Mylodon is an extinct genus of ground sloth belonging to the family Mylodontidae, known from southern South America. With a total length of 3 to 4 m and a body mass of 1-2 tonnes, it is one of the largest mylodontids (though it was considerably exceeded in size by the mylodontid Lestodon).
The oldest finds probably date to the Lower Pleistocene; however, most of the fossil remains date to the Late Pleistocene. Its distribution ranged from southern Bolivia and the Pampas in the north southwards to the southernmost part of Patagonia at around 52-53 degrees south, the furthest south of any Pleistocene ground sloth, with some of the best known remains of the species being from the eponymous Cueva del Milodon (cave of Mylodon) in southern Chile.
In addition to skeletal remains, Mylodon is also known from preserved skin and hair. Preserved dung (Coprolite) indicates that Mylodon was a primarily a grazer, feeding on grasses and sedges.
Mylodon has generally only a single recognised species, Mylodon darwini, which was described by Richard Owen in 1840 based on remains collected in the Pampas by Charles Darwin (for whom the species is named) during the Voyage of the Beagle. Some recent authors suggest that there were two species, with M. darwini restricted to the Pampas, with the Patagonian remains belonging to the separate species Mylodon listai.
Mylodon went extinct at the end of the Late Pleistocene-beginning of the Holocene, around 12-10,000 years ago, as part of the end-Pleistocene extinction event, along with other ground sloths and most large animals across the Americas. Mylodon chronologically overlapped with Paleo-Indians, the first humans to inhabit the Americas, evidence from several sites suggest that they may have hunted Mylodon. The extinction of Mylodon may be the result of climatic change, hunting by Paleoindians, or a combination of both factors.
Well preserved samples of Mylodon remains have been discovered in the Cueva del Milodón site in Patagonia, Chile along the southern flank of Cerro Benítez in the year 1896. Associated with bones of other early Patagonian animals, these remains of Mylodon date from an era earlier than 10,000 BC. C. Michael Hogan (2008) Cueva del Milodon, Megalithic Portal The American Museum of Natural History has exhibited a sample of Mylodon dung from Argentina with a note that reads "deposited by Theodore Roosevelt".
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, Mylodon's 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 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 often caniniform while the rear teeth were more molariform. Within the sloth, this structure of the teeth can be called original. A special feature of Mylodon was that the upper canine-like tooth of each row was completely regressed and only the molar-like four rear teeth were found here. In the lower row of teeth, the anterior caniniform tooth was transformed into a molariform. The dentition thus consisted of a total of 16 teeth. This is somewhat reminiscent of Paramylodon, in which the upper canine-shaped teeth were also missing, but the lower ones had retained their strikingly pointed shape. In contrast to this, Glossotherium and Lestodon had the original decayed teeth. The flat, flap-like and largely indented structure of the molariform teeth can be emphasized as a characteristic of the mylodonts, which clearly differs from that of the Megatheriidae and Megalonychidae with their two transverse raised ridges per tooth. The shapes of the teeth present in Mylodon were simpler. In the upper jaw row, they had a rather round to oval outline, in the lower jaw row a more diamond-shaped outline. The typically more complex bilobed design of the molar-like teeth of Glossotherium and Lestodon, caused by a central constriction, only occurred on the lower rearmost tooth in Mylodon. In general, the rows of teeth diverged to the front, and the teeth were very high crowned (hypsodont). The upper row of teeth ranges in length from 10.9 to 13.3 cm, the lower row was between 12.0 and 15.0 cm in length.
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.
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 2 g. 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. 1–55
The first occurrence of Mylodon may have been in the Lower Pleistocene, but finds are rather rare.
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 numerous has a large share. Other finds belong to camels such as Lama, Equidae such as Hippidion or South American ungulates such as Macrauchenia, in addition, several predators are represented, including the jaguar, Smilodon as a member of the saber-toothed cats, and the giant bear form Arctotherium. Some of the mammal bones have marks that were originally associated with human activity, but the current view is that they are more likely due to predator browsing. In addition to faunal remains, the cave also held a myriad of botanical material. It also yielded one of the most extensive data sequences from the Upper Pleistocene. Several radiocarbon dates, measured from a wide variety of Mylodon finds, span a period from about 16,700 to 10,200 years ago. The upper data are among the most recent obtained directly from finds of the sloth vertebrate.
Below is a phylogenetic tree of the Mylodontidae, based on the work of Boscaini et al.. 2019.
