Discovery
''Mylodon'' was named byDescription
General
''Mylodon'' was a large representative of the Mylodontidae. Its total length was estimated to be around 3 to 4 m. Based on the size of the skull, a weight between 1 and 2 tonnes is assumed, with an approximate estimate of 1.65 tonnes. Thus, ''Mylodon'' had about the size of related forms such as '' Glossotherium'' or '' Paramylodon'', but was significantly smaller than the giant '' Lestodon''. In terms of physique, it largely corresponded with the other large ground-living sloths.Skull and dentition features
Especially in the construction of the skull, ''Mylodon'' differed significantly from other related forms. Its length varied between 59 and 71.5 cm, which is significantly longer than ''Glossotherium'' or ''Lestodon''. At the skull it was between 16.5 and 22.5 cm wide, in the front nasal area between 11.3 and 15.5 cm. The height of the posterior skull was 14.0 to 19.0 cm and the anterior 15.0 to 23.5 cm. The skull was thereby elongated and narrow, unlike ''Glossotherium'' and ''Lestodon'' that had a short and very broad skull. The extraordinary length of the skull of ''Mylodon'' was mainly due to elongations in the rostrum. Seen from above, the rostrum narrowed towards the front. This is where the most important difference to most of the other representatives of the Mylodontidae can be found: The nasal bone was long and narrow and curved downwards in the front area. At the front end, it connected to the middle jawbone, which was lengthened by an appendage, and which in turn fused with the upper jaw. This resulted in a completely closed nasal arch in adult individuals, which is largely unknown in other sloths. In comparison, the skulls of ''Glossotherium'' and ''Lestodon'', but also of ''Paramylodon'', showed a nasal area, seen from above, which was rather short and looked clearly cut off when viewed from the side; the roof of the skull was largely straight in ''Mylodon'', only a slight indentation could occur above the orbit. On parietal, significant temporal lines were present, but no head crest formed. The zygomatic arch was slim, the anterior attachment began above the third and fourth molars. It did not form a solid end with the rear arch attachment. As is usual with sloths, the front arch base consisted of three appendages: one ascending, one horizontal, and one descending, the former of which was the longest. The rear arch formed a triangular plate. The occiput bent at an angle of 120° from the roof of the skull. The underside of the occiput was at about the level of the occlusal plane. When viewed from behind, the occiput appeared almost circular and not as depressed as in ''Glossotherium'' and ''Lestodon''. The palate was narrow and was more or less triangularly oriented towards the front of the skull. Numerous small bone openings were characteristic here. The glenoid pit, in which the joint of the lower jaw engages, corresponding to that of other mylodonts with its weak form, but this provided free rotation overall. The lower jaw of ''Mylodon'' varied in length between 42 and 48 cm. It was elongated, more noticeable than in ''Glossotherium'' and ''Lestodon'', since in ''Mylodon'' the area in front of the teeth, in particular, is strongly elongated. The horizontal bone body increased continuously in height towards the rear, below the last molar it was about 10.5 to 12.7 cm. The symphysis at the front end for the jointing of the two halves of the lower jaw was about 12.4 cm long. Here the lower edge of the body of the lower jaw rose at an angle so that the anterior end of the symphysis was above the occlusal plane of the teeth. As with other sloths, the symphysis extended forward, it ended slightly rounded. According to the rostrum of the skull, ''Mylodons symphysis was narrow and not as wide as in ''Glossotherium'' and ''Lestodon''. The mandible foramen opened shortly behind the symphysis. The ascending branch started behind the last molar and formed an angle of 140° to the occlusal plane. The crown process rose up to 20 cm. In contrast, the articular process was lower, roughly at the level of the occlusal plane, resulting in a low cranial-mandibular connection. The angular process at the rear end of the lower jaw was clearly visible. Sometimes it tipped down and was below the lower edge of the horizontal bone body. The upper side of the angular process does not reach the occlusal plane. The dentition of ''Mylodon'' differs greatly from that of the other placental mammals and usually consists of five teeth at the top and four teeth at the bottom per jaw arch, meaning a total of 18 teeth. In the mylodonts, the first tooth was oftenPostcrania
