''Cramauchenia'' is an extinct

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...

litoptern Litopterna (from grc, λῑτή πτέρνα "smooth heel") is an extinct order of fossil hoofed mammals from the Cenozoic era. The order is one of the five great orders of South American ungulates that were endemic to the continent, until the ...

South American ungulate. ''Cramauchenia'' was named by

Florentino Ameghino Florentino Ameghino (born Giovanni Battista Fiorino Giuseppe Ameghino September 19, 1853 – August 6, 1911) was an Argentine naturalist, paleontologist, anthropologist and zoologist, whose fossil discoveries on the Argentine Pampas, especiall ...

. The name has no literal translation. Instead, it is an

anagram An anagram is a word or phrase formed by rearranging the letters of a different word or phrase, typically using all the original letters exactly once. For example, the word ''anagram'' itself can be rearranged into ''nag a ram'', also the word ...

of the name of a related genus ''

Macrauchenia ''Macrauchenia'' ("long llama", based on the now-invalid llama genus, ''Auchenia'', from Greek "big neck") was a large, long-necked and long-limbed, three-toed native South American mammal in the order Litopterna. The genus gives its name to its ...

''. This genus was initially discovered in the

Sarmiento Formation The Sarmiento Formation (Spanish: ''Formación Sarmiento''), in older literature described as the Casamayor Formation, is a geological formation in Chubut Province, Argentina, in central Patagonia, which spans around 30 million years from the mid- ...

in the

Chubut Province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ran ...

, in

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, t ...

, and later it was found in the

Chichinales Formation The Chichinales Formation is a geological formation in Río Negro Province, Argentina which dates from the Late Oligocene to the Early Miocene, around 23 to 17.5 million years ago. It predominantly consists of pyroclastic deposits, which were depo ...

in the

Río Negro Province Río Negro (, ''Black River'') is a province of Argentina, located in northern Patagonia. Neighboring provinces are from the south clockwise Chubut, Neuquén, Mendoza, La Pampa and Buenos Aires. To the east lies the Atlantic Ocean. Its c ...

and the

Cerro Bandera Formation The Cerro Bandera Formation is a geological formation in Neuquén Province, Argentina, in northern Patagonia, which dates to the Early Miocene, around 21 to 17.5 million years ago. It predominantly consists of pyroclastic deposits, which were dep ...

Neuquén Neuquén (; arn, Nehuenken) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rive ...

, also in Argentina, in sediments assigned to the SALMA

Colhuehuapian The Colhuehuapian age is a period of geologic time (21.0–17.5 Ma) within the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Deseadan and precedes the Santacrucian age. ...

(in the Early Miocene), as well as the Agua de la Piedra Formation in Mendoza, in sediments dated to the

Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fol ...

(during the Late Oligocene). In 1981 Soria made ''C. insolita'' a junior synonym of ''C. normalis''. A specimen of ''C. normalis'' was described in 2010 from Cabeza Blanca (Chubut, Argentina) in the Sarmiento Formation, in sediments assigned to the

SALMA (Upper Oligocene).

Description

This animal had an appearance vaguely similar to that of a small

llama The llama (; ) (''Lama glama'') is a domesticated South American camelid, widely used as a meat and pack animal by Andean cultures since the Pre-Columbian era. Llamas are social animals and live with others as a herd. Their wool is so ...

or perhaps that of a stocky

antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mamm ...

. The

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...

of this animal was relatively elongated and provided with a slightly recessed nasal opening, which would indicate the presence of a strong, muscular lip, likely prehensile. In similar but larger and more recent forms, such as '' Theosodon'' and '' Scalabrinitherium'', this lip gradually developed, eventually giving rise to a possibly proboscis-like structure with ''

''.

Taxonomy

''Cramauchenia'' was first described by

1902 Events January * January 1 ** The Nurses Registration Act 1901 comes into effect in New Zealand, making it the first country in the world to require state registration of nurses. On January 10, Ellen Dougherty becomes the world's ...

