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.
Rod monochromacy is a rare condition in vertebrates characterized by the absence of cone photoreceptor cells. The resulting phenotype is colourblindness and low acuity vision in dim-light and blindness in bright-light conditions. Early reports of xenarthrans (armadillos, sloths and anteaters) suggest that they are rod monochromats, but this has not been tested with genomic data. We searched the genomes of Dasypus novemcinctus (nine-banded armadillo), Choloepus hoffmanni (Hoffmann's two-toed sloth) and Mylodon darwinii (extinct ground sloth) for retinal photoreceptor genes and examined them for inactivating mutations. We performed PCR and Sanger sequencing on cone phototransduction genes of 10 additional xenarthrans to test for shared inactivating mutations and estimated the timing of inactivation for photoreceptor pseudogenes. We concluded that a stem xenarthran became an long-wavelength sensitive-cone monochromat following a missense mutation at a critical residue in SWS1, and a stem cingulate (armadillos, glyptodonts and pampatheres) and stem pilosan (sloths and anteaters) independently acquired rod monochromacy early in their evolutionary history following the inactivation of LWS and PDE6C, respectively. We hypothesize that rod monochromacy in armadillos and pilosans evolved as an adaptation to a subterranean habitat in the early history of Xenarthra. The presence of rod monochromacy has major implications for understanding xenarthran behavioural ecology and evolution.
Northwestern Argentina contains one of the most complete continental late Neogene (ca. 9–2.58 Ma) fossiliferous sequences in South America, especially in the current territories of the Catamarca, Tucumán and Jujuy provinces. More precisely in Jujuy Province several localities bearing mainly fossil mammals have been reported at the Quebrada de Humahuaca in the Uquía, Maimará and Tilcara formations, in which the clade Xenarthra (Mammalia) is well-represented. In this scenario, the fossiliferous potential of other localities of Jujuy Province are less known, especially in those areas located at the northwest end of Argentina, bordering Bolivia in the Northern Puna. A new late Neogene fossiliferous locality near Calahoyo (3639 m.a.s.l), Jujuy Province, is here reported. The materials, belonging to Xenarthra, were exhumed from the base of the Tafna Formation which was deposited in a sedimentary basin by alluvial and/or fluvial currents, undergoing transitions of various lacustrine episodes. The taxa include the Tardigrada (Megatheriidae) and the Cingulata sp. (Glyptodontidae) and (Dasypodidae). From a biostratigraphic viewpoint, this assemblage suggests a Late Miocene-Pliocene age for the base of the Tafna Formation, and partially contradicts the supposed Plio-Pleistocene age of this unit. Finally, the new specimens here described indicate that Xenarthra were taxonomically and ecologically diverse during the late Neogene in the northwest end of Argentina, since they are represented by at least three main lineages (sloths, glyptodontids and armadillos).
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.
Riostegotherium yanei from the Itaborai Basin, Brazil, is the oldest known Xenarthra. This paper aims to describe the internal morphology of the osteoderms of Riostegotherium yanei from the perspective of histology and micro-CT approaches, expanding the available data on cingulate osteoderm microstructure. Seven osteoderms of R. yanei were used for the internal microstructure description and eight of Dasypus novemcinctus for comparison. The osteoderms of Riostegotherium yanei lacks the diploe-like structure typical of glyptodonts but has a three-layered structure composed of two layers of non-Haversian compact bone enclosing a central layer of primary and secondary osteons. This internal organization is distinct from other Astegotheriini of comparable age, but similar to Dasypus. The 3D reconstruction of Riostegotherium yanei revealed two patterns of internal organization. Pattern 1 of movable osteoderm is composed of large remodeled areas at the base and a more compact bone at the tongue; in Pattern 2 (both movable and buckler), the internal cavities are much smaller, more numerous, and more interconnected to each other. In one buckler osteoderm, the cavities are organized somewhat radially with a compact central region (Pattern 1). Pattern 1 of both movable and buckler osteodenns resemble that of Dasypus.
In Argentina, Chaetophractus villosus has a wide distribution that overlaps with agricultural areas where soybean is the predominant crop. In such areas the pesticide Roundup Full II (R) (RU) is widely applied. The genotoxic effect of its active ingredient glyphosate (RU is 66.2% glyphosate) on the peripheral blood lymphocytes of C. villosus was tested over a range of concentrations (280, 420, 560, 1120 mu mol/L). Culture medium without glyphosate served as negative control, while medium containing mitomycin C served as positive control. Genetic damage was characterized in terms of the percentage of cells with chromosome aberrations (CA), the mean number of sister chromatid exchanges (SCE) per cell, and the modification of cell proliferation kinetics via the calculation of the replication index. Significant increases (p < 0.0001) were seen in the CA frequency and the mean number of SCEs per cell compared to negative controls at all the RU concentrations tested. Chromatid breaks, the only form of CA observed, under the 560 mu mol/L RU conditions and in presence of mitomycin C were four to five times more common than at lower concentrations, while no viable cells were seen in the 1120 mu mol/L treatment. The mean number of SCEs per cell was significantly higher under the 280 mu mol/L RU conditions than the 420 or 560 mu mol/L RU conditions; cells cultivated in the presence of MMC also showed significantly more SCEs. All the RU concentrations tested (except in the 1120 mu mol/L RU treatment [no viable cells]) induced a significant reduction in the replication index (p < 0.0001). The present results confirm the genotoxic effects of RU on C. villosus lymphocytes in vitro, strongly suggesting that exposure to RU could induce DNA damage in C. villosus wildlife.
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.
The phylogenetic positions of the 4 clades, Euarchontoglires, Laurasiatheria, Afrotheria, and Xenarthra, have been major issues in the recent discussion of basal relationships among placental mammals. However, despite considerable efforts these relationships, crucial to the understanding of eutherian evolution and biogeography, have remained essentially unresolved. Euarchontoglires and Laurasiatheria are generally joined into a common clade (Boreoeutheria), whereas the position of Afrotheria and Xenarthra relative to Boreoeutheria has been equivocal in spite of the use of comprehensive amounts of nuclear encoded sequences or the application of genome-level characters such as retroposons. The probable reason for this uncertainty is that the divergences took place long time ago and within a narrow temporal window, leaving only short common branches. With the aim of further examining basal eutherian relationships, we have collected conserved protein-coding sequences from 11 placental mammals, a marsupial and a bird, whose nuclear genomes have been largely sequenced. The length of the alignment of homologous sequences representing each individual species is 2,168,859 nt. This number of sites, representing 2840 protein-coding genes, exceeds by a considerable margin that of any previous study. The phylogenetic analysis joined Xenarthra and Afrotheria on a common branch, Atlantogenata. This topology was found to fit the data significantly better than the alternative trees.