Eremotherium
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''Eremotherium'' (from Greek for "steppe" or "desert beast": ἔρημος "steppe or desert" and θηρίον "beast") is an extinct genus of giant
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 Caribb ...
, in the family
Megatheriidae Megatheriidae is a family of extinct ground sloths that lived from approximately 23 mya—11,000 years ago. Megatheriids appeared during the Late Oligocene (Deseadan in the SALMA classification), some 29 million years ago, in South America ...
, the largest and most heavily built family of sloths. ''Eremotherium'' lived in the southern parts of North America and the northern parts of South America from the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Late Pleistocene The Late Pleistocene is an unofficial age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a stratigraphic perspective. It is intended to be the fourth division of the Pleistocene Epoch withi ...
, around 10,000 years ago. ''Eremotherium'' was widespread in tropical and subtropical lowlands and lived there in partly open and closed landscapes, while its close relative '' Megatherium'' lived in more temperate climes. Both genera reached the size of today's elephants and were among the largest mammals in the Americas. Characteristic of ''Eremotherium'' was its robust physique with comparatively long limbs and front and hind feet especially for later representatives- three fingers. However, the skull is relatively gracile, the teeth are uniform and high-crowned. Like today's sloths, ''Eremotherium'' was a pure herbivore who dined on leaves and grasses, though it may have had semi-aquatic habits like modern hippos based on isotopic data. ''Eremotherium'' was a generalist that could adapt its diet to the respective local and climatic conditions of many regions. Finds of ''Eremotherium'' are common and widespread, with fossils being found as far north as South Carolina in the United States and as far south as Rio Grande Do Sul, and many complete skeletons have been unearthed. Only two valid species are known, ''Eremotherium laurillardi'' and ''E. eomigrans,'' the former was named by prolific Danish paleontologist Peter Lund in 1842 based on a tooth of a juvenile individual that had been collected from Pleistocene deposits in caves in Lagoa Santa, Brazil alongside fossils of thousands of other megafauna. Lund originally named it as a species of its relative ''Megatherium'', though Austrian paleontologist Franz Spillman later created the genus name ''Eremotherium'' after noticing its distinctness from other megatheriids.


