Recent studies show Xenarthra to be even more isolated systematically from other placental mammals than traditionally thought. The group not only represents 1 of 4 primary placental clades, but proposed links to other fossorial mammal taxa (e.g., Pholidota, Palaeanodonta) have been contradicted. No unambiguous Paleocene fossil xenarthran remains are known, and Eocene remains consist almost exclusively of isolated cingulate osteoderms and isolated postcrania of uncertain systematic provenance. Cingulate skulls are unknown until the late middle Eocene, and the oldest sloth and anteater skulls are early Oligocene and early Miocene age, respectively; there are no nearly complete xenarthran skeletons until the early Miocene. Ecological reconstructions of early xenarthrans based on extant species and the paleobiology of extinct Neogene taxa suggest the group’s progenitors were myrmecophagous with digging and perhaps some climbing adaptations. The earliest cingulates were terrestrial diggers and likely myrmecophagous but soon diverged into numerous omnivorous lineages. Early sloths were herbivores with a preference for forested habitats, exhibiting both digging and climbing adaptations. We attribute the rarity of early xenarthran remains to low population densities associated with myrmecophagy, lack of durable, enamel-covered teeth, and general scarcity of fossil localities from tropical latitudes of South America. The derivation of numerous omnivorous and herbivorous lineages from a myrmecophagous ancestor is a curious and unique feature of xenarthran history and may be due to the peculiar ecology of the native South American mammal fauna. Further progress in understanding early xenarthran evolution may depend on locating new Paleogene fossil sites in northern South America. Los estudios sistemáticos recientes muestran que, a nivel sistemático, los xenartros están aún más aislados de otros mamíferos placentarios de lo que se pensaba tradicionalmente. El grupo no sólo representa una de las cuatro ramas principales de los Placentalia, sino que también se han refutado las hipótesis previas de posibles conexiones con otros taxones de mamíferos fosoriales (por ejemplo Pholidota, Palaenodonta). No se conocen restos fósiles inequívocos de xenartros del Paleoceno y los restos provenientes del Eoceno consisten casi exclusivamente de osteodermos aislados de cingulados y restos postcraneanos aislados de origen sistemático incierto. No se conocen cráneos razonablemente completos de cingulados hasta finales del Eoceno medio; los cráneos más antiguos de perezosos y osos hormigueros provienen del Oligoceno temprano y del Mioceno temprano, respectivamente; y no existen esqueletos completos o casi completos de ninguno de los 3 linajes hasta el Mioceno temprano. Reconstruimos la ecología de los primeros xenartros basándonos en las especies actuales y lo que se sabe de la paleobiología del Mioceno y de los taxones extintos más recientes. Nuestros resultados sugieren que los primeros xenartros eran mirmecófagos y poseían adaptaciones para cavar y tal vez para trepar. Los primeros cingulados eran cavadores terrestres y probablemente mirmecófagos, pero pronto divergieron en numerosos linajes omnívoros. Nuestras reconstrucciones indican que los primeros perezosos eran herbívoros con preferencia de hábitats boscosos, tal vez exhibiendo adaptaciones tanto para cavar como para trepar. Atribuimos la rareza de restos de los primeros xenartros a varios factores: bajas densidades poblacionales asociadas a hábitos mirmecófagos; falta de dientes duraderos y cubiertos de esmalte; y una escasez general de localidades de mamíferos tempranos de las latitudes tropicales de América del Sur. La derivación de numerosos linajes omnívoros y herbívoros de un ancestro mirmecófago es un rasgo curioso y único de la historia de los xenartros y puede deberse a la peculiar ecología de la fauna de mamíferos sudamericanos. Los nuevos avances en la comprensión de la evolución temprana de los xenartros podrían depender de la localización de nuevos sitios fósiles paleógenos en áreas de tierras bajas poco accesibles del norte de América del Sur.
