The dermal armor of mylodontid sloths (Mammalia, Xenarthra) from Cueva del Milodón (Última Esperanza, Chile).

Dermal ossifications (osteoderms, dermal ossicles, osteoscutes) appear independently in various tetrapod lineages. In mammals, however, dermal ossifications are only present in some members of Xenarthra. This clade includes Cingulata (armadillos and their relatives), and Pilosa, including Vermilingua (anteaters) and Folivora (sloths). In extant xenarthrans, osteoderms are invariably present in cingulates whereas they are absent in pilosans. Among extinct sloths, however, a limited number of taxa possessed dermal ossifications. Records of mummified skins of ground sloths bearing osteoderms found in Cueva del Milodón (Southern Chile), with a late Pleistocene age, allowed us to analyze their micro- and macroscopic morphology. The main goal of this study is to closely examine a portion of a mylodontid skin portion using radiography. The arrangement, morphology and internal structure of the ossicles are analyzed and the results are discussed in the context of previous research. The results we obtained indicate that ossicles vary in shape and size, and the integument has four different patterns of arrangement of the ossicles: that is, areas without ossicles, disorganized ossicles, rows, and mosaic areas. The latter has two variants, with clusters of ossicles forming rosettes or stars. Thin sections of the ossicles allowed us to recognize and describe anatomical features of the bone and its mode of growth. Finally, paleobiological and functional considerations of the dermal armor are discussed along with its phylogenetic and chronological implications.

Phylogenetic signal analysis in the basicranium of Ursidae (Carnivora, Mammalia).

Ursidae is a monophyletic group comprised of three subfamilies: Tremarctinae, Ursinae and Ailuropodinae, all of which have a rich geographical distribution. The phylogenetic relationships within the Ursidae group have been underexamined, especially regarding morphological traits such as the basicranium. Importantly, the basicranium is a highly complex region that covers a small portion of the skull, combining both structural and functional aspects that determine its morphology. Phylogenetic hypotheses of the Ursidae (including Tremarctinae) have been made based on morphological characters that considers skull, mandible and teeth features, while specific characters of the auditory region and basicranium have not been taken into account. To do this, we analyse the shape and size macroevolution of the basicranium of Ursidae, testing its morphological disparity in a phylogenetic context, which is quantified by means of the phylogenetic signal. We investigated phylogenetical autocorrelation by shape (depicted by Principal Components Analysis scores from previous published analyses) and basicranium size (depicted by centroid size, CS) using an orthonormal decomposition analysis and Abouheif C mean. The main advantages of these methods are that they rely exclusively on cladogram topology and do not require branch-length estimates. Also, an optimisation of the ancestral nodes was performed using TNT 1.5 software. In relation to the phylogenetic signal, both methods showed similar results: the presence of autocorrelation was detected in PC1 and PC2, while in PC3, PC4 and PC5 and in the size of the basicranium (CS), the absence of autocorrelation occurred. The most significant nodes (where there is autocorrelation) are the basal nodes 'Ursidae' and 'Ursinae-Tremarctinae'. Within this last group, distinctive basicranium morphology is observed, being more conservative in Tremarctinae than in Ursinae. The differences between these subfamilies could be related to historical events involving varying food and environmental preferences. The high phylogenetic signal in the node Tremarctinae probably indicates that the basicranium configuration of these bears was obtained early in their evolutionary history. Finally, our results of the basicranium and skull length ratios indicate that in Tremarctinae, the basicranium size was not determined by phylogeny but instead by other factors, such as adaptive responses to climatic changes and competition with other carnivores.

Comparative osteology and functional morphology of the forelimb of Cyonasua (Mammalia, Procyonidae), the first South American carnivoran.