History and naming

The taxonomic history of ''Eremotherium'' largely involves it being confused with '' Megatherium'' and the naming of many additional species that are actually synonymous with ''E. laurillardi.'' For many years fossils from the genus have been known, with records from as early as 1823 when fossil collectors J. P. Scriven and Joseph C. Habersham collected several teeth, skull, and mandible fragments, including a nearly complete set of mandibles, from Quaternary age deposits in Skidaway Island,
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in the United States. The fossils were not described until 1852 however, when American paleontologist named ''Megatherium mirabile'', based on the specimens (specimen numbers
USNM The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with ...
825-832 + 837) but the species has since been synonymized with ''Eremotherium laurillardi''. The first published discovery was only a year after ''M. mirabile'' was discovered, when portions of 2 teeth that had been also collected from Skidaway Island were referred to ''Megatherium'' later in 1823 by Dr. Samuel L. Mitchell.Hodgson, W. B., & Habersham, J. C. (1846). ''Memoir on the Megatherium, and Other Extinct Gigantic Quadrupeds of the Coast of Georgia: With Observations on Its Geologic Feature'' (Vol. 10). Barlett & Welford. 20 more fossils from the island were reported in 1824 by naturalist
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, including mandibular, limb, and dental remains, that now reside at the
Lyceum of Natural History The New York Academy of Sciences (originally the Lyceum of Natural History) was founded in January 1817 as the Lyceum of Natural History. It is the fourth oldest scientific society in the United States. An independent, nonprofit organization wit ...
in New York, that had also been collected by Joseph C. Habersham. Several other discoveries from Georgia and
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
were described as ''Megatherium'' throughout the 1840s and 1850s, like in 1846 when
Savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
scholar William B. Hodgson described some "''Megatherium''" fossils from Georgia that had been donated by Habersham, including portions of several skulls, in a collection that included fossils of several other Pleistocene megafauna like
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...
s and
bison Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North A ...
. These were all described in more detail by
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later was a professor of natural history at Swarthmore ...
in 1855, but they were not all referred to ''Eremotherium'' until the late 20th century. In 1842,
Richard Harlan Richard Harlan (September 19, 1796 – September 30, 1843) was an American paleontologist, anatomist, and physician. He was the first American to devote significant time and attention to vertebrate paleontology and was one of the most importan ...
named a new species of the turtle ''Chelonia, Chelonia couperi'', based on a supposed
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 ...
, or thigh bone, that had been found in the Brunswick Canal in Glynn County, Georgia and dated to the Pleistocene. It was not until 1977 that further analysis demonstrated that the "femur" was actually a clavicle from ''Eremotherium''. It is unknown, which publication was published first - according to the regulations of the ICZN, the species name of the first publication would have priority, even if it was attached to another genus - but the species name ''E. couperi'' is rarely used, while ''E. laurillardi'' is more widely used and has been adopted by more scientists. Fossils from South America were first described by Danish paleontologist and founder of Brazilian paleontology
Peter Wilhelm Lund Peter Wilhelm Lund (14 June 1801 – 25 May 1880) was a Danish paleontologist, zoologist, and archeologist. He spent most of his life working and living in Brazil. He is considered the father of Brazilian paleontology as well as archaeology. He ...
when he established a new species of ''Megatherium'' based on two teeth (specimen number ZMUC 1130 and 1131) from Lapa Vermella, a cave in the valley of the Rio de la Velhas in the Brazilian state of Minas Gerais under the name ''Megatherium laurillardi'', the first named species now assigned to ''Eremotherium''. Lund diagnosed the species based on the size of the teeth, which were only a quarter the size of ''Megatherium americanum'', the greatest representative of ''Megatherium'', and he believed that it was a tapir-sized animal.Lund, P.W., 1842. Blik paa Brasiliens Dyreverden för Sidste Jordomvaeltning. Tredie Afhandling: Forsaettelse af Pattedyrene. Det Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Math. Afd. 9, 137–208. Today, the teeth are considered to be from a juvenile of ''E. laurillardi'' and adults reached or exceeded the size of ''M. americanum''. Two years earlier, Lund had already figured teeth found at Lapa Vermella, which he assigned to ''Megatherium americanum'' due to their dimensions, which he figured alongside those of ''M. laurillardi'' in the 1842 publication.Lund, P. W. (1840). Nouvelles recherches sur la faune fossile du Brésil. In ''Annales des Sciences Naturelles'' (Vol. 13, pp. 310-319). They also have been referred to ''Eremotherium laurillardi''. For many years, ''E. laurillardi'''s holotype was speculated to actually have come from a dwarf species of ''Eremotherium'' while the larger fossils belonged to another distinct species like ''E. rusconii,'' a species that was erected by Samuel Schaub in 1935 for giant fossils from Venezuela, though it was initially thought to be a species of ''Megatherium''. However, this view is mostly contradicted and argues that at least in the Late Pleistocene in South and North America there was only a single species, ''E. laurillardi'', which had a strong
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
. Discoveries of extensive material of ''Eremotherium'' at sites such as those at Nova Friburgo in Brazil and
Daytona Beach Daytona Beach, or simply Daytona, is a coastal resort-city in east-central Florida. Located on the eastern edge of Volusia County near the Atlantic coastline, its population was 72,647 at the 2020 census. Daytona Beach is approximately nort ...
in Florida further prove that the two were synonymous and lacked any major differences between populations. Fossils of ''Eremotherium'' from Mexico were first described in 1882 by French scientist Alfred Duges, though they consisted only of a fragmentary left femur, as a new species of the South American ''
Scelidotherium ''Scelidotherium'' is an extinct genus of ground sloth of the family Scelidotheriidae, endemic to South America during the Late Pleistocene epoch. It lived from 780,000 to 11,000 years ago, existing for approximately . Description It is chara ...
'', naming it ''S. guanajatense.'' The femur had been found in Pleistocene deposits in
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, Mexico, but the fossil has since been lost and the species is a synonym of ''E. laurillardi.'' Another species that is currently considered valid was described in 1997 by Canadian zoologist Gerardo De Iuliis and French paleontologist Pierre-Antoine St-Andréc based on a single, approximately 39 cm long femur from the Pleistocene strata in Ulloma,
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as ''Eremotherium sefvei,'' though it was first described in 1915 as a fossil of ''Megatherium''. ''E. sefvei'''s geologic aging is less definite can only be placed in the general Pleistocene, but it is the smallest representative of ''Eremotherium'' and all post-Miocene megatheriids. Two years later in 1999, De Iuliis and Brazilian paleontologist Carlos Cartelle erected another species of ''Eremotherium'' now seen as valid, ''E. eomigrans'', based on a partial skeleton, the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
, that had been unearthed from the latest
Blancan The Blancan North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 4,750,000 to 1,806,000 years BP, a period of .Latest Pliocene) layers of
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,
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, USA, though many other fossils from the area were referred to it. Many of the fossils were isolated and had been recovered from sinkholes, river canals, shorelines, and hot springs, with few of the specimens being associated skeletons. So far, the latter has only been found in North America and reached a size similar to ''E. laurillardi'', but comes from the Pliocene and Early Pleistocene and bares a pentadactyl, or five fingered, hand in contrast to the tridactyl hands of ''Megatherium'' and ''E. laurillardi''. The genus name ''Eremotherium'' was not erected until 1948 by Franz Spillmann, erecting a new species, ''E. carolinese'', as the type species of the genus based on a 65 cm long skull with associated lower jaw, both fossils come from the Santa Elena Peninsula in Ecuador, and the species name was after the local village of Carolina.Spillmann, F. (1948). Beitrge zur Kenntnis eines neuen gravigraden Riesensteppentieres (Eremotherium carolinenese gen. et. spec. nov.), seines Lebensraumes und seiner Lebensweise. ''Palaeobiologica'', ''8''(3), 231-279. Although it was the type species of the genus for many years, the species has since been synonymized with ''E. laurillardi'' and has been replaced by it as the type species. The generic name ''Eremotherium'' is derived from the Greek words ἔρημος (Erēmos "Steppe", "desert") and θηρίον (Thērion "animal") after the landscape in Santa Elena Peninsula that ''E. carolinese'' was unearthed from. The following year, French taxonomist
Robert Hoffstetter Robert Julien Hoffstetter (11 June 1908 in Fargniers – 29 December 1999 in Gennevilliers) was a French taxonomist and herpetologist who was influential in categorizing reptiles Reptiles, as most commonly defined are the animals in the clas ...
introduced the genus ''Schaubia'' for Samuel Schaub's ''Megatherium rusconii'' because he recognized its generic distinctness from ''Megatherium'', though the genus name was preoccupied, so it was renamed ''Schaubtherium'' the following year. It was not until 1952 that he recognized similarities to Spillmann's ''Eremotherium'' and synonymized the two. Another dubious genus and species, ''Xenocnus cearensis'', was dubbed in 1980 by Carlos de Paula Couto based on a partial unciform (wrist bone), though he mistook as the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
(tarsal bone) of a megalochynid, that had been found in Pleistocene deposits in Itapipoca, Brazil.DE, P. (1980). FOSSIL PLEISTOCENE TO SUB-RECENT MAMMALS FROM NORTHEASTERN BRAZIL. I: EDENTATA MEGALONYCHIDAE. Paula Couto even created a new subfamily, Xenocninae, for the genus, but reanalysis in 2008 proved that the fossil was instead from ''Eremotherium laurillardi''.