The specimen described herein and assigned to 'Xyophorus' sp. (Mammalia, Xenarthra, Tardigrada) was collected in the locality Cerro Zeballos, northwestern Chubut Province, Argentina. The fossiliferous sediments bearing the specimen are correlated with Collón Curá Formation. The specimen has the features described for other members of 'Xyophorus' (e.g. shape and size of the molariforms, relationship between diastema length, m1 and m2 length) and has a Diastema Length/Tooth Row Length index (DL/TRL index) of ca. 14, between that of 'X.' villarroeli (12.07) from the Mauri Formation, Bolivia (ca. 10.3 Ma) and that of 'X.' bondesioi (16.45) from Arroyo Chasicó Formation, Argentina (ca. 10-8.7 Ma). The relationship between DL/TRL index and age of the bearing sediments, would suggest a Tortonian age (late Miocene) for the deposits of Collón Curá Formation at Cerro Zeballos, which results in a 'younger age' compared to the middle Miocene age traditionally accepted for the Collón Curá Formation bearing the Colloncuran fauna sensu stricto. Although no absolute ages for Cerro Zeballos are available yet, the geographic proximity of Cerro Zeballos to Cushamen River (with levels dated at ca. 11.2 Ma) supports the tentative Tortonian age indicated by the presence of 'Xyophorus' sp.
Glyptodonts were giant (some of them up to ~2400 kg), heavily armoured relatives of living armadillos, which became extinct during the Late Pleistocene/early Holocene alongside much of the South American megafauna. Although glyptodonts were an important component of Cenozoic South American faunas, their early evolution and phylogenetic affinities within the order Cingulata (armoured New World placental mammals) remain controversial. In this study, we used hybridization enrichment and high‐throughput sequencing to obtain a partial mitochondrial genome from Doedicurus sp., the largest (1.5 m tall, and 4 m long) and one of the last surviving glyptodonts. Our molecular phylogenetic analyses revealed that glyptodonts fall within the diversity of living armadillos. Reanalysis of morphological data using a molecular ‘backbone constraint’ revealed several morphological characters that supported a close relationship between glyptodonts and the tiny extant fairy armadillos (Chlamyphorinae). This is surprising as these taxa are among the most derived cingulates: glyptodonts were generally large‐bodied and heavily armoured, while the fairy armadillos are tiny (~9–17 cm) and adapted for burrowing. Calibration of our phylogeny with the first appearance of glyptodonts in the Eocene resulted in a more precise timeline for xenarthran evolution. The osteological novelties of glyptodonts and their specialization for grazing appear to have evolved rapidly during the Late Eocene to Early Miocene, coincident with global temperature decreases and a shift from wet closed forest towards drier open woodland and grassland across much of South America. This environmental change may have driven the evolution of glyptodonts, culminating in the bizarre giant forms of the Pleistocene.
The hyoid apparatus reflects aspects of the form and function of feeding in living and extinct organisms and, despite the availability of information about this structure for Xenarthra, it remains little explored from an evolutionary perspective. Here we compare the morphology of the hyoid apparatus in xenarthrans, describing its general morphology and variation in each major clade and score these variations as phylogenetic characters, which were submitted to ancestral states reconstructions. The general hyoid morphology of Xenarthra consists of a v-bone (basihyal fused with the thyrohyals) and three paired bones (stylohyal, epihyal and ceratohyal), which are unfused in the majority of taxa. The clade-specific morphology observed here, allowed us to obtain additional synapomorphies for all major clades of Xenarthra (Cingulata, Pilosa, Folivora and Vermilingua), for Glyptodontididae, and for Nothrotheriidae. The fusion of hyoid elements are convergentelly achieved among the diphyletic extant tree sloths, some extinct ground sloths and glyptodontids. Despite the heavy influence of adaptive evolution related to feeding habits, the morphology of the hyoid apparatus proved to be a valuable source of phylogenetic information.