Recent molecular sequence results obtained using collagen and mitochondrial DNA extracted from fossils indicate that the closest living relatives of Mylodon are the two-toed sloths of genus Choloepus. This revelation came as a surprise, since morphological analyses had previously suggested that two-toed sloths were close to and Megalonyx, now regarded as representing two separate and distant branches of the sloth evolutionary tree.
Some authors suggest that there were two species, with M. darwini restricted to the Pampas, with the Patagonian remains belonging to the separate species Mylodon listai.
A skull including mandible found at Pergamino in Buenos Aires Province served the Danish zoologist Johannes Theodor Reinhardt (1816-1882) in 1879 as the basis for a comprehensive description. Characteristic of the skull was the narrow snout and a closed nasal arch, which was formed by the firm adhesion of the nasal bone to the middle jaw bone. Furthermore, the dentition consisted of a total of 16 teeth, the upper front canine-like tooth was reduced in each case, in the lower jaw, however, sat four molar-like teeth each. Reinhardt noticed similarities to M. darwinii in the construction of the lower jaw, but in the skull design his find deviated clearly from the broad-nosed M. robustus by the narrow snout. However, according to Reinhardt, there were matches in Owen's Glossotherium skull fragment with corresponding skull sections in M. robustus. Based on the clear parallels between M. darwinii and his narrow-beaked skull finding, Reinhardt propagated the new genus Grypotherium with Grypotherium darwinii as the type species.Johannes Theodor Reinhardt: Beskrivelse af Hovedskallen af et Kæmpedovendyr, Grypotherium darwini. In: Det Kongelige Danske Videnskabernes Selskabs Skrifter. 5 Række. Naturvidenskabelig og Mathematisk Afdeling 12 (5), 1879, pp. 351–381 ( archive.org ). A different approach was taken by Florentino Ameghino (1854-1911) about ten years later. He confirmed in 1889, the separation of both M. darwinii and M. robustus not only at the species level, but also at the genus level. Deviating from Reinhardt, but agreeing with Owen, he considered the mandible of M. darwinii and the skull fragment of Glossotherium to belong together. Since in this scenario Glossotherium possessed priority over Mylodon (Owen mentioned the former over the latter in 1840) and Grypotherium, Ameghino introduced the species Glossotherium darwinii. In contrast, he left the status of M. robustus untouched.Florentino Ameghino: Contribución al conocimiento de los mamíferos fósiles de la República Argentina. In: Actas de la Academia Nacional de Ciencias. Volume 6, 1889, pp. 1-1027 (pp. 734-744). Arthur Smith Woodward (1864-1944), in turn, followed Reinhardt's reasoning. In a paper published in 1900, he presented findings from ground sloths of southern Patagonia and at the same time revised Charles Darwin's collection. In doing so, he equated the mandible of M. darwinii with Reinhardt's Grypotherium and subsequently recovered Grypotherium darwinii. Smith Woodward assigned the skull fragment of Glossotherium to M. robustus by analogy with Reinhardt.
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% grasses and 5 to 20% sedges. 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.
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 Tolypeutes armadillos, which seldom build their own burrows, but can do so. The previous analyses for Mylodon resulted in a much shorter olecranon, which accounts for only about 22% of the total length of the ulna. However, the fact that proportional estimates for Mylodon refer to a not fully grown specimen is problematic. Other clues can be derived from the construction of the hand. In Mylodon, for example, the metacarpal bones of the second and third ray are very delicate, in contrast to Glossotherium. A weakly pronounced central ray does not seem to support a digging activity, as this is usually most strongly developed in underground mammals. However, the distal articular facet of the third metacarpal bone is flat, which means that the middle finger is generally stiff and stable. The same articulation surface on the second metacarpal is significantly more rounded and thus supports greater mobility of the finger when gripping. This obviously resulted in functional differences between the individual rays of the hand. The rare signs of wear and tear on the last phalanx, which are isolated from the Cueva del Milodón several times, can serve as an additional indicator of digging activities.
In various coprolites produced by Mylodon, eggs of nematodes are preserved. The eggs are ovaloid in shape with lengths of almost 50 μm in length and 29 μm in thickness. In addition, individual beetles could be detected.
Taxonomy
History of research
First description
Glossotherium, Grypotherium and Paramylodon
Paleobiology
Diet
Locomotion
Predation and Parasites
Extinction
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