Description


Size

''Eremotherium'' rivalled the closely related '' Megatherium'' in size, reaching an overall length of and a height of while on all fours, but could attain a height of about when it reared up on its hind legs. Weight estimates vary, with a range of .Mário, Dantas (2017) ''Isotopic paleoecology of the Pleistocene megamammals from the Brazilian Intertropical Region: Feeding ecology (δ13C), niche breadth and overlap'' p.152Franz Spillmann: Contributions to the knowledge of a new gravigrade giant steppe animal (Eremotherium carolinense gen. et sp. nov.), its habitat and its way of life. Palaeobiologica 8, 1948, pp. 231-279 In any case, it is one of the largest land-dwelling mammals of that time in
the Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
, along with the
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family ( Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...
ns that migrated from
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelag ...
.Richard M. Fariña, Sergio F. Vizcaíno and Gerardo de Iuliis: Megafauna. Giant beasts of Pleistocene South America. Indiana University Press, 2013, pp. 1-436 (pp. 216-218) ISBN 978-0-253-00230-3Sergio F. Vizcaíno, M. Susasna Bargo and Richard A. Fariña: Form, function, and paleobiology in xenarthrans. In: Sergio F. Vizcaíno and WJ Loughry (eds.): The Biology of the Xenarthra. University Press of Florida, 2008, pp. 86-99 As a ground-dwelling sloth, it had relatively shorter and stronger limbs compared to modern arboreal sloths and also had a longer tail.M Susana Bargo, Sergio F Vizcaíno, Fernando M Archuby and R Ernesto Blanco: Limb bone proportions, strength and digging in some Lujanian (Late Pleistocene-Early Holocene) mylodontid ground sloths (Mammalia, Xenarthra). Journal of Vertebrate Paleontology 20(3), 2000, pp. 601-610