Extant procyonids only inhabit the Americas and are represented by six genera (Procyon, Nasua, Nasuella, Bassaricyon, Potos, and Bassariscus); all of them, except Bassariscus, are present in South America. The first records correspond to the early Miocene in North America (NA) and the late Miocene in South America (SA). Cyonasua was the first carnivoran to enter SA from NA, before the Great American Biotic Interchange, and went extinct in the early middle Pleistocene. This extinct procyonid is recorded in several localities of Argentina, and also in Venezuela. Paleobiological studies of procyonids are interesting from evolutionary and biogeographical viewpoints. In this study, the pectoral girdle and forelimb of 10 specimens of Cyonasua are described and compared with extant South American procyonids using a qualitative approach. Additionally, four functional morphology indexes were calculated for them and compared with an ecologically diverse sample of living carnivorans. Results indicate that Cyonasua most resembles Nasua nasua and Procyon cancrivorus, even though the extinct procyonid possessed peculiar features. Cyonasua had robust and relatively short forelimb bones, with strong stabilized joints, and movements associated with the sagittal plane, which suggest a tendency toward terrestrial habits, related to their ability to resist relatively high bending and shearing stresses. However, some features indicate a freedom in their range of movements, with moderate supination ability, compatible with climbing. When combined with previous analyses of dietary habits and estimated body mass, the morphology of Cyonasua would be well suited for digging and prey manipulation, allowing them to prey on small and relatively large-sized vertebrates, as well as to avoid some of the predators that were dominant in the Cenozoic of South America.

Muscular reconstruction and functional morphology of the hind limb of santacrucian (Early Miocene) sloths (Xenarthra, Folivora) of Patagonia.

This article presents a morphofunctional analysis of the hind limb of Santacrucian (Early Miocene) sloths from southernmost Patagonia (Argentina). These fossil sloths were mid sized to large animals, ranging from 40 to 120 kg, and their postcranial skeleton was markedly different in shape compared with that of extant tree sloths, which vary from 2 to 10 kg. The functional anatomy of the hind limb of Santacrucian sloths was compared with that of living xenarthrans (tree sloths, anteaters, and armadillos), which involved reconstruction of the hind limb musculature and comparative and qualitative morphofunctional analyses, and hypotheses on the biological role of the hind limb in terms of preferences in substrate, posture, and strategies of locomotion were formulated. The hind limb of Santacrucian sloths bears strong resemblances to that of living South American anteaters in stoutness of skeletal elements, form of the characteristics related to muscular and ligamentous attachments, and conservative, pentadactylous strong-clawed pes. The musculature was very well developed, allowing powerful forces, principally in entire limb adduction, crus flexion and extension, pes extension, and toe prehension. These functional features, together with those of the forelimb, are congruent with climbing behavior, and support the hypothesis that Santacrucian sloths were good but slow climbing mammals. However, their climbing strategies were limited, owing principally to their comparatively large body size, and they relied to a large extent on their powerful musculature and curved manual and pedal unguals for both moving and standing on the arboreal supports.

Muscular reconstruction and functional morphology of the forelimb of early Miocene sloths (Xenarthra, Folivora) of Patagonia.

Early Miocene sloths are represented by a diversity of forms ranging from 38 to 95 kg, being registered mainly from Santacrucian Age deposits in southern-most shores of Patagonia, Argentina. Their postcranial skeleton differs markedly in shape from those of their closest living relatives (arboreal forms of less than 10 kg), Bradypus and Choloepus. In order to gain insight on functional properties of the Santacrucian sloths forelimb, musculature was reconstructed and a comparative, qualitative morphofunctional analysis was performed, allowing proposing hypotheses about biological role of the limb in substrate preferences, and locomotor strategies. The anatomy of the forelimb of Santacrucian sloths resembles more closely extant anteaters such as Tamandua and Myrmecophaga, due to the robustness of the elements, development of features related to attachment of ligaments and muscles, and conservative, pentadactylous, and strong-clawed manus. The reconstructed forelimb musculature was very well developed and resembles that of extant Pilosa (especially anteaters), although retaining the basic muscular configuration of generalized mammals. This musculature allowed application of powerful forces, especially in adduction of the forelimb, flexion and extension of the antebrachium, and manual prehension. These functional properties are congruent with both climbing and digging activities, and provide support for proposed Santacrucian sloths as good climbing mammals, possibly arboreal or semiarboreal, being also capable diggers. Their climbing strategies were limited, thus these forms relied mainly on great muscular strength and curved claws of the manus to move cautiously on branches.