Postcranial skeletons are far rarer in ''Mylodon'' than in the other large mylodontid sloths. As a result, the skeleton is less well documented. Only individual elements of the spine, such as the atlas and various thoracic vertebrae, have been described. The humerus was massive and extremely long at 46 to 48 cm. The joint head, the diameter of which was over 10 cm, stood out due to its hemispherical, but laterally somewhat flattened shape. A distinct deltopectoral ridge ran down the shaft, which acted as an anchor point for the shoulder muscles. As with many ground sloths, the lower end of the joint extended far and brought it here to a width of almost 26 cm. In part, this was caused by a massive internal epicondyle. The articular surfaces (capitulum and trochlea) were almost perpendicular to each other and did not form such an obtuse angle as in ''Glossotherium''. The cubit was built gracefully. Their length was around 37 cm. The olecranon, i.e. the upper articular process, took up about 8.1 cm of it, which corresponds to about 22% of the total length and is significantly less than in comparison with ''Glossotherium'' and ''Lestodon''. It was laterally narrowed, which is also found in ''Paramylodon''. The spoke largely resembled that of Glossotherium and was compact and straight built with a length of about 30 cm. The head was oval in shape with a prominent lip. The pelvis was extremely expansive and 114 cm wide between the two iliac bones. The thigh bone measured between 55 and 59 cm in length. It was typical of ground sloths, being flat in shape. Its width decreased significantly on the shaft, the lowest value was reached just below the midpoint. Here the width was about 18 cm, the thickness about 7.5 cm. The joint ends, on the other hand, were markedly wider, around 30 cm at the knee end and around 26 cm at the foot end. The thighbone reached the shin with only about half of its length, a characteristic of mylodonts. This bone, too, was clearly flat with a thickness that was only half the value of the width at the shaft. The fibula is so far only fragmented. It was drawn in on the shaft and widened at the joint ends, with the upper joint end showing more pronounced curves than in ''Glossotherium''.Lucas Kraglievich: Contribución al conocimiento de ''Mylodon darwini'' Owen y especies afines. Revista del Museo de La Plata 34, 1934, pp. 255-292 The hand comprised a total of five digits (I to V), whereby the metacarpal bone was fused with the large polygonal bone on the first digit. This created the so-called Metacrapal Carpal Complex (MCC for short), which is typical for many ground sloths. As a special feature of the wrist, the pea bone was clearly flat, its shape resembled that of ''Glossotherium'', but differed from the corresponding bone of other Mylodonts with spherical, walnut-like or a pyramidal shape. The fourth digit had formed the longest metacarpal bone, while that of the fifth was only slightly shorter. The respective bones measured there around 12.5 and 10.7 cm in length. As with ''Glossotherium'' and ''Paramylodon'', only the three inner digits were probably clawed, but only of the second digit have all bone elements been documented. The metacarpal bone was 7.8 cm long and was built very gracefully. The first phalanx was extremely short and only about 2.5 cm long, the second was about 4.2 cm long and the third at least 11.5 cm. It was tubular and went forward into an extension on which the claw rested. The first phalanges of the two outer digits were significantly reduced in length. Only individual root bones of the foot, such as the talus, are present.Integument
''Mylodon'' is one of the few extinct mammals that has mummified skin remains. The most important location for such finds is the Cueva del Milodón in the Chilean province of Última Esperanza, where the first skin parts were brought to light at the end of the 19th century.Otto Nordenskjöld (with the participation of other authors): Scientific results of the Swedish expedition to the Magellan lands 1895-1897, under the direction of Dr. Otto Nordenskjöld. Volume II: Zoology. Stockholm, 1899, pp. 1–170 (especially pp. 149–170) Individual pieces have lengths of up to 150 cm, but have shrunk through drying processes. Its thickness is up to 1.5 cm in some places, but it is usually around 1 cm. The skin is densely covered with stiff, slightly wavy hair, with only the top hair being developed, while the undercoat is missing. This feature is similar to the two-toed sloths but less so than the three-toed sloths, which possess an undercoat. The length of the individual hairs vary between 5 and sometimes over 20 cm with the shortest in the area of the back of the head, medium-length hair on the back and very long hair on the limbs. Their known color ranges from yellowish to reddish-brown. The hair shafts are uniformly tubular, at the upper