Skull

The skull of ''Eremotherium'' was large and massive, but lighter in build compared to ''Megatherium''. A complete skull measured 65 cm in length and was up to 33 cm wide at the
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
es. In the area of the back teeth it reached 19 cm in height. The forehead line was clearly straight and not as wavy as in ''Megatherium''. The
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Ea ...
was shortened compared to the skull of ''Megatherium'', giving it an overall truncated cone appearance. Further differences to ''Megatherium'' existed at the
premaxillary bone The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
: In ''Eremotherium'' this had an overall triangular shape and was only loosely connected to the upper jaw, whereas in ''Megatherium'' the premaxillary bone had a quadrangular shape, as well as a firm connection to the upper jaw.Cástor Cartelle and Gerardo De Iuliis: Eremotherium laurillardi: The Panamerican Late Pleistocene megatheriid sloth. Journal of Vertebrate Paleontology 15(4), 1995, pp. 830–841 ( online ) The
occipital bone The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cer ...
is semicircular in posterior view and sloped backwards in lateral view. The articular surfaces as the point of attachment of the cervical spine curved far outwards and were relatively larger than in tree sloths and numerous other ground sloths. The
parietal bones The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is named ...
had a far outward curved shape, which was partly caused by the large
cranial cavity The cranial cavity, also known as intracranial space, is the space within the skull that accommodates the brain. The skull minus the mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the hu ...
with a volume of 1600 cm³. The strong zygomatic arch was closed, unlike today's sloths, but like the latter it had a massive bony outgrowth pointing downwards and backwards from the anterior base of the arch. In addition, a third outgrowth protruded diagonally upwards. The downward pointing bony process was clearly steeper than in other sloths. The
eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
was shallow and small and slightly lower than in ''Megatherium'' or modern sloths.Cástor Cartelle and Gerardo De Iuliis: Eremotherium laurillardi (Lund) (Xenarthra, Megatheriidae), the Panamerican giant ground sloth: Taxonomic aspects of the ontogeny of skull and dentition. Journal of Systematic Palaeontology 4 (2), 2006, pp. 199-209Gerardo De Iuliis and Cástor Cartelle: A new giant megatheriine ground sloth (Mammalia: Xenarthra: Megatheriidae) from the late Blancan to early Irvingtonian of Florida. Zoological Journal of the Linnean Society 127, 1999, pp. 495-515Virginia L Naples and Robert K McAfee: Reconstruction of the cranial musculature and masticatory function of the Pleistocene panamerican ground sloth Eremotherium laurillardi (Mammalia, Xenarthra, Megatheriidae). Historical Biology: An International Journal of Paleobiology 24 (2), 2012, pp. 187-206 The lower jaw was about long, both halves were connected by a strong symphysis, which extended forward in a spatulate shape and ended in a rounded shape. Typical for all representatives of the
Megatheriidae Megatheriidae is a family of extinct ground sloths that lived from approximately 23 mya—11,000 years ago. Megatheriids appeared during the Late Oligocene (Deseadan in the SALMA classification), some 29 million years ago, in South America ...
was the clearly downward curved course of the lower edge of the bone body, which resulted from the different length of the teeth. In ''Eremotherium'' this caused the lower jaw to be deep below the symphysis, 15 cm below the second tooth and 12.5 cm below the fourth. The thickness of the curvature of the lower margin of the mandible increased significantly in the course of individual development, but the ratio of the height of the mandibular body to the length of the tooth row remained largely the same. This differs markedly from ''Megatherium'', in which the height of the mandible increased not only in absolute terms, but also relatively in relation to the length of the dentition.Cástor Cartelle, Gerardo De Iuliis and François Pujos: Eremotherium laurillardi (Lund, 1842) (Xenarthra, Megatheriinae) is the only valid megatheriine sloth species in the Pleistocene of intertropical Brazil: A response to Faure et al., 2014. Comptes Rendus Palevol 14, 2014, pp. 15-23 The mandibular body was also very thick, leaving little space for the tongue. The crown process rose up to , and the articular process was only slightly lower. At the posterior, lower end there was a strong, clearly notched angular process, the upper edge of which was approximately at the level of the masticatory plane. At the anterior edge of the lower jaw there was a strong
mental foramen The mental foramen is one of two foramina (openings) located on the anterior surface of the mandible. It is part of the mandibular canal. It transmits the terminal branches of the inferior alveolar nerve and the mental vessels. Structure The ...
. The dentition was typical for sloths, but in contrast to today's representatives it consisted of completely
homodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
teeth, which is a characteristic feature of megatherians. Each branch of the jaw had 5 teeth in the upper jaw and 4 in the lower jaw, so in total ''Eremotherium'' had 18 teeth. They resembled molars and, except for the front one, were quadrangular in shape, usually a good long in large individuals and very high-crowned (''hypsodont'') with a height of . They had no roots and grew throughout their entire life. The enamel was also missing. However, two transverse, sharp-edged ridges were typically formed on the chewing surface to help grind food. The entire upper row of teeth grew up to long, while the lower reached up to .Martine Faure, Claude Guérin and Fabio Parenti: Sur l'existence de deux specèces d'Eremotherium E. rusconii (Schaub, 1935) et E. laurillardi (Lund, 1842) dans le Pléistocène supérieur du Brésil intertropical. Comptes Rendus Palevol 13 (4), 2014, pp. 259-266M. Susana Bargo, Gerardo de Iuliis and Sergio F. Vízcaino: Hypsodonty in Pleistocene ground sloths. Acta Palaeontologica Polonica 51 (1), 2006, pp. 53-61


