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 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.
Recently, dental microwear analysis has been successfully employed to xenarthran teeth. Here, we present new data on use wear features on 16 molariforms of Orophodon hapaloides and Octodontotherium grande. These taxa count among the earliest sloths and are known from the Deseadan SALMA (late Oligocene). Modern phylogenetic analyses classify Octodontotherium and Orophodon within Mylodontoidea with whom they share lobate cheek teeth with an outer layer of cementum and a thick layer of orthodentine. Similar target areas of 100μm2 were analyzed on the orthodentine surface of each tooth by stereomicroscopic microwear and by SEM microwear. Results were unlike those of extant sloths (stereomicroscopic microwear: Bradypus, Choloepus) and published data from fossil sloths (SEM microwear: Acratocnus, Megalonyx, Megatherium, Thinobadistes); thus, both approaches independently indicate a different feeding ecology for the Oligocene taxa. The unique microwear results suggest that both taxa fed on plant material with low to moderate intrinsic toughness (foliage, twigs) but also proposes intake of tougher food items (e.g., seeds). Frequent gouging of the tooth surfaces can be explained by exogenous influence on microwear, such as possible intake of abrasive grit. We suggest an unspecialized herbivorous diet for Octodontotherium and Orophodon utilizing diverse food resources of their habitat. These interpretations support the reconstruction of (1) Deseadan environments as open habitats with spreading savannas/grasslands and (2) both taxa as wide-muzzled bulk feeders at ground level.
To investigate the evolution of xenarthran epaxial muscles, fresh specimens of the North American Common long-nosed armadillo Dasypus novemcinctus and of a marsupial, the Virginia opossum Didelphis virginiana, were dissected. Data from one fixed specimen of a two-toed sloth Choloepus didactylus were also used for comparison, because it is a xenarthran exhibiting a highly derived locomotor mode. The opossum was used to represent a more generalized mammalian condition. Each of the three mammalian epaxial muscle groups, the iliocostalis, longissimus dorsi, and transversospinalis, was removed and its mass was determined. All data were corrected for body mass and length. Unpaired, one-tailed t-tests showed the average mass of the iliocostalis and transversospinalis of Dasypus to be significantly larger than the mass of the same muscles in Didelphis, whereas the average mass of the longissimus dorsi was not statistically different between the two species. In agreement with pronounced lateral bending and de-emphasized dorso-ventral flexion and extension, Choloepus also had a relatively large iliocostalis and small longissimus. Our limited data suggest that this condition was inherited from non-arboreal and probably digging early xenarthrans. We believe the relatively larger iliocostalis and transversospinalis muscles in Dasypus can be attributed to the need to provide vertical stabilization of the trunk and resist lateral reaction forces generated by digging. Thus, for Xenarthra it represents a synapomorphy linked to adaptations for fossoriality.
Numerous climatic fluctuations occurred during the Cenozoic (last 66 Ma BP); some of them were drastic (e.g., during the Eocene-Oligocene boundary) while others were more gradual (e.g., late Tertiary cooling), but both have deep effect on the biotas. Armadillos are exclusively from the Americas; they have an old evolutionary history in South America and faunal replacement and/or local extinctions were detected, linked with climatic fluctuations. The global cooling of the late Eocene - early Oligocene coincides with a well-documented faunal turnover of Dasypodinae by Euphractinae in Patagonia. During cold and arid periods of the Quaternary, Euphractinae and Tolypeutinae moved more than once to the eastern Pampean Region, and Dasypodinae moved northward to central Brazil or even further north to the Guyana Region. During interglacial periods some armadillos went extinct locally and/or moved to Patagonia (Zaedyus), central Argentina (Tolypeutes matacus, Chaetophractus vellerosus), or from the north to Mesopotamia and the Pampean Region (Dasypus). Since the end of the Pleistocene/early Holocene, human activity has strongly impacted armadillo populations. Currently, the eastern Pampean Region (Argentina) is characterized by the presence of the couple C. villosus - D. hybridus (probably established since the late Holocene), but during the Pleistocene was Z. pichiy – T. matacus while Z. pichiy - C. villosus characterized early-middle Holocene. This work serves as evidence that paleozoological studies can be used to assess responses of biological systems to large scale perturbations and is the basis for studying future species distributions, in order to identify species in danger of extinction and establish management actions.
Background: Extant sloths present an evolutionary conundrum in that the two living genera are superficially similar (small-bodied, folivorous, arboreal) but diverged from one another approximately 30 million years ago and are phylogenetically separated by a radiation of medium to massive, mainly ground-dwelling, taxa. Indeed, the species in the two living genera are among the smallest, and perhaps most unusual, of the 50+ known sloth species, and must have independently and convergently evolved small size and arboreality. In order to accurately reconstruct sloth evolution, it is critical to incorporate their extinct diversity in analyses. Here, we used a dataset of 57 species of living and fossil sloths to examine changes in body mass mean and variance through their evolution, employing a general time-variable model that allows for analysis of evolutionary trends in continuous characters within clades lacking fully-resolved phylogenies, such as sloths. Results: Our analyses supported eight models, all of which partition sloths into multiple subgroups, suggesting distinct modes of body size evolution among the major sloth lineages. Model-averaged parameter values supported trended walks in most clades, with estimated rates of body mass change ranging as high as 126 kg/million years for the giant ground sloth clades Megatheriidae and Nothrotheriidae. Inclusion of living sloth species in the analyses weakened reconstructed rates for their respective groups, with estimated rates for Megalonychidae (large to giant ground sloths and the extant two-toed sloth) were four times higher when the extant genus Choloepus was excluded. Conclusions: Analyses based on extant taxa alone have the potential to oversimplify or misidentify macroevolutionary patterns. