Macrauchenia ("long
llamaThe llama is a South American camelid, widely used as a pack animal by the Incas and other natives of the Andes mountains. In South America llamas are still used as beasts of burden, as well as for the production of fiber and meat....
", based on the now superseded Latin term for llamas
Auchenia, from Greek terms which literally mean "Big Neck") was a long-necked and long-limbed, three-toed
South AmericaSouth America is the southern continent of the Americas, situated entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere...
n
ungulateUngulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...
mammalMammals are a class of vertebrate animals whose females are characterized by the possession of mammary glands while both males and females are characterized by sweat glands, hair, three middle ear bones used in hearing, and a neocortex region in the brain.Mammals are divided into three main...
, typifying the order
LitopternaThe Litopterna is an order of fossil hoofed mammals from the Tertiary period that displays toe reduction. Three-toed, and even a one-toed horselike form developed....
. The oldest fossils date back to around 7 million years ago, and
M. patagonica disappears from the fossil record during the late
PleistoceneThe Pleistocene is the epoch from 2.588 million to 12 000 years BP covering the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
, around 20,000 years ago.
M.
Macrauchenia ("long
llamaThe llama is a South American camelid, widely used as a pack animal by the Incas and other natives of the Andes mountains. In South America llamas are still used as beasts of burden, as well as for the production of fiber and meat....
", based on the now superseded Latin term for llamas
Auchenia, from Greek terms which literally mean "Big Neck") was a long-necked and long-limbed, three-toed
South AmericaSouth America is the southern continent of the Americas, situated entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere...
n
ungulateUngulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...
mammalMammals are a class of vertebrate animals whose females are characterized by the possession of mammary glands while both males and females are characterized by sweat glands, hair, three middle ear bones used in hearing, and a neocortex region in the brain.Mammals are divided into three main...
, typifying the order
LitopternaThe Litopterna is an order of fossil hoofed mammals from the Tertiary period that displays toe reduction. Three-toed, and even a one-toed horselike form developed....
. The oldest fossils date back to around 7 million years ago, and
M. patagonica disappears from the fossil record during the late
PleistoceneThe Pleistocene is the epoch from 2.588 million to 12 000 years BP covering the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
, around 20,000 years ago.
M. patagonica was the best known member of the family
MacraucheniidaeMacraucheniidae is a family in the Litopterna order of extinct South American ungulates. The recessed nasal bones of their skulls suggest that they may have had a small proboscis, or trunk. Their hooves were similar to those of rhinoceroses today, with a simple ankle joint and three digits on each...
, and is known only from
fossilFossils are the preserved remains or traces of animals, plants, and other organisms from the remote past. The totality of fossils, both discovered and undiscovered, and their placement in fossiliferous rock formations and sedimentary layers is known as the fossil record...
finds in
South AmericaSouth America is the southern continent of the Americas, situated entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere...
, primarily from the Lujan Formation in
ArgentinaArgentina, officially the Argentine Republic , is the second largest country in South America, constituted as a federation of 23 provinces and an autonomous city, Buenos Aires. It is the eighth largest country in the world by land area and the largest among Spanish-speaking nations, though Mexico,...
. The original specimen was discovered by
Charles DarwinCharles Robert Darwin FRS was an English naturalist who realised and presented compelling evidence that all species of life have evolved over time from common ancestors, through the process he called natural selection...
during the voyage of the
Beagle. In life,
Macrauchenia resembled a humpless camel with a short trunk, though it is not closely related to either camels or
proboscideaProboscidea is an order containing one living family, Elephantidae, and several recently extinct families, Gomphotheriidae , possibly Stegodontidae and Mammutidae...
ns.
History
Macrauchenia appeared in the fossil record some 7 million years ago in South America (in the
MioceneThe Miocene is a geological epoch of the Neogene period and extends from about 23.03 to 5.33 million years before the present . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the...
epoch). It is likely that
Macrauchenia arose from either
TheosodonTheosodon is an extinct genus of litoptern mammal from the early Miocene of South America.Theosodon bore a superficial resemblance the modern guanaco, and was around in length. It had a long neck and tapir-like, three-toed feet...
or
Promacrauchenia.