Postcrania

Almost all of the poscranial skeleton is known. The vertebrae were massively shaped, both at the vertebral bodies and at the lateral
transverse processes The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
. However, the vertebral bodies were compressed in length, so that the tail appeared rather short overall and generally did not exceed the length of the lower limb sections. 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 r ...
represented a long tube with a bulky lower joint end. The total length was about . Distinctive, ridge-like muscle attachments on the middle shaft were typical. The forearm bones had much shorter lengths, with the
spoke A spoke is one of some number of rods radiating from the center of a wheel (the hub where the axle connects), connecting the hub with the round traction surface. The term originally referred to portions of a log that had been riven (split ...
measuring about 67 cm, and 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 ...
in length.Gerardo De Iuliis: Toward the morphofunctional understanding of the humerus of Megatheriinae: The identity and homology of some diaphyseal humeral features (Mammalia, Xenarthra, Megatheriidae). Senckenbergiana biologica 83, 2003, pp. 69-78 Massive was 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 ...
, which had the broad build characteristic of megatherians and was narrowed in front and behind. It had an average length of 74 cm, the largest bone found so far was long and wide. The third trochanter, a prominent muscle attachment point on the shaft, typical of xenarthrans, was absent in ''Eremotherium'' as in all other megatherians. The
shinbone The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
and
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
were only fused together at the upper end and not also at the lower end as in ''Megatherium''. In this case, the tibia became about 60 cm long.H. Gregory McDonald: Xenarthran skeletal anatomy: primitive or derived? Senckenbergiana biologica 83, 2003, pp. 5-17 The forelegs ended in hands with three fingers (III to V). The two inner phalanges (I and II) were fused together with some elements of the
carpus In human anatomy, the wrist is variously defined as (1) the carpus or carpal bones, the complex of eight bones forming the proximal skeletal segment of the hand; "The wrist contains eight bones, roughly aligned in two rows, known as the carpal ...
, such as the great polygonal leg, to form a unit, the metacarpal-carpal complex (MCC).Gerardo De Iuliis and Cástor Cartelle: The medial carpal and metacarpal elements of Eremotherium and Megatherium (Xenarthra: Mammalia). Journal of Vertebrate Paleontology 14, 1994, pp. 525-533 Thus, ''Eremotherium'' clearly deviates from ''Megatherium'' and other closely related forms, which possessed four-fingered hands. In ''Eremotherium'', the
metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ar ...
of the third digit was the shortest, measuring 19 cm in length, while those of the fourth and fifth were almost the same length, and respectively. The
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly ...
(the third phalanx) of the third and fourth fingers had a long and pointedly curved shape, which suggests correspondingly long claws. The fifth finger had only two phalanges and consequently no claw was formed there. (An exception is the older form ''E. eomigrans'', whose hands, in contrast to other megateria, were still five-fingered, with claws on digits I to IV.) The foot, as in all megatheriids, was also three-fingered (digits III to V). It resembled the hand with an extremely short
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
of the third finger. That of the fourth finger reached , that of the fifth in length. Deviating from the hand, only the middle digit (III) had three phalanges with a terminal phalanx bearing a long claw. The two outer digit had only two phalanges. This structure of the foot is typical for evolved megatherians.Diego Brandoni, Alfredo A. Carlini, Francois Pujos, and Gustavo J. Scillato-Yané: The pes of Pyramiodontherium bergi (Moreno & Mercerat, 1891) (Mammalia, Xenarthra, Phyllophaga): The most complete pes of a Tertiary Megatheriinae. Geodiversitas 26 (4), 2004, pp. 643–659François Pujos and Rodolfo Salas: A systematic reassessment and paleogeographic review of fossil Xenarthra from Peru. Bulletin de l'Institut Français d'Études Andines 33 (2), 2004, pp. 331-377


Palaeobiology


Locomotion

The predominantly quadrupedal locomotion took place on inwardly turned feet, with the entire weight resting on the outer, fifth and possibly fourth phalanges (a pedolateral gait), whereby the talus was subject to massive reshaping.H. Gregory McDonald: Evolution of the Pedolateral Foot in Ground Sloths: Patterns of Change in the Astragalus. Journal of Mammal Evolution 19, 2012, pp. 209-215Néstor Toledo, Gerardo De Iuliis, Sergio F. Vizcaíno and M. Susana Bargo: The Concept of a Pedolateral Pes Revisited: The Giant Sloths Megatherium and Eremotherium (Xenarthra, Folivora, Megatheriinae) as a Case Study. Journal of Mammalian Evolution 25 (4), 2018, pp. 525-537, doi:10.1007/s10914-017-9410-0 Likewise, the hands were turned inwards, in a position somewhat resembling the forefeet of the similarly clawed
Chalicotheriidae Chalicotheres (from Greek '' chalix'', "gravel" and '' therion'', "beast") are an extinct clade of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene until the Early Ple ...
, a now extinct group of
odd-toed ungulates Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
.Giuseppe Tito and Gerardo De Iuliis: Morphofunctional aspects and paleobiology of the manus in the giant ground sloth Eremotherium Spillmann 1948 (Mammalia, Xenarthra, Megatheriidae). Senckenbergiana biologica 83 (1), 2003, pp. 79-94 It also suggests that locomotion was rather slow. It was also unable to perform digging activities, as has been demonstrated for other large ground sloths, which can also be seen in the construction of the forearm, just as the manipulation of objects was minimised due to the limited ability of the fingers to move in relation to each other. However, ''Eremotherium'' was able to stand up on its hind legs and pull branches and twigs with its hands, for example to reach the foliage of tall trees for feeding,Giuseppe Tito: New remains of Eremotherium laurillardi (Lund, 1842) (Megatheriidae, Xenarthra) from the coastal region of Ecuador. Journal of South American Earth Sciences 26, 2008, pp. 424-434 as well as defensive strikes with its long claws were possible. The standing up was supported by the strong tail, similar to what is still the case today with
armadillos 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, along ...
and
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 ...
. The massive tail vertebrae in the front area of the tail suggest a strong musculature. Among other things, this concerns the
coccygeus muscle The coccygeus muscle or ischiococcygeus is a muscle of the pelvic floor, located posterior to levator ani and anterior to the sacrospinous ligament. Structure The coccygeus muscle is posterior to levator ani and anterior to the sacrospino ...
, which attaches to the
ischium The ischium () form ...
and fixes the tail. Less well developed, on the other hand, were the epaxial muscles, which could cause the tail to straighten up.


