clades known to have reached Antarctica, the other being astrapotheres.

Polymorphis is an extinct genus of litopterns belonging to the family Macraucheniidae. It lived during the Middle Eocene of Argentina.

<i>Tetramerorhinus</i> Extinct genus of litopterns

Tetramerorhinus is an extinct genus of proterotheriid litoptern that lived during the Early and Middle Miocene in what is now Argentina and Peru.

Pternoconius is an extinct genus of macraucheniid litoptern from the Late Oligocene and Early Miocene of Argentina. Fossils of this genus have been found in the Sarmiento Formation of Argentina.

References

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  2. ↑ Püschel, Hans P.; Martinelli, Agustín G. (December 2023). "More than 100 years of a mistake: on the anatomy of the atlas of the enigmatic Macrauchenia patachonica". Swiss Journal of Palaeontology . 142 (1): 16. Bibcode:2023SwJP..142...16P. doi: 10.1186/s13358-023-00279-1 . ISSN   1664-2376.
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  6. 1 2 3 4 de Oliveira, Karoliny; Araújo, Thaísa; Rotti, Alline; Mothé, Dimila; Rivals, Florent; Avilla, Leonardo S. (March 2020). "Fantastic beasts and what they ate: Revealing feeding habits and ecological niche of late Quaternary Macraucheniidae from South America". Quaternary Science Reviews. 231: 106178. Bibcode:2020QSRv..23106178D. doi:10.1016/j.quascirev.2020.106178. S2CID   213795563.
  7. ↑ Moyano, Silvana Rocio; Giannini, Norberto Pedro (November 2018). "Cranial characters associated with the proboscis postnatal-development in Tapirus (Perissodactyla: Tapiridae) and comparisons with other extant and fossil hoofed mammals". Zoologischer Anzeiger. 277: 143–147. Bibcode:2018ZooAn.277..143M. doi: 10.1016/j.jcz.2018.08.005 . hdl: 11336/86349 .
  8. ↑ Souza Lobo, Leonardo; Lessa, Gisele; Cartelle, Cástor; and Romano, Pedro S. R. (September 2017). "Dental eruption sequence and hypsodonty index of a Pleistocene macraucheniid from the Brazilian Intertropical Region". Journal of Paleontology. 91 (5): 1083–1090. Bibcode:2017JPal...91.1083S. doi:10.1017/jpa.2017.54. ISSN   0022-3360.
  9. ↑ Fernicola, Juan C., Vizcaino, Sergio F., & De Iuliis, Gerardo (2009). The fossil mammals collected by Charles Darwin in South America during his travels on board the HMS Beagle. Revista De La Asociación Geológica Argentina, 64(1), 147-159. Retrieved from https://revista.geologica.org.ar/raga/article/view/1339
  10. ↑ Darwin, Charles (2001). Keynes, Richard D. (ed.). Charles Darwin's Beagle diary. Cambridge: Cambridge University Press. p. 214. ISBN   9780521003179 . Retrieved 19 December 2008.
  11. ↑ "Darwin Correspondence Project - Letter 238 — Darwin, C. R. to Henslow, J. S., Mar 1834". Archived from the original on 2009-01-16. Retrieved 2008-12-19.
  12. ↑ Owen 1838 , p. 35
  13. ↑ Souza Lobo, Leonardo; Lessa, Gisele; Cartelle, Cástor; and Romano, Pedro S. R. (September 2017). "Dental eruption sequence and hypsodonty index of a Pleistocene macraucheniid from the Brazilian Intertropical Region". Journal of Paleontology. 91 (5): 1083–1090. Bibcode:2017JPal...91.1083S. doi:10.1017/jpa.2017.54. ISSN   0022-3360.
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  15. 1 2 Scherer, Carolina; Pitana, Vanessa; Ribeiro, Ana Maria (2009-12-28). "Proterotheriidae and Macraucheniidae (Litopterna, Mammalia) from the Pleistocene of Rio Grande do Sul State, Brazil" (PDF). Revista Brasileira de Paleontologia. 12 (3): 231–246. doi:10.4072/rbp.2009.3.06.
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  25. 1 2 3 4 5 6 7 8 9 10 11 Blanco, R. Ernesto; Jones, Washington W.; Yorio, Lara; Rinderknecht, Andrés (October 2021). "Macrauchenia patachonica Owen, 1838: Limb bones morphology, locomotory biomechanics, and paleobiological inferences". Geobios. 68: 61–70. Bibcode:2021Geobi..68...61B. doi:10.1016/j.geobios.2021.04.006.
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