Sloths, like other xenarthrans, are an extremely interesting group of mammals that, after a long history of evolution and diversification in South America, became established on islands in the Caribbean and later reached North America during the Great American Biotic Interchange. In all three regions, they were part of the impressive Pleistocene megafauna. Most taxa became extinct and only two small, distantly related tree-dwelling genera survived. Here we incorporate several recently described genera of sloths into an assembled morphological data supermatrix and apply Bayesian inference, using phylogenetic and morphological clock methods, to 64 sloth genera. Thus, we investigate the evolution of the group in terms of the timing of divergence of different lineages and their diversity, morphological disparity and biogeographical history. The phylogeny obtained supports the existence of the commonly recognized clades for the group. Our results provide divergence time estimates for the major clades within Folivora that could not be dated with molecular methods. Lineage diversity shows an early increase, reaching a peak in the Early Miocene followed by a major drop at the end of the Santacrucian (Early Miocene). A second peak in the Late Miocene was also followed by a major drop at the end of the Huayquerian (Late Miocene). Both events show differential impact at the family level. After that, a slight Plio-Pleistocene decline was observed before the marked drop with the extinction at the end of the Pleistocene. Phenotypic evolutionary rates were high during the early history of the clade, mainly associated with Mylodontidae, but rapidly decreased to lower values around 25 Ma, whereas Megalonychidae had lower rates at the beginning followed by a steady increase, peaking during the Late Miocene and the Pliocene. Morphological disparity showed a similar trend, with an early increase, followed by a slowly increasing phase through the Late Oligocene and Early Miocene, and ending with another increase beginning at the middle of the Miocene. Biogeographic analysis showed southern South America as the most probable area of origin of the clade and the main region in which the early diversification events took place. Both Megatheriinae and Nothrotheriinae basal nodes were strongly correlated with Andean uplift events, whereas the early history of Mylodontidae is closely associated with southern South America and also shows an early occupation of the northern regions. Within Megalonychidae, our results show Choloepus as a descendant of an island dispersing ancestor and a probable re-ingression to South America by a clade that originated in Central or North America.
Human–megafauna interaction in the Americas has great scientific and ethical interest because of its implications on Pleistocene extinction. The Arroyo del Vizcaíno site near Sauce, Uruguay has already yielded over 1000 bones belonging to at least 27 individuals, mostly of the giant sloth Lestodon . The assemblage shows some taphonomic features suggestive of human presence, such as a mortality profile dominated by prime adults and little evidence of major fluvial transport. In addition, several bones present deep, asymmetrical, microstriated, sharp and shouldered marks similar to those produced by human stone tools. A few possible lithic elements have also been collected, one of which has the shape of a scraper and micropolish consistent with usage on dry hide. However, the radiocarbon age of the site is unexpectedly old (between 27 and 30 thousand years ago), and thus may be important for understanding the timing of the peopling of America.
For over 200 years, fossils of bizarre extinct creatures have been described from the Americas that have ranged from giant ground sloths to the ‘native’ South American ungulates, groups of mammals that evolved in relative isolation on South America. Ground sloths belong to the South American xenarthrans, a group with modern although morphologically and ecologically very different representatives (anteaters, armadillos and sloths), which has been proposed to be one of the four main eutherian clades. Recently, proteomics analyses of bone collagen have recently been used to yield a molecular phylogeny for a range of mammals including the unusual ‘Malagasy aardvark’ shown to be most closely related to the afrotherian tenrecs, and the south American ungulates supporting their morphological association with condylarths. However, proteomics results generate partial sequence information that could impact upon the phylogenetic placement that has not been appropriately tested. For comparison, this paper examines the phylogenetic potential of proteomics-based sequencing through the analysis of collagen extracted from two extinct giant ground sloths, Lestodon and Megatherium. The ground sloths were placed as sister taxa to extant sloths, but with a closer relationship between Lestodon and the extant sloths than the basal Megatherium. These results highlight that proteomics methods could yield plausible phylogenies that share similarities with other methods, but have the potential to be more useful in fossils beyond the limits of ancient DNA survival.
Glyptodonts are a group of extinct xenarthrans with several anatomical features that make them one of the most bizarre groups of mammals. By the late 19th century, some authors began to analyze the brain of Pleistocene glyptodonts using natural endocranial casts. These studies revealed the small size of the brain of the large Pleistocene forms. However, the evolution of the brain in glyptodonts and how it fits in a phylogenetic context has not been analyzed. In order to evaluate the evolution of the brain in this group, we described the first digital endocranial cast of the late Miocene glyptodont Pseudoplohophorus absolutus and compared it with digital endocranial casts of the Pleistocene glyptodonts Glyptodon, Doedicurus, and Panochthus and the extant armadillos Dasypus, Euphractus, Chaetophractus, and Zaedyus. The endocast morphology of P. absolutus is similar to that of Pleistocene glyptodonts: large olfactory bulbs, a small cerebrum with a single neocortical sulcus, and a large cerebellum. However, the relative brain size is larger than in the Pleistocene forms, with values of the encephalization quotient (EQ) close to that of extant armadillos. A comparison between xenarthrans orders shows that Cingulata (glyptodonts and armadillos) have lower EQ values than Pilosa (sloths and vermilinguas). This could possibly be related to certain restrictions and benefits imposed by the presence of the carapace in cingulates. Furthermore, because the carapace restricts the development of the cervical musculature that supports the skull, the small size of the brain in glyptodonts could be a trade-off (along with others) to reduce the weight of the skull.
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