Social behaviour

Due to some group finds of several individuals at individual sites, such as in El Bajión in
Chiapas Chiapas (; Tzotzil and Tzeltal: ''Chyapas'' ), officially the Free and Sovereign State of Chiapas ( es, Estado Libre y Soberano de Chiapas), is one of the states that make up the 32 federal entities of Mexico. It comprises 124 municipalities ...
in
Mexico Mexico (Spanish language, Spanish: México), officially the United Mexican States, is a List of sovereign states, country in the southern portion of North America. It is borders of Mexico, bordered to the north by the United States; to the so ...
with four animals or in Tanque Loma on the Santa Elena in
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ' ...
with 22 individuals, some scientists discuss whether ''Eremotherium'' possibly lived and roamed in small, herd-like groups.Bruno Andrés Than-Marchese, Luis Enrique Gomez-Perez, Jesús Albert Diaz-Cruz, Gerardo Carbot-Chanona and Marco Antonio Coutiño-José: Una nueva localidad con restos de Eremotherium laurillardi (Xenarthra: Megateriidae) in Chiapas, Mexico: possible evidence de gregarismo en la especie. VI Jornadas Paleontológicas y I Simposio de Paleontología en el Sureste de México: 100 years de paleontología en Chiapas, 2012, p. 50Emily L Lindsey, Erick X Lopez Reyes, Gordon E Matzke, Karin A Rice, and H Gregory McDonald: A monodominant late-Pleistocene megafauna locality from Santa Elena, Ecuador: Insight on the biology and behavior of giant ground sloths. Palaeogeography, Palaeoclimatology, Palaeoecology, 2020, p. 109599, Especially in Tanque Loma, the individuals recorded are composed of at least 15 adults and six juveniles. They were all found in close association in a single horizon, and they are interpreted as being contemporary with each other. The possible group was thought to have gathered at a waterhole and died there relatively abruptly due to an unknown event. On the other hand, sometimes clustered occurrences of ''Eremotherium'' such as the 19 individuals from the
sinkhole A sinkhole is a depression or hole in the ground caused by some form of collapse of the surface layer. The term is sometimes used to refer to doline, enclosed depressions that are locally also known as ''vrtače'' and shakeholes, and to openi ...
of Jirau in
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 ...
are considered to be accumulations over a long period of time.Hermínio Ismael de Araújo-Júnior, Kleberson de Oliveira Porpino, Celso Lira Ximenes and Lílian Paglarelli Bergqvist: Unveiling the taphonomy of elusive natural tank deposits: A study case in the Pleistocene of northeastern Brazil. Palaeogeography, Palaeoclimatology, Palaeoecology 378, 2013, pp. 52-74 In the case of the likewise giant ground sloth '' Lestodon'' from central South America, experts also interpret mass accumulations of remains of different individuals in part as evidence of phased group formation.Rodrigo L. Tomassini, Claudia I. Montalvo, Mariana C. Garrone, Laura Domingo, Jorge Ferigolo, Laura E. Cruz, Dánae Sanz-Pérez, Yolanda Fernández-Jalvo, and Ignacio A. Cerda: Gregariousness in the giant sloth Lestodon (Xenarthra ): multi‑proxy approach of a bonebed from the Last Maximum Glacial of Argentine pampas. Scientific Reports 10, 2020, p. 10955, doi:10.1038/s41598-020-67863-0 Living tree sloths live solitary lives.Adriano Garcia Chiarello: Sloth ecology. An overview of field studies. In: Sergio F. Vizcaíno and WJ Loughry (eds.): The Biology of the Xenarthra. University Press of Florida, 2008, pp. 269-280


