Lestodon
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''Lestodon'' is an extinct genus of megafaunal
ground sloth Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. The term is used to refer to all extinct sloths because of the large size of the earliest forms discovered, compared to existing tree sloths. The Caribbe ...
from
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
during the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
periods. Its fossil remains have been found 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, th ...
,
Paraguay Paraguay (; ), officially the Republic of Paraguay ( es, República del Paraguay, links=no; gn, Tavakuairetã Paraguái, links=si), is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to th ...
,
Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, República Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...
,
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
,
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
, and
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. Measuring approximately from snout to tail tip, it is estimated to have weighed . It was a herbivore and primarily fed on the grasses on the South American plains and is thought to perhaps have used its semi-bipedal stance to obtain foliage from trees. ''Lestodon'' is placed as member of the
Mylodontidae Mylodontidae is a family of extinct South American and North American ground sloths within the suborder Folivora of order Pilosa The order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes the anteaters a ...
as indicated by the lobed form of the last tooth in the dentition.


Etymology

The genus name ''Lestodon'' derives from the Greek for "robber tooth".


Evolutionary adaptations


Cranial

The
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
morphologically reflects its evolutionary history as well as its daily activities. At the caudal end, the sloth’s head connects to its neck with a class one lever. 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, the ...
is two to three orders of magnitude larger than that of living species of sloth, but the muzzle itself shows highly positive
allometry Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Julian Huxley in 1932. Overview Allom ...
to the size of the skull in comparison to modern sloths and even other megafauna of the same order
Pilosa The order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes the anteaters and sloths (which includes the extinct ground sloths). The name comes from the Latin word for "hairy". Origins and taxonomy The b ...
. ''L. armatus'' had a large, blunt, and square
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
well adapted for foliage consumption. Its muzzle and
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
show a significant adaptation to this diet. The shape of the muzzle aided the ground sloth in the grazing necessary to sustain the metabolic activity of its large body; square, flat muzzles are associated with bulk feeders while pointed snouts are adaptive features of precision eaters, like the long sloping snout of modern
anteaters Anteater is a common name for the four extant mammal species of the suborder Vermilingua (meaning "worm tongue") commonly known for eating ants and termites. The individual species have other names in English and other languages. Together with ...
, which are also in the same class
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 ...
(Bargo ''et al.'', 2006). The mandible, a class three lever, has a shape that indicates evolutionary pressure for strength over speed. To sustain its caloric needs with only plant matter, the sloth would have needed to be able to consume a high quantity of food with little precision. The species would dig for its food, and would have accidentally ingested a large amount of gritty dirt and soil particulate. This gives them a large crown height, called
hypsodonty Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritt ...
, that is unsuitable for strong bite forces. While the mandibular muscles would have been well developed to support the allometrically positive
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
of ''L. armatus'', they would have also been extremely weak (Bargo ''et al.'', 2006).


Appendicular skeleton

The pursuit of bulk plant matter is reflected in the stance, movement, and total bone structure of the ground sloth. In order to dislodge a large amount of foliage from the earth, ''L. armatus'' would have used its flat, spade-like
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
in a hook-and-pull fashion to turn over soil and vegetation. Once again, the evolutionary design of the giant sloth favors quantity and force to quality and speed in its manus. While the giant sloth was not fully bipedal, its forelimbs show evolutionary pressure for specialized functions over walking and could easily shift its center of gravity. The
olecranon The olecranon (, ), is a large, thick, curved bony eminence of the ulna, a long bone in the forearm that projects behind the elbow. It forms the most pointed portion of the elbow and is opposite to the cubital fossa or elbow pit. The olecranon ...
of ''L. armatus'', for example, is longer allometrically than species whose forelimbs are primarily adapted for movement; the longer olecranon on the
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
maximizes the leverage to the moment arm of the
triceps The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of 3 parts: the medial, lateral, and long head. It is the muscle principally responsibl ...
, the extensor of the forearm, but is proportionally much longer than is necessary for walking (Bargo ''et al.'', 2000). The extension of the forearm would have been very strong, but not very fast. Triceps are the primary muscle activated during the digging motion, but the odd articulation of the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
to the
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
, and therefore the orientation of the
teres major The teres major muscle is a muscle of the upper limb. It attaches to the scapula and the humerus and is one of the seven scapulohumeral muscles. It is a thick but somewhat flattened muscle. The teres major muscle (from Latin ''teres'', meaning ...
, also strengthens the force during pulling. Compared to humans, the scapula is tilted laterally, with the medial margin at a higher elevation than the acrimonion. This orientation maximizes the power of the teres major during digging. Energy storage is possible in the
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
s of the triceps, produces power necessary to execute the digging behavior Lestodon armatus uses to acquire the bulk of its diet. The spinal column of ''L. armatus'' is arranged parallel to the ground, making the angle between the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
s and the majority of the mass of the megafauna slightly more than orthogonal. On the femur, the femoral
condyle A condyle (;Entry "condyle"
in
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 ...
, like anteaters, show a similar design. All species within this order have specialized forelimbs and a center of gravity shifted caudally towards the
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
from other quadrupeds, but rostrally when compared to fully bipedal humans. The
lumbar vertebrae The lumbar vertebrae are, in human anatomy, the five vertebrae between the rib cage and the pelvis. They are the largest segments of the vertebral column and are characterized by the absence of the foramen transversarium within the transverse p ...
are said to be xenarthrous, the eponymous characteristic of the magnorder, and that means that they come into contact with each other at more points to stabilize the pelvis and the bulk of the caudally shifted mass. It would have been unable to gallop. In humans, pelvic bones are not fused together in a solid block, and this allows us greater variability of motion from side to side. Though there is reduced mobility, xenarthrous fusion provides a stable platform for digging.