end they form blunt tips. As with today's sloths, the hair did not have a pith (medulla). In contrast to the hair of the two-toed sloth, they lack their characteristic longitudinal ribbing. The mylodonts are the only representatives of the sloths to have bony plates embedded in their skin. Such structures, called osteoderms, are known today to a greater extent only in armadillos. In contrast to the outer armor of the armadillos, the bone platelets of the mylodonts were rather loosely scattered. Hermann Burmeister published the first finds of individual osteoderms of ''Mylodon'' as early as the 1860s.Hermann Burmeister: skin armor at Mylodon. Archives for anatomy, physiology and scientific medicine 1865, pp. 334–336Hermann Burmeister: Fauna Argentina. Primera party. Mamiferos fósiles. Lista de los mamiferos fósiles del terreno diluviano. Anales del Museo Público de Buenos Aires 1, 1867, pp. 87–300 (p. 173) The remains of skin found in the caves of Última Esperanza give an impression of how they were embedded in the skin and distributed over the body. The bone platelets are all located in the lower section of the skin, while the hairs originate in the upper sections. The distribution turned out to be very inconsistent. Some areas with a dense array of osteoderms contain between 83 and 95 platelets per 10 cm². For others, however, the number is very thin. However, even with a close arrangement, the osteoderms never unite to form a closed shell, but are always separated from one another by individual skin folds. In accordance with the armadillos' shells, the bone platelets form a single layer and do not appear stacked. Since all skin residues were found isolated from the body skeletons, it is sometimes difficult to assign the skin areas with a dense and thin arrangement of bone platelets to a specific part of the body. However, it can be assumed that the back was largely armored and the stomach was free. In the sections with dense osteoderm formation, these were larger than in the clear areas. The bone platelets of ''Mylodon'' were mostly of irregular oval shape with dimensions of 0.5 to 2.5 cm in length, 0.3 to 1.8 cm in width and 0.2 to 1.1 cm in thickness, with weights of a maximum of 2g. On the surface, they showed individual dimples. In cross-section, they consisted of numerous bundles of fibers mixed with hard bone blades (osteoma). This made their structure much simpler than that of the armadillos, and they probably lacked the keratin layer known from the armadillos. In principle, the osteoderms of ''Mylodon'' were similar to those of other large mylodonts.Wilhelm Branco: The application of X-rays in paleontology. Treatises of the Royal Prussian Academy of Sciences Berlin 1906, pp. 1–55Distribution and important fossil finds
Overview and origins
''Mylodon'' was mainly distributed in the southern part of South America. Fossil finds are available from Argentina, Chile, Bolivia, Uruguay and Brazil. Thus, the colonized regions include very far southern sites on the island of Tierra del Fuego as well as most of Patagonia northward to the Pampa region. Its southern limit reached the range at about 53° southern latitude. The Tres Arroyos site on Tierra del Fuego and the region around Cueva del Milodón in southwestern Patagonia are among the southernmost known records of a sloth representative in the Pleistocene.Claudio Latorre: Paleontología de mamíferos del alero Tres Arroyos 1, Tierra del Fuego. In: Anales del Instituto de la Patagonia. Volume 26, 1998, pp. 77-90.H. Gregory McDonald, Gerardo de Iuliis: Fossil history of sloths. In: Sergio F. Vizcaíno, WJ Loughry (Ed.): The Biology of the Xenarthra. University Press of Florida, 2008, pp. 39-55. In the Pampa region, the northern limit was found approximately at the Chuí River in the southeastern Brazilian state of Rio Grande do Sul around 30 degrees south latitude. Even more northerly points of discovery, such as Ñuapua in Bolivia, are tangent to the 20th parallel south. Finds reported from Paraguay, however, are considered rather uncertain. The first occurrence of ''Mylodon'' may have been in the Lower Pleistocene, but finds are rather rare. During this period, the possibly closely related form '' Archaeomylodon'' also occurred in the Pampas region, whose foremost canine teeth of the upper dentition were greatly reduced in size, but not yet completely reduced. Among the early and more northerly finds of ''Mylodon'' is, for example, a skull from the El Palmar Formation in the Argentine province of Entre Ríos, which dates to the end of the last warm period about 80,000 years ago. Also from the northern distribution areas two partial skeletons are worth mentioning, one of which was found at the Río Anisacate in the Argentine province of Córdoba and the other in Arroyo Quequén Salado near Oriente in the Argentine province of Buenos Aires. Mainly in the Pampas, there was an overlap in the occurrence of ''Mylodon'' with the two other major mylodontid sloth representatives '' Glossotherium'' and '' Lestodon'' during the Upper Pleistocene. However, actual co-occurrence is rarely attested. These include the important archaeological site of Paso Otero in Buenos Aires Province, the locality of Arroyo de Vizcaíno in southern Uruguay, and the Chuí River.Important Upper Pleistocene finds