Morphology and function of the hyoid apparatus of fossil xenarthrans (mammalia).

The analysis of the hyoid apparatus of fossil xenarthrans provides insight on the form of the tongue and its function in food intake and intraoral processing. The hyoid apparatus of xenarthrans is notable for fusion among its elements. The presence of a V-bone, a complex consisting of fused basihyal and thyrohyal bones, is a consistent and probably synapomorphic feature of xenarthrans. Fusion of other elements is variable in fossil xenarthrans. Most fossil sloths retain independent elements, as in living dasypodids and mammals generally. Among nothrotheriids, the elements are slender and their articular surfaces indicate considerable mobility, and the relatively long and horizontal orientation of the geniohyoid muscle suggests considerable tongue protrusion. Among mylodontines, such as Paramylodon and Glossotherium, the elements indicate relatively mobile articulations, except between the stylo- and epihyals. The relatively posterior placement of the apparatus and the length and alignment of the geniohyoid muscle indicate considerable capacity for tongue protrusion. Scelidotherium, however, had rigidly articulated stylohyal and epihyal, and the apparatus lies farther anteriorly, which together with the elongated, steeply inclined mandibular symphysis, indicates a relatively shorter geniohyoid muscle and thus more limited capacity for tongue protrusion. A similar situation is indicated for Megatherium, casting doubt on the classical reconstruction of this sloth as having a long prehensile tongue. Among cingulates Prozaedyus resembles living dasypodids, indicating considerable tongue protrusion important in food acquisition and intake. More extensive fusion of hyoid elements occurs in the cingulates Glyptodon and Proeutatus, in which the stylohyal and epihyal at least, are fused into a single element termed the sigmohyal. The presence of this element supports recent proposals of a sister-group relationship between glyptodonts and eutatines. The rigidity of the apparatus suggests limited tongue protrusion, but the tongue, in glyptodonts at least, was a powerful structure important for intraoral manipulation of food.

Muzzle of South American Pleistocene ground sloths (Xenarthra, Tardigrada).

Sloths are among the most characteristic elements of the Cainozoic of South America and are represented, during the Pleistocene, by approximately nine genera of gigantic ground sloths (Megatheriidae and Mylodontidae). A few contributions have described their masticatory apparatus, but almost no attention has been paid to the reconstruction of the muzzle, an important feature to consider in relation to food intake, and particularly relevant in sloths because of the edentulous nature of the muzzle and its varied morphology. The relationship between dietary habits and shape and width of the muzzle is well documented in living herbivores and has been considered an important feature for the inference of alimentary styles in fossils, providing an interesting methodological tool that deserves to be considered for xenarthrans. The goal of this study was to examine models of food intake by reconstructing the appearance and shape of the muzzle in five species of Pleistocene ground sloths (Megatherium americanum, Glossotherium robustum, Lestodon armatus, Mylodon darwini, and Scelidotherium leptocephalum) using reconstructions of the nasal cartilages and facial muscles involved in food intake. The preservation of the nasal septum, and the scars for muscular attachment in the rostral part of the skulls, allow making a conservative reconstruction of muzzle anatomy in fossil sloths. Wide-muzzled ground sloths (Glossotherium and Lestodon) had a square, nonprehensile upper lip and were mostly bulk-feeders. The lips, coupled with the tongue, were used to pull out grass and herbaceous plants. Narrow-muzzled sloths (Mylodon, Scelidotherium, and Megatherium) had a cone-shaped and prehensile lip and were mixed or selective feeders. The prehensile lip was used to select particular plants or plant parts.