NotoungulataThe notoungulates are an extinct order of hoofed mammals that were native to South America. The order includes the huge Toxodon. Due to the isolated nature of South America, many notoungulates evolved along the lines of convergent evolution into forms that resembled mammals on other continents...
and
LitopternaThe Litopterna is an order of fossil hoofed mammals from the Tertiary period that displays toe reduction. Three-toed, and even a one-toed horselike form developed....
were two ancient orders of ungulates which only occurred in South America. Many of these species became extinct through competition with invading North American ungulates during the
Great American InterchangeThe Great American Interchange was an important paleozoogeographic event in which land and freshwater fauna migrated from North America via Central America to South America and vice versa, as the volcanic Isthmus of Panama rose up from the sea floor and bridged the formerly separated continents...
, after the establishment of the
Central AmericaManagua
Guatemala City
San Salvador
San Pedro Sula
Panama City
San José, Costa Rica
Santa Ana, El Salvador
León
San Miguel|-|}...
n land bridge. A few survivors of this invasion were the litopterns
Macrauchenia and
XenorhinotheriumXenorhinotherium , a Brazilian species in the Macraucheniidae family, was related to Macrauchenia patachonica of Patagonia.-Name:...
and the large notungulates
ToxodonToxodon is an extinct mammal of the late Pliocene and Pleistocene epochs about 2.6 million to 16,500 years ago. It was indigenous to South America, and was probably the most common large-hoofed mammal in South America at the time of its existence....
and
MixotoxodonMixotoxodon is an extinct toxodontid notoungulate, known from a single species M. larensis. Mixotoxodon is the only notoungulate known to have migrated out of South America during the Great American Interchange. Its fossils have been found in both northern South America and throughout Central...
. These last original South American hoofed animals died out eventually at the end of the Pleistocene, along with numerous other large animals on the American continent (such as American
elephantElephants are large land mammals in two genera of the family Elephantidae: Elephas and Loxodonta. Three species of elephant are living today: the African Bush Elephant, the African Forest Elephant and the Asian Elephant...
s,
horseThe horse is a hoofed mammal, a subspecies of one of seven extant species of the family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today...
s,
camelCamels are even-toed ungulates within the genus Camelus. The dromedary or Arabian camel has a single hump, and the Bactrian camel has two humps. They are native to the dry desert areas of western Asia, and central and east Asia, respectively...
s,
saber-toothed catSaber-toothed cat refers to extinct subfamilies of Machairodontinae , Barbourofelidae , and Nimravidae as well as two marsupial families that were found worldwide from the Eocene-Pleistocene epochs , existing for approximately .The Nimravidae are the oldest entering the landscape around 42 mya and...
s and
ground slothGround sloths are a diverse group of extinct sloths, in the mammalian superorder Xenarthra. Their most recent survivors lived in the Antilles, where it has been proposed they may survived until 1550 AD; however, the youngest AMS radiocarbon date reported is 4190 BP, calibrated to c. 4700 BP for...
s). As this genus was the last of the litopterns, its extinction ended that line of mammals.
Anatomy
Macrauchenia had a somewhat
camelCamels are even-toed ungulates within the genus Camelus. The dromedary or Arabian camel has a single hump, and the Bactrian camel has two humps. They are native to the dry desert areas of western Asia, and central and east Asia, respectively...
-like body, with sturdy legs, a long neck and a relatively small head. Its feet, however, more closely resembled those of a modern
rhinocerosRhinoceros , often colloquially abbreviated rhino, is a name used to group five extant species of odd-toed ungulates in the family Rhinocerotidae. Two of these species are native to Africa and three to southern Asia. Three of the five species—the Javan, Sumatran and Black Rhinoceros—are...
, and had three hoofs each. It was a relatively large animal, with a body length of around .
One striking characteristic of
Macrauchenia is that, unlike most other mammals, the openings for nostrils on its skull were atop the head, leading some early scientists to believe that, much like a
whaleWhale is the common name for marine mammals of the order Cetacea. The term whale is sometimes used to refer to all cetaceans, but in more common English usage it generally excludes the members of the Delphinoidea superfamily, such as dolphins and porpoises...
, it used these nostrils as a form of snorkel. Soon after some more recent findings, this theory was rejected. An alternative theory is that the animal possessed a trunk, perhaps to keep dust out of the nostrils.
One insight into
Macrauchenias 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. It is speculated that since Macrauchenia
lived in an environment much like the savannaA savanna, or savannah, is a grassland ecosystem characterized by the trees being sufficiently small or widely spaced so that the canopy does not close...
s of modern-day Africa, it may have had a tawny coat to match the color of dried grass.