As with many of the other large ground sloths, most of the ''Mylodon'' material is from the Upper Pleistocene, with a focus toward the end of the last glacial period. It is also the phase when ''Mylodon'' again disappeared from the fossil record. From a global perspective, numerous larger animals became extinct during the transition from the Pleistocene to the Holocene, which is why this event is considered a Quaternary extinction wave. In South America, this coincides with the first appearance of humans. Whether the two are causally related is the subject of much controversy. In addition to potential hunting and possible landscape overprinting by early human hunter-gatherer groups, climatic changes may also have had an influence. Numerous archaeological sites, especially in the Pampa region and in the Patagonian area, are between 13,500 and 10,000 years old. The majority of these attest to at least a coexistence of humans and ground sloths over an extended period of time. Direct associations of human cultural products and fossil remains of ''Mylodon'' are found, among others, at Gruta del Indio in the eastern foothills of the Andes, at Piedra Museo or Las Buitreras, all in Argentina, and at Tres Arroyos in Tierra del Fuego, respectively. ''Mylodon'' is often represented by isolated osteoderms, bones or in the form of coprolites, while human remains are limited to stone artifacts and/or hearths. Whether this also involved a more or less intensive raw material use of sloth bones on the part of humans is in many cases unproven. Numerous bone marks that were originally interpreted as anthropogenically caused are, according to recent studies, due to predation. Evidence of direct hunting by humans of the large ground sloths is even more difficult. One piece of evidence is often considered to be Quebrada de Quereo, a site on an ancient coastline in northern Chile. From here come, among other things, skeletal remains of two individuals of ''Mylodon'', distributed in each case over a narrowly defined area, but in two different stratigraphic units and at a spatial distance of 21 m from each other. One of the individuals was associated with about 70 stone objects, whose anthropogenic origin is under discussion. No cut marks are found on the bones as evidence of any human manipulation. The age of the site is given as 11,600 to 10,900 years before present.Donald Jackson S .: Evaluating evidence of cultural associations of Mylodon in the semiarid region of Chile. In: L. Miotti, M. Salemme, M. Flegenheimer (Eds.): Where the south winds blow: ancient evidence of Paleo South Americans. Texas A&M University, 2003, pp. 77-81. One of the most important sites is the Cueva del Milodón near Lago Sofía in the Chilean province of Última Esperanza, known mainly for its surviving skin remains. It is part of a whole system of caves in the region, such as the Cueva del MedioHugo G. Nami, Calvin J. Heusser: Cueva del Medio: A Paleoindian Site and Its Environmental Setting in Southern South America. In: Archaeological Discovery. Volume 3, 2015, pp. 62-71. or the Cueva Chica, which line the southern flank of the 556 m high Cerro Benitez like pearls. Cueva del Milodón is a large cave 250 m long, 140 m wide and 30 m high at the entrance and 10 m at the back, respectively. It was discovered in 1895 by the German captain Hermann Eberhard, who also found the first skin remains. The great importance of these finds led to the cave, initially known as "Cueva Eberhardt", being subsequently visited and explored by numerous scientists. As a result, a large number of finds accumulated over time, among which ''Mylodon'' with bone remains, and numerousTaxonomy
''Mylodons close relatives include the ground sloths of the genera ''Glossotherium'' and ''Paramylodon''. The latter genus has often been confused with ''Glossotherium'', but ''Paramylodon'' is a distinct genus that was restricted to theHistory of research
First description
The taxonomic history of ''Mylodon'' is complex. It has involved confusion and equivocation with other mylodont forms such as '' Glossotherium'' and '' Paramylodon'' over a long period of time. Some of this complexity can be attributed to the first describer of the genus itself, who is''Glossotherium'', ''Grypotherium'' and ''Paramylodon''