, based on fossils found in the Sarmiento Formation of Argentina. It is a primitive representative of the

Macraucheniidae Macraucheniidae is a family in the extinct South American ungulate order Litopterna, that resembled various camelids. The reduced nasal bones of their skulls was originally suggested to have housed a small proboscis, similar to that of the sa ...

, a group of South American mammals belonging to the Litopterna, with forms similar to those of

camelids Camelids are members of the biological family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant members of this group are: dromedary camels, Bactrian camels, wild Bactrian camels, llamas, alpacas, vicuñas, ...

, in spite not being closely related. ''Cramauchenia'' is known for the sole species ''C. normalis'' another species, ''C. insolita,'' was initially described as a separate species, but following a study by Soria 1981 it has since been attributed to the type species. Furthermore, ''Cramauchenia'' has been assigned to the Cramaucheniinae, a subfamily including the most basal macraucheniids, however, many recent studies tend to indicate that the subfamily is

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

, with ''Cramauchenia'' being most closely related to ''

Pternoconius ''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. Etymology The genus name, ''Pternoconius'', ...

''. The following position of the

is based on McGrath ''et al.'' 2018, showing the position of ''Cramauchenia''.

Paleoenvironment

Fossils of ''Cramauchenia'' have been found in various fossiliferous stratigraphic units in South America. Several specimens come from the

in the

Golfo San Jorge Basin The Golfo San Jorge Basin ( es, Cuenca del Golfo San Jorge) is a hydrocarbon-rich sedimentary basin located in eastern Patagonia, Argentina. The basin covers the entire San Jorge Gulf and an inland area west of it, having one half located in Sant ...

in central

Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and ...

, with other finds from the Agua de la Piedra, Cerro Bandera, and the

of the

Neuquén Basin Neuquén Basin ( es, Cuenca Neuquina) is a sedimentary basin covering most of Neuquén Province in Argentina. The basin originated in the Jurassic and developed through alternating continental and marine conditions well into the Tertiary. The ba ...

. In the Chichinales Formation, which is known for its local mammal fauna, ''Cramauchenia'' would have coexisted with

astrapotheres 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 R ...

, the notoungulates '' Cochilius volvens'', ''

Colpodon ''Colpodon'' is an extinct genus of herbivorous mammal, belonging to the order Notoungulata. It lived during the Early Miocene, in what is today Argentina and Chile, in South America. Description This animal is almost exclusively known from c ...

'', '' Hegetotheriopsis'', '' Hegetotherium'' and '' Protypotherium'', the

rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...

'' Australoprocta'', '' Caviocricetus'', '' Eoviscaccia'', and '' Willidewu'', the

armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, al ...

s ''

Proeutatus ''Proeutatus'' is an extinct genus of xenarthran, belonging to the order Cingulata. It lived during the Early Miocene, and its fossilized remains were discovered in South America. Description This animal was vaguely similar to the extant six ...

'' and ''

Stenotatus ''Stenotatus'' is an extinct genus of cingulate, belonging to the family Dasypodidae. It lived from the Early to the Late Miocene in South America. Description Stenotatus was a small to medium-sized armadillo, not exceeding 4 kilograms in weigh ...

'', and the

sparassodont Sparassodonta (from Greek to tear, rend; and , gen. , ' tooth) is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now tho ...

'' Cladosictis''. Bird remains from the formation are comparatively poor. A part of a

tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These sm ...

has previously been classified as an undetermined species of psilopterine phorusrhacid. Other birds include an undetermined

wading bird 245px, A flock of Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflats in order to foraging, ...

, '' Opisthodactylus horacioperezi'', a species of rhea, and '' Patagorhacos'', a phorusrhacid. During the Miocene the area likely consisted of open but wooded environment with temperate climate and a proximity to freshwater. The Sarmiento Formation has provided a wide assemblage of mammals, including the astrapotheres ''

Astrapotherium ''Astrapotherium'' ("lightning beast") is an extinct genus of South American mammals that vaguely resembled a small elephant or large tapir. However, it was unrelated to elephants or tapirs, but was instead related to other extinct South America ...