Xenarthrans comprised an ecologically significant and diverse group of small to gigantic sized terrestrial insectivorous, omnivorous and herbivorous mammals during the Cenozoic in South America and during the Pleistocene in North America. Their peculiar tooth morphology has proven to be challenging for palaeodietary analyses of this group. Here we introduce a new approach to this problem by utilising the recently developed mesowear angle analysis for xenarthran palaeodietary analyses. The method is based on recording the relief of worn teeth as angles measured from the occlusal surfaces. We compare our results with other lines of evidence of extant and fossil xenarthran diets, based on direct observation, orthodentine microwear analyses and analyses of fossilised faecal material. Our results support previous findings and hypotheses on fossil xenarthran diets, but also provide new information on the diversity of dietary preferences in the diverse assemblages of large Pleistocene xenarthrans such as ground sloths and glyptodonts.
The orientation of the semicircular canals of the inner ear in the skull of vertebrates is one of the determinants of the capacity of this system to detect a given rotational movement of the head. Past functional studies on the spatial orientation of the semicircular canals essentially focused on the lateral semicircular canal (LSC), which is supposedly held close to horizontal during rest and/or alert behaviors. However, they generally investigated this feature in only a few and distantly related taxa. Based on 3D‐models reconstructed from µCT‐scans of skulls, we examined the diversity of orientations of the LSC within one of the four major clades of placental mammals, that is, the superorder Xenarthra, with a data set that includes almost all extant genera and two extinct taxa. We observed a wide diversity of LSC orientations relative to the basicranium at both intraspecific and interspecific scales. The estimated phylogenetic imprint on the orientation of the LSC was significant but rather low within the superorder, though some phylogenetic conservatism was detected for armadillos that were characterized by a strongly tilted LSC. A convergence between extant suspensory sloths was also detected, both genera showing a weakly tilted LSC. Our preliminary analysis of usual head posture in extant xenarthrans based on photographs of living animals further revealed that the LSC orientation in armadillos is congruent with a strongly nose‐down head posture. It also portrayed a more complex situation for sloths and anteaters. Finally, we also demonstrate that the conformation of the cranial vault and nuchal crests as well as the orientation of the posterior part of the petrosal may covary with the LSC orientation in Xenarthra. Possible inferences for the head postures of extinct xenarthrans such as giant ground sloths are discussed in the light of these results. Orientation of the lateral semicircular canal of the inner ear within the skull of extant xenarthrans (Mammalia). Upper skull: in the extant armadillo Dasypus kappleri . Lower skull: in the extant three‐toed sloth Bradypus tridactylus . The red lines mark the plane of the lateral semicircular canal, which is more oblique in armadillos than in sloths (relative to the palate).
The relationship between humerus shape and the modes of exploring substrate among extinct and extant Pilosa (especially anteaters and ground sloths) were investigated here. We used geometric morphometrics and discriminant analyses to relate morphological patterns and their possible ecological categories. Our results suggest that plesiomorphic taxa such as Nothrotheriidae, most Megalonychidae and basal Megatheriidae tend to have more slender humerus, associated to generalist habitus (climbing, swimming and digging activities), and while Mylodontidae developed specialized digging habitus. Additionally, we inferred ground sloths which inhabited the Brazilian territory during the Quaternary likely occupied at least four different niches. Mammals display morphofunctional adaptations on the limbs which are reflected on their modes of substrate exploration. Herein, we analyzed the humerus morphology of ground sloths and anteaters. Our results suggest that most of the Pleistocene Mylodonts were fossorial taxa, while most of the Santacrucian sloths plus extant anteaters were semiarboreal or semiaquatic taxa. The Pleistocene Megatheriidae should be ambulatory. Mammals display morphofunctional adaptations on the limbs which are reflected on their modes of substrate exploration. Herein, we analyzed the humerus morphology of ground sloths and anteaters. Our results suggest that most of the Pleistocene Mylodonts were fossorial taxa, while most of the Santacrucian sloths plus extant anteaters were semi‐arboreal or semi‐aquatic taxa. The Pleistocene Megatheriidae should be ambulatory