Owen, in his 1840 work, established the genus ''Glossotherium'' in addition to ''Mylodon'', but without designation of a specific species. The basis here was a posterior skull fragment taken from the riverbed of the Arroyo Sarandi in thePaleobiology
Diet
The mylodontids (particularly ''Mylodon'' itself) are often considered to be pronounced grazers because of their dental structure with flat chewing surfaces on the molar-like teeth. This is also supported by the high (hypsodont) tooth crowns and the wide mouth with numerous shapes. The ungulates are mostly used as analogous examples, in which shapes with high tooth crowns and broad-lipped mouths are usually grass-eating, such as various cattle, horses or the white rhinoceros. In contrast, those with low tooth crowns and narrow snouts such as the duiker or the black rhinoceros feed largely selective from various leaves and other soft vegetable foods. In contrast to other large mylodontid sloths such as ''Glossotherium'', ''Paramylodon'' or ''Lestodon'', the mouth of ''Mylodon'' is relatively narrow. A special feature is the closed nasal arch, which is heavily roughened in its front area and thus offers muscle attachment points for a mobile upper lip. Something similar can be said about individual depressions in the vicinity of the infraorbital foramen, which also functioned as starting points for individual muscle strands in the nose and lip area. Maybe ''Mylodon'' was more well-adapted to a mixed-vegetation diet, which was picked up with the help of a movable upper lip. The loss of the front teeth in the upper row of teeth also leads to the assumption that, comparable to cattle, there was a horn-like structure on the middle jawbone that could be used to pluck the food. The entire anterior cranial structure of ''Mylodon'' is relatively solid, combined with a partially ossified nasal septum, it can be assumed that relatively high chewing forces acted when the food was chopped up. In contrast to the sometimes huge representatives of the Megatheriidae, the joint between the lower jaw and the skull of the Mylodonts was relatively low, roughly at the chewing level of the teeth. The resulting decreasing lever arm of the masseter muscle experiences through the structure of the zygomatic arch, mainly of the descending process, a certain compensation, so that there should have been only minor differences to the Megatheria with regard to the biting force. The extended mandibular joint allows a wide freedom of movement when chewing. Against this, however, is the zygomatic arch, which is not closed and therefore could only withstand the opposing forces of the masseter and musculus pterygoideus to a limited extent. It can therefore be assumed that forwards and backwards directed chewing movements dominated in ''Mylodon''. The flat tooth crowns lead to a comparatively small size of the total available chewing surface. In ''Mylodon'', this amounts to a good 1320 mm² corresponding to other mylodonts of the same size. The Indian rhinoceros, which is comparable in terms of its dimensions, has, on the other hand, double to four times the value with 2660 to 5190 mm². The situation is similar with the hippopotamus, the total surface area of which is between 3290 and 5410 mm². The small total occlusal surface of the teeth in ''Mylodon'' probably resulted in a rather low processing capacity for the food in the mouth. This can result in either a high rate of fermentation in the gastrointestinal tract and/or a very slow metabolism concluded. The latter is the case with today's sloths. This is due to the long passage time of the food of up to a week through the large, multi-chambered stomach. It can be assumed that this also applies to the extinct sloths. Possibly this made the stomach of the mylodonts a functional equivalent to the complex stomach of the ruminants, whereby a long passage time of the food enabled efficient digestion, in which even more difficult to access nutrients could be provided, for example from foods with a greater fiber content. Such a digestive system could reduce the amount of processed food in the mouth and thus ultimately also have compensated for the small total chewing surface in ''Mylodon''. Direct analysis of the food resources used is possible, among many other things, due to the numerous dung residues in the form of coprolites. These are available not only from the Cueva del Milodón in the Chilean part of Patagonia, but also from other caves. The coprolites of ''Mylodon'' have a diameter of up to 18 cm. Investigations of the plant residues showed 80 to 95% sweet grasses and 5 to 20% sour grasses. Herbaceous plants, on the other hand, could only be detected in traces. Accordingly, ''Mylodon'' led, at least in southwestern Patagonia, a diet consisting almost exclusively of grasses. The food is reflected in the paleohabitat, as pollen analyzes show that the landscape at that time was tundra-like in character and was therefore almost free of trees with only a few low bushes. Occasional evidence of false beeches is interpreted as pollen carried by the wind.Calvin J. Heusser, Luis A. Borrero and José A. Lanata: Late Glacial vegetation at Cueva del Mylodon. Anales del Instituto de la Patagonia (Ciencias Naturales series) 21, 1992, pp. 97-102 A 2021 study on stable isotope ratios concluded that ''Mylodon'' must have been at least sporadically omnivorous.Locomotion