Distribution

Fossils of ''Lestodon'' have been found in:''Lestodon''
at
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.org
* Luján, Pampean and
Monte Hermoso Formation Monte may refer to: Places Argentina * Argentine Monte, an ecoregion * Monte Desert * Monte Partido, a ''partido'' in Buenos Aires Province Italy * Monte Bregagno * Monte Cassino * Montecorvino (disambiguation) * Montefalcione Portugal * Monte ...
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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, th ...
*
Tarija Formation Tarija or San Bernardo de la Frontera de Tarixa is a city in southern Bolivia. Founded in 1574, Tarija is the largest city and capital and municipality within the Tarija Department, with an airport (Capitán Oriel Lea Plaza Airport, (TJA)) offe ...
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Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
*
Rio Grande do Sul Rio Grande do Sul (, , ; "Great River of the South") is a Federative units of Brazil, state in the South Region, Brazil, southern region of Brazil. It is the Federative_units_of_Brazil#List, fifth-most-populous state and the List of Brazilian st ...
and
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Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
*
Paraguay Paraguay (; ), officially the Republic of Paraguay ( es, República del Paraguay, links=no; gn, Tavakuairetã Paraguái, links=si), is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to th ...
*
Dolores Dolores, Spanish for "pain; grief", most commonly refers to: * Our Lady of Sorrows or La Virgen María de los Dolores * Dolores (given name) Dolores may also refer to: Film * ''Dolores'' (2017 film), an American documentary by Peter Bratt * ' ...
and
Sopas Formation ''Sopas'' is a Filipino macaroni soup made with elbow macaroni, various vegetables, and meat (usually chicken), in a creamy broth with evaporated milk. It is regarded as a comfort food in the Philippines and is typically eaten during breakfast, ...
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Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, República Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...


Evidence of hunting

Researchers working at the Arroyo del Vizcaíno site near
Sauce, Uruguay Sauce is a small city and municipality in the Canelones Department of southern Uruguay. Geography Location The city is located on Km. 36 of Route 6 and its intersection with Routes 67, 86 and 107. It is part of the wider metropolitan area of ...
suggested that ''Lestodon'' was hunted by humans about 30,000 years ago. This was based on analysis of ''Lestodon'' bones. Deep slash markings on some of them were suggested to be from the use of human stone tools. However, the date is significantly older than the oldest accepted dates for humans in South America around 15,000 years ago, and there is no evidence for stone tools at the site, making a natural origin for the marks more likely.


References

{{Taxonbar, from=Q604501 Prehistoric sloths Prehistoric placental genera Pliocene xenarthrans Pleistocene xenarthrans Pliocene mammals of South America Pleistocene mammals of South America Lujanian Ensenadan Montehermosan Neogene Argentina Neogene Venezuela Fossils of Venezuela Pleistocene Argentina Fossils of Argentina Pleistocene Bolivia Fossils of Bolivia Pleistocene Brazil Fossils of Brazil Pleistocene Paraguay Fossils of Paraguay Pleistocene Uruguay Fossils of Uruguay Fossil taxa described in 1855