Here, we review population genetic, phylogeographic, and phylogenetic studies on xenarthrans and show how this information fits in current discussions about patterns of diversification within the Neotropics. Specifically, we focus on how the genetic diversity of xenarthrans has been shaped by important historical processes such as Andean uplift, the Great American Biotic Interchange, and fluctuating linkages between the Amazon and Atlantic forests. We also describe latitudinal patterns of differentiation within the Atlantic forest and discuss how these might have been generated. Even with the modest amount of information currently available, our comparative analyses indicate 3 things: the Andes may have promoted events of intraspecific divergence for at least 2 xenarthran species; the biogeographic history of the Neotropical rain forests influenced the divergence of clades in sloths; and inter- and intra-specific genetic patterns reveal a very high diversity in xenarthrans, probably higher than currently recognized from morphological data. Finally, we highlight Xenarthra as an appropriate model for investigating biogeographic patterns in the Neotropics and also point to additional directions to be taken in future studies of this unique mammal group. Revisamos los estudios realizados sobre genética de poblaciones, filogeografía y filogenia de los Xenarthra y mostramos como la información generada se ajusta a la discusión actual acerca de los patrones de diversificación en el Neotrópico. Enfocamos nuestro análisis en la influencia de 4 eventos pasados importantes relacionados con la dinámica del paisaje Neotropical y la diversificación de la fauna, sobre la diversidad genética de los Xenarthra; estos son el levantamiento de los Andes, el gran intercambio de fauna del Pleistoceno, la relación entre el bosque Amazónico y el bosque Atlántico y los patrones latitudinales de diferenciación en el bosque Atlántico. Aunque la información disponible aún es escasa, nuestro análisis comparativo indica que los Andes pudieron haber promovido eventos de diversificación intraespecífica, al menos en 2 especies de Xenarthra; que la historia biogeográfica del bosque húmedo Neotropical tuvo influencia sobre la divergencia de linajes de perezosos; y que los patrones genéticos intra e interespecíficos revelan una gran diversidad en Xenarthra, probablemente mayor que la actualmente reconocida con datos morfológicos. Finalmente, resaltamos a los Xenarthra como un modelo apropiado para investigar patrones biogeográficos en el Neotrópico y sugerimos líneas adicionales a considerarse en futuros estudios sobre este grupo único de mamíferos.
Remains of peltephilid cingulates from the late Oligocene (Deseadan, South American Land Mammal Age) of Salla, Bolivia, are described and organized as two morphs, the larger referred to a new taxon, Ronwolffia pacifica, and the smaller as indeterminate. A fairly well-preserved cranium serves as the holotype for Ronwolffia pacifica, with referred material consisting of jaws, osteoderms, and a partial pelvis. Ronwolffia is recognized by a combination of characters, some of which are regarded as general placental traits compared to some distinctive features of the well-known Santacrucian species of Peltephilus. Such generalized traits in Ronwolffia include tendencies for eight (rather than seven) mandibular teeth, unfused mandibular symphysis, incompletely ossified auditory bulla, and a low occiput and cranial vault. Like those of other peltephilids, the temporomandibular joint (TMJ) is low, but, unlike typical armadillos and the genotypic Peltephilus strepens, the glenoid fossa forms part of the wall of the external acoustic porus. Similar crowding of the TMJ and porus is noted in Peltephilus pumilus and Peltephilus ferox. Terminology related to the classification of xenarthrans is considered. Dasypodoidea Gray, 1821 is herein used for the crown clade that includes armadillos and glyptodonts, with Cingulata designating the total clade (crown + stem); that is, taxa more closely related to Dasypus than to any pilosan taxon (sloth or anteater). It is also desirable to clearly discriminate between the crown and total clade Xenarthra; thus Xenarthra is herein used exclusively for the crown, with the biogeographically inspired name, Americatheria, being proposed for the total clade; that is, taxa more closely related to Dasypus than to any members of Afrotheria or Boreotheria.