In general, large mylodonts are ground-dwelling animals. The lower section of the hind leg, which is very short compared to the upper, is also found in ''Mylodon'', whose tibia is 27 cm in length and only half as long as the thigh bone, 59 cm in length. In comparison, the Megatheriidae possess significantly longer lower leg portions, about the almost equally-proportioned '' Pyramiodontherium'' possessing to a 47 cm long shin to a 49 cm long femur. Possibly these differences in the hind leg structure result in much more agile locomotion in the Megatheria in relation to the mylodonts. Similar to other large ground sloths, the hand of ''Mylodon'' made contact with the ground with the outer side edge and thus sat up rotated. This is indicated by the long metacarpal bones of the external digits and the decreasing number of phalanges on them. The special hand position protected the long claws of the inner digits, which did not penetrate the ground while walking. A functionally similar but fundamentally different hand position can be found in the ankle duct of the distantly related present-day great anteater. The elbow joint was pointed slightly outwards when standing on four feet and the arms were thus angled slightly inwards, which is evident from the position of the olecranon yields. The hands came to rest slightly within the width of the elbow. Such an orientation of the arms can effectively support the large mass of ''Mylodon''. As a result, the hands would also be in a line with the feet, which is also conveyed, among other things, by footprints from ''Paramylodon''. The laterally limited articular surface of the femoral head severely restricted the mobility of the hindlimbs. The same applies to the forearm, the straight spoke with the laterally elongated head of which did not allow any major rotational movements. These features can be interpreted as adaptations to a purely terrestrial lifestyle. Finally, the muscle attachment points on the first cervical vertebra referenced, which are more developed than for example with ''Paramylodon''. Correspondingly, the occipital joint surfaces are also somewhat further apart. Both can be interpreted as meaning that the more massive skull of ''Mylodon'', caused by the lengthening of the snout region, required greater muscle support. For some of the mylodonts of South America, such as ''Glossotherium'', a partially burrowing way of life is being reconstructed, which results from the construction of the foreleg, among other things. An indicator for this is the upper articular process (olecranon) of the ulna. The longer the olecranon, the higher the leverage of the forearm, since more attachment surface is available for the forearm muscles. In ''Glossotherium'', the olecranon takes up up to 35% of the total length of the ulna. The resulting ability to dig would be comparable to that of the ''Predation and Parasites
Especially in southern and southwestern Patagonia, numerous bone changes in finds of ''Mylodon'' can be proven to be caused by predatory animals. This includes, above all, the remains from the Cueva del Milodón in southwestern Chile. Some caves in their immediate vicinity, such as Cueva Lago Sofía 4 and Cueva Chica, are interpreted as clumps of predators. The same applies to the Cueva del Puma or the Cueva Fell in the Pali-Aike area of southern Chile. Some of the caves mainly contain smaller skeletal elements such as hand and foot bones or bone plates, which indicate that only part of the carcass was carried into the shelter. Whether this is the result of direct foraging or scavenging cannot be determined in many cases. Other caves, in turn, contained a larger proportion of young ''Mylodon'' animals. The largest predators occurring at that time are the puma and theReferences
{{Taxonbar, from=Q310869 Prehistoric sloths Pleistocene xenarthrans Prehistoric placental genera Pleistocene first appearances Holocene extinctions Pleistocene mammals of South America Lujanian Ensenadan Uquian Pleistocene Argentina Fossils of Argentina Pleistocene Chile Fossils of Chile Fossil taxa described in 1840 Taxa named by Richard Owen