Diet

''Eremotherium'' possessed extremely high-crowned teeth, which, however, did not reach the dimensions of those of ''Megatherium''. As the teeth lack enamel, this hypsodonty may not be an expression of specialisation on grass as food, unlike mammals with enamel in their teeth. The different expression of high-crownedness in the two large ground sloths is probably rather to be sought in adaptation to divergent habitats—more tropical lowlands in ''Eremotherium'' and more temperate regions in '' Megatherium''. From an
anatomical Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
point of view, the only moderately wide snout and the large total chewing surface of the teeth advocate a diet adapted to mixed plant foods. The average surface area of all teeth available for chewing food is 11,340 mm², which roughly corresponds to the values of the closely related ''Megatherium'', but clearly exceeds those of the '' Lestodon'', which is also giant but has a much broader snout. The latter genus belongs to the more distantly related Mylodontidae and was probably a specialised grazer. Moreover, the total purchase area is within the range of variation of present-day
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantida ...
s, some of which also prefer mixed plant diets.Sergio F Vizcaíno, M Susana Bargo and Guillermo H Cassini: Dental occlusal surface area in relation to body mass, food habits and other biological features in fossil xenarthrans. Ameghiniana 43 (1), 2006, pp. 11-26Mário AT Dantas and Adaiana MA Santos: Inferring the paleoecology of the Late Pleistocene giant ground sloths from the Brazilian Intertropical Region. Journal of South American Earth Sciences 117, 2022, p.103899, doi:10.1016/j.jsames.2022.103899 Support for this view comes from various
isotopic analysis Isotope analysis is the identification of isotopic signature, abundance of certain stable isotopes of chemical elements within organic and inorganic compounds. Isotopic analysis can be used to understand the flow of energy through a food we ...
on the teeth of ''Eremotherium''. Thus, the animals probably fed on grass in rather open landscapes, but on foliage in largely closed forests.Mário André Trindade Dantas, Rodrigo Parisi Dutra, Alexander Cherkinsky, Daniel Costa Fortier, Luciana Hiromi Yoshino Kamino, Mario Alberto Cozzuol, Adauto de Souza Ribeiro and Fabiana Silva Vieira: Paleoecology and radiocarbon dating of the Pleistocene megafauna of the Brazilian Intertropical Region. Quaternary Research 79, 2013, pp. 61-65 Carbon isotopes and stereo microwear analysis suggest that an individual from the Late Pleistocene (34,705-33,947 cal yr BP), of
Goiás Goiás () is a Brazilian state located in the Center-West region. Goiás borders the Federal District and the states of (from north clockwise) Tocantins, Bahia, Minas Gerais, Mato Grosso do Sul and Mato Grosso. The state capital is Goiâ ...
, Brazil, was a mixed feeder, suggesting a high proportion of shrubs and trees, this is in contrast to the presumed diet from specimens from
Northeast Brazil The Northeast Region of Brazil ( pt, Região Nordeste do Brasil; ) is one of the five official and political regions of the country according to the Brazilian Institute of Geography and Statistics. Of Brazil's twenty-six states, it comprises ni ...
, which had a diet of C4
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
plants. A 2020 discovery in Ecuador found 22 individuals ranging in age from juveniles to adults preserved together in anoxic marsh sediments, suggesting that ''Eremotherium'' may have been gregarious.


Relationship with humans and extinction

The disappearance of ''Eremotherium'' coincides with the Quaternary Extinction Event, which saw the arrival of humans in the Americas and the extinction of many megafauna, large or giant animals of an area, habitat, or geological period, extinct and/or extant that were larger than or a comparable size to humans, such as
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...
s,
glyptodont Glyptodonts are an extinct subfamily of large, heavily armoured armadillos. They arose in South America around 48 million years ago and spread to southern North America after the continents became connected several million years ago. The best-k ...
s, and other ground sloths. One of the latest finds of ''Eremotherium'' is from Ittaituba on Rio Tapajós, a tributary of the Amazon, that dates to 11,340 BP and includes several skull and lower jaw fragments. In a similar period, the finds at Barcelona in the Brazilian state of Rio Grande do Norte come from strata dating from 11,324 to 11,807 years ago. The amount of hunting by Paleoindians enacted on ''Eremotherium'' is a subject of incomplete, partly controversial, scientific discourse, but joint finds of ''Eremotherium'' and early human inhabitants of the region are very rare. A possible indication human altercation is a tooth of ''Eremotherium'' that may had been edited by Paleoindians was unearthed from a doline on the site of the São-José farm in the Brazilian state of Sergipe.Hubbe, A., Haddad-Martim, P. M., Hubbe, M., & Neves, W. A. (2012)
Comments on:“An anthropogenic modification in an Eremotherium tooth from northeastern Brazil”.
''Quaternary International'', ''269'', 94-96.
However, Caninde Paleoindian artifacts date to 8950 and 5570 BP, which has led to two hypotheses: (1) ''Eremotherium'' lived until the early Holocene and interacted with humans, as evidenced by a potential discovery in Piaui, northeastern Brazil; or; (2) Humans migrated into the Americas earlier than typically regarded, about 15,000 BP. The anthropogenic origin of the brands is also discussed.