Macrauchenia
is known, like its relative, TheosodonTheosodon is an extinct genus of litoptern mammal from the early Miocene of South America.Theosodon bore a superficial resemblance the modern guanaco, and was around in length. It had a long neck and tapir-like, three-toed feet...
, to have had a full set of 44 teeth.
Diet and behavior
Macrauchenia
was an herbivore, likely living on leaves from trees or grasses. Scientists believe that, because of the forms of its teeth, it ate using its trunk to grasp leaves and other food. It is also believed that it lived in herds like modern-day wildebeestThe wildebeest , also called the gnu , is an antelope of the genus Connochaetes. It is a hooved mammal. Wildebeest is Dutch for wild beast....
or antelopeAntelope is a term referring to many even-toed ungulate species found in the family Bovidae. The term does not refer to a monophyletic group, as not all members of Bovidae are considered antelope. Instead, the term refers to a ‘miscellaneous’ group within the family encompassing the species which...
, the better to escape predators.
Predators
When Macrauchenia
first arose, it would have been preyed upon by the largest of native South American predators, terror birds such as AndalgalornisAndalgalornis steulleti was a species of giant flightless predatory birds of the family Phorusrhacidae or "terror birds" that lived in Argentina....
, and carnivorous marsupialMarsupials are an infraclass of mammals, characterized by a distinctive pouch , in which females carry their young through early infancy.- History :...
s such as ThylacosmilusThylacosmilus was a genus of sabre-toothed metatherian predators that first appeared during the Miocene. Remains of the animal have been found in parts of South America, primarily Argentina...
and BorhyaenaBorhyaena is an extinct genus of South American marsupial, living between 20 and 15 million years ago.Borhyaena was a large, bear-like marsupial predator of up to long and with an estimated average weight of . A typical borhyaenid, it had flat feet and a heavy build...
. During the late PlioceneThe Pliocene epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present....
/Early Pleistocene, the Panama Isthmus formed, allowing predators of North AmericaNorth America is the northern continent of the Americas, situated in the Earth's northern hemisphere and in the western hemisphere. It is bordered on the north by the Arctic Ocean, on the east by the North Atlantic Ocean, on the southeast by the Caribbean Sea, and on the west by the North Pacific...
n origin, such as the puma, the jaguarThe Jaguar, Panthera onca, is a big cat, a feline in the Panthera genus. It is the only Panthera species found in the Americas. The jaguar is the third-largest feline after the tiger and the lion, and the largest and most powerful feline in the Western Hemisphere...
and the saber-toothed catSaber-toothed cat refers to extinct subfamilies of Machairodontinae , Barbourofelidae , and Nimravidae as well as two marsupial families that were found worldwide from the Eocene-Pleistocene epochs , existing for approximately .The Nimravidae are the oldest entering the landscape around 42 mya and...
, Smilodon populatorSmilodon populator was a machairodontine saber-toothed cat species. Smilodon populator first appeared in South America about 1 million years ago and became extinct about 10,000 years ago...
, to emigrate into South America and replace the native forms.
It is presumed that Macrauchenia
dealt with its predators primarily by outrunning them, or, failing that, kicking them with its long, powerful legs, much like modern-day vicuñaThe vicuña or vicugna is one of two wild South American camelids, along with the guanaco, which live in the high alpine areas of the Andes. It is a relative of the llama, and is now believed to be the wild ancestor of domesticated alpacas, which are raised for their fiber...
or camelCamels are even-toed ungulates within the genus Camelus. The dromedary or Arabian camel has a single hump, and the Bactrian camel has two humps. They are native to the dry desert areas of western Asia, and central and east Asia, respectively...
s. Its potential ability to twist and turn at high speed could have enabled it to evade pursuers.
Fossil evidence
Macrauchenia
was first discovered on 9 February 1834 at Port St JulianPuerto San Julián, also known historically as Port St Julian, is a natural harbour in Patagonia in the Santa Cruz Province of Argentina located at . In the days of sailing ships it formed a stopping point, 180 km south of Puerto Deseado...
in PatagoniaPatagonia is a geographic region containing the southernmost portion of South America. Located in Argentina and Chile, it comprises the southernmost portion of the Andes mountains to the west and south, and plateaux and low plains to the east. The name Patagonia comes from the word patagón used by...
(Argentina) by Charles DarwinCharles Robert Darwin FRS was an English naturalist who realised and presented compelling evidence that all species of life have evolved over time from common ancestors, through the process he called natural selection...
