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.

Species distribution models (SDMs) for the last interglacial (LIG), the global last glacial maximum (LGM) and the Holocene climatic optimum (HCO) were generated for three extinct South American Pleistocene mylodontid giant sloths, Glossotherium robustum, Lestodon armatus and Mylodon darwinii. They are recorded co-occurring in some localities including Arroyo del Vizcaíno site (AdV) in Uruguay. Co-occurrence records were studied based on the overlap of their generated areas of potential distributions, and compared with the available biome reconstructions of South America during the LGM to analyze their distribution patterns, ecological requirements and possible interactions between them. Our results suggest that these sloths could have co-existed mainly in the Chaco-Paraná Basin and the plains in the Río de la Plata area. Areas of high suitability were observed for submerged parts of the continental shelf that were exposed during the LGM showing an overall increase in potential habitat compared to the LIG and HCO. This suggests that there was a drastic reduction in total available areas of preferred habitat at the end of the Pleistocene. The co-occurrence of these sloths at the AdV site suggests the presence of vegetation indicative of mainly open, cold to temperate habitats but with mixed patches typical of humid climates.

The moment arms of jaw muscles and the hypsodonty index (HI) have been widely used to infer diet preferences in extinct species. Here, we intend to evaluate the use of the masseter moment arm and HI as dietary proxies in herbivorous ungulates and analyse its relations to diet preference, habitat selection and two continuous variables: percentage of grass in natural diet (%Grass) and annual precipitation (PREC). Masseter moment arm (Mm) showed a strong correlation to diet preference and %Grass but not for habitat selection and PREC. Contrarily, HI showed similar correlation with diet preference and %Grass but also with habitat selection and PREC. The results show that Mm predominately reflects adaptation to tougher, rich in grass diets, while HI, as previously found, indicates adaptation to the wear effect of both diet and environment. This approach has been useful in identifying the habits of two extinct very large Pleistocene South American ungulates. Macrauchenia patachonica, an extinct litoptern, is interpreted as an openhabitat high browser and Toxodon platensis, an extinct notoungulate, as a grazer. Figure 6 Relationships between (a) the masseter moment arm and (b) the hypsodonty index and the mean annual precipitation at the centre of a species' distribution for the studied species. L. Varela and R. A. Fariña Masseter moment arm as a dietary proxy Journal of Zoology 296 (2015) 295-304