Classification

''Eremotherium'' is a genus of the extinct ground sloth family Megatheriidae, which includes large to very large sloths in the group Folivora, which, together with the Megalonychidae and the Nothrotheriidae, form the superfamily Megatherioidea. The Megatherioidea also includes the three-toed sloths of the genus '' Bradypus'', one of the two sloth genera still alive today. ''Eremotherium'''s closest relative in Megatheriidae is the namesake of the family ''Megatherium'', which was endemic to South America, slightly larger, and preferred more open habitats than ''Eremotherium''. ''Pyramiodontherium'' and ''Anisodontherium'' are also part of this subfamily, but are smaller and older, dating to the Late Miocene of Argentina. All of these genera belong to the subfamily Megatheriinae, which includes the largest and most derived sloths. The direct phylogenetic ancestor of ''Eremotherium'' is unknown, but may be linked to '' Proeremotherium'' from the Codore Formation in Venezuela, which dates to the Pliocene. The genus has numerous characteristics that are akin to those of ''Eremotherium'', but are more primitive. Little is known about the evolution of the genus ''Eremotherium''. It may have evolved in the Early Pliocene in South America, where only a few sites from this period are known, and dispersed by crossing the Isthmus of Panama, i.e. the formation of the land bridge connecting North and South America, in the course of the Great American Biotic Interchange. The oldest fossils come from the Pliocene of the southern United States in North America, suggesting that the species instead evolved there before colonizing South America. The discovery of ''Proeremotherium'' also supports this hypothesis, indicating that these or other close ancestors of ''Eremotherium'' first migrated to North America and evolved there, then moved back southward to South America after the formation of the Isthmus of Panama, similar to the glyptodont ''
Glyptotherium ''Glyptotherium'' (from Greek for 'grooved or carved beast') is a genus of glyptodont (an extinct group of large, herbivorous armadillos) that lived from the Early Pliocene, about 4.9 million years ago, to the Early Holocene, around 7,000 yea ...
''. The following phylogenetic analysis of Megatheriinae within Megatheriidae was conducted by Brandoni ''et al.,'' 2018 that was modified from Varela ''et al.'' 2019 based on lower molariform and astragalus morphology:


Fossil distribution

Finds of ''Eremotherium'' are widespread, but are limited to the tropical and subtropical lowland regions of the Americas. Fossils have come from more than 130 sites. In South America, fossils have been unearthed from as far west as northern Peru via Ecuador, Colombia to the east in Guayana and the Amazon basin. The southernmost discovery was in the Brazilian state of Rio Grande Do Sul, while further south in the Pampas and in mountainous states, the close relative ''Megatherium'' lived. In North America, fossils are known from the southern United States in Georgia to Texas to South Carolina and on the east coast from Florida to New Jersey. The oldest finds known so far come from the US state of Florida and belong to the more uncommon species ''E. eomigrans,'' which were found in Haile in a water-filled doline (locality 7c), 6 km northeast of Newberry in Alachua County. They comprise several partial skeletons and several isolated bones and are between 2.1 and 1.8 million years old, which corresponds to the beginning of the Lower Pleistocene. Additional finds from Florida such as from the Kissimmee River, which are too unspecific in order to be able to precisely date, the maximum age is 2.5 million years and thus from the transition from the Pliocene to the Pleistocene.


References

{{Taxonbar, from=Q907433 Prehistoric sloths Prehistoric placental genera Pliocene xenarthrans Pleistocene xenarthrans Zanclean first appearances Holocene extinctions Pliocene mammals of North America Pleistocene mammals of North America Rancholabrean Irvingtonian Blancan Neogene Costa Rica Neogene United States Pleistocene Costa Rica Pleistocene El Salvador Pleistocene Guatemala Pleistocene Honduras Pleistocene Mexico Pleistocene Nicaragua Pleistocene Panama Pleistocene United States Fossils of Costa Rica Fossils of El Salvador Fossils of Guatemala Fossils of Honduras Fossils of Mexico Fossils of Nicaragua Fossils of Panama Fossils of the United States Pliocene mammals of South America Pleistocene mammals of South America Lujanian Ensenadan Uquian Chapadmalalan Montehermosan Neogene Venezuela Pleistocene Bolivia Pleistocene Brazil Pleistocene Colombia Pleistocene Ecuador Pleistocene Venezuela Fossils of Bolivia Fossils of Brazil Fossils of Colombia Fossils of Ecuador Fossils of Venezuela Fossil taxa described in 1948 Taxa named by Peter Wilhelm Lund