'' and ''

Parastrapotherium ''Parastrapotherium'' is an extinct genus of South American land mammal that existed from the Late Oligocene (Deseadan SALMA) to the Early Miocene (Colhuehuapian SALMA). The genus includes some of the largest and smallest known astrapotherian, bu ...

'', the notoungulates '' Argyrohippus'', ''

Cochilius ''Cochilius'' is an extinct genus of interatheriid notoungulate that lived between the Late Oligocene and the lower Miocene in what is now Argentina. Description The skull and skeleton show features also found in other similar contemporary o ...

'', ''

'', '' Interatherium'', '' Pachyrukhos'' and '' Protypotherium'', the fellow litopterns ''

Lambdaconus ''Lambdaconus'' is a genus of proterotheriid from the Late Oligocene to Early Miocene of Argentina. The type species is ''L. suinus'', named in 1897 by Ameghino, with referred species including ''L. lacerum'', named as ''Proterotherium ''Prot ...

'', ''

Paramacrauchenia ''Paramacrauchenia'' is an extinct genus of proterotheriid litopterns from the Early Miocene of what is now Argentina and Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the souther ...

'', '' Proheptaconus'', ''

Prolicaphrium ''Prolicaphrium'' is an extinct genus of proterotheriid litoptern that lived during the Early Miocene, in what is now Argentina. Fossils have been found in the Sarmiento Formation of Argentina. Description This animal may have been vaguely si ...

'', ''

'', '' Tetramerorhinus'' and '' Theosodon'', the

xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Ex ...

ns '' Hapaloides'', '' Holomegalonyx'', '' Nematherium'', '' Peltephilus'', ''

'', '' Proschismotherium'', ''

Prozaedyus ''Prozaedyus'' is an extinct genus of chlamyphorid armadillo that lived during the Middle Oligocene and Middle Miocene in what is now South America. Description It was a small-sized animal, and its life appearance was probably similar to th ...

'', '' Stegotherium'', and ''

'', the

metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as we ...

ns '' Acyon'', '' Acrocyon'', '' Arctodictis'', ''

Borhyaena ''Borhyaena'' is an extinct genus of South American metatherian, living between 17.5 and 15.5 million years ago in Patagonia, Argentina ( Santa Cruz and Sarmiento Formations) and Chile (Río Frias Formation).Cladosictis'' '' Palaeothentes'', ''

'', and '' Sipalocyon'', the rodents '' Acarechimys'', '' Acaremys'', '' Caviocricetus'', '' Eosteiromys'', '' Eoviscaccia'', '' Hypsosteiromys'', '' Neoreomys'', '' Paradelphomys'', '' Parasteiromys'', '' Perimys'', '' Prospaniomys'', '' Prostichomys'', '' Protacaremys'', '' Protadelphomys'', '' Sarremys'' and '' Soriamys'', and the

primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...

s ''

Homunculus A homunculus ( , , ; "little person") is a representation of a small human being, originally depicted as small statues made out of clay. Popularized in sixteenth-century alchemy and nineteenth-century fiction, it has historically referred to the ...

'', '' Mazzonicebus'' and '' Tremacebus''. The late-surviving

meridiolestida Meridiolestida is an extinct clade of mammals known from the Cretaceous and Cenozoic of South America and possibly Antarctica. They represented the dominant group of mammals in South America during the Late Cretaceous. Meridiolestidans were morp ...

n ''

Necrolestes ''Necrolestes'' ("grave robber" or "thief of the dead") is an extinct genus of mammals, which lived during the Early Miocene in what is now Argentine Patagonia. It is the most recent known genus of Meridiolestida, an extinct group of mammals mo ...

'' was also present.

References

External links

* {{Taxonbar, from=Q3753628 Macraucheniids Miocene mammals of South America Oligocene mammals of South America Colhuehuapian Deseadan Neogene Argentina Paleogene Argentina Fossils of Argentina Fossil taxa described in 1902 Taxa named by Florentino Ameghino Prehistoric placental genera Golfo San Jorge Basin Neuquén Basin Sarmiento Formation Cerro Bandera Formation Chichinales Formation