, when HMS Beagle
HMS Beagle was a Cherokee class 10-gun brig-sloop of the Royal Navy, named after the beagle, a breed of dog. She was launched on 11 May 1820 from the Woolwich Dockyard on the River Thames, at a cost of £7,803...
was surveying the port during the voyage of the Beagle
The second voyage of HMS Beagle from 27 December 1831 to 2 October 1836 was the second survey expedition of HMS Beagle, under captain Robert FitzRoy who had taken over command of the ship on its first voyage after her previous captain committed suicide...
. As a non-expert he tentatively identified the leg bones and fragments of spine he found as "some large animal, I fancy a MastodonMastodons or mastodonts were large tusked mammal species of the extinct genus Mammut found in Asia, Africa, Europe, North America and Central America from the Oligocene through Pleistocene, 33.9 mya to 11,000 years ago. The American mastodon is the most recent and best known species of the group...
". In 1837, soon after the Beagle's
return, the anatomist Richard OwenSir Richard Owen KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...
revealed that the bones including vertebrae from the back an neck were actually from a gigantic creature resembling the LlamaThe llama is a South American camelid, widely used as a pack animal by the Incas and other natives of the Andes mountains. In South America llamas are still used as beasts of burden, as well as for the production of fiber and meat....
and the camelCamels are even-toed ungulates within the genus Camelus. The dromedary or Arabian camel has a single hump, and the Bactrian camel has two humps. They are native to the dry desert areas of western Asia, and central and east Asia, respectively...
, which Owen named Macrauchenia patachonica
. In naming it, Owen noted the original Greek terms Μακρος (large or long), and αυχην (neck) as used by IlligerJohann Karl Wilhelm Illiger was a German entomologist and zoologist.Illiger was the son of a merchant in Brunswick. He studied under the entomologist Johann Hellwig, and later worked on the zoological collections of Johann Centurius Hoffmannsegg...
as the basis of Auchenia
as a generic name for the Llama, VicugnaThe vicuña or vicugna is one of two wild South American camelids, along with the guanaco, which live in the high alpine areas of the Andes. It is a relative of the llama, and is now believed to be the wild ancestor of domesticated alpacas, which are raised for their fiber...
and so on. The find was one of the discoveries leading to the inception of Darwin's theoryThe inception of Darwin's theory occurred during an intensively busy period which began when Charles Darwin returned from the survey voyage of the Beagle, with his reputation as a fossil collector and geologist already established...
. Since then, more Macrauchenia
fossils have been found, mainly in Patagonia, but also in BoliviaBolivia, officially Plurinational State of Bolivia , is a landlocked country in central South America. It is bordered by Brazil to the north and east, Paraguay and Argentina to the south, and Chile and Peru to the west....
, ChileChile, officially the Republic of Chile , is a country in South America occupying a long, narrow coastal strip between the Andes mountains to the east and the Pacific Ocean to the west. It borders Peru to the north, Bolivia to the northeast, Argentina to the east, and the Drake Passage in the far...
and VenezuelaVenezuela , officially titled Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It is a continental mainland with numerous islands located off its coastline in the Caribbean Sea...
.
Cultural references
Macrauchenia
is featured in the episode "Saber-tooth" of the documentary Walking with BeastsWalking with Beasts is a 2001 six-part television documentary produced by the BBC in the United Kingdom, narrated by Kenneth Branagh. In North America it has been retitled Walking with Prehistoric Beasts, and the original Discovery Channel broadcast was narrated by Stockard Channing...
, and individuals are featured in the 2002 Blue Sky film Ice AgeIce Age is a American computer-animated film created by Blue Sky Studios and released by 20th Century Fox in 2002. It was directed by Carlos Saldanha and Chris Wedge from a story by Michael J. Wilson...
and its sequel, the 2006 film Ice Age: The MeltdownIce Age: The Meltdown, also known as Ice Age 2: The Meltdown or simply as Ice Age 2, is the sequel to the computer-animated Ice Age. It was produced by Blue Sky Studios for 20th Century Fox, and premiered in Belgium on March 1, 2006. It was eventually released in 70 countries, with the last...
. It was included in the simulation game Zoo Tycoon: Complete Collection as part of the Dinosaur Digs Theme Pack and in Wildlife Park 2: Crazy Zoo as a cloneable beast.
External links