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.
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.
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.
Ungual phalanges (the most distal bone within a limb) and claws (the overlying corneous sheath) from the third digit of the forefoot of selected Pleistocene ground sloths (Lestodon armatus, Glossotherium robustum, Scelidotherium leptocephalum and Megatherium americanum) are analysed, as well as those of some living xenarthrans for actualistic comparison, aiming at testing hypotheses of substrate usage and locomotor behaviour. The third digits were chosen for this study because of its size and nearly perfect bilateral symmetry, which increases the possibilities of revealing functional differences between taxa. The analyses performed were of inner and external curvature, the strength indicator and the mechanical advantage. The mechanical advantage indicates that the four ground sloths' species were well adapted for strenuous activities, such as digging, in which force rather than velocity is optimised. Their strength indicator shows expected values for their body size, while in Mylodon darwinii the value obtained was lower than expected. In the two curvature analyses L. armatus, G. robustum and M. americanum fall within the group of armadillos that dig, whereas S. leptocephalum does not, this might be due to a difference in the movements performed while performing an activity such as digging or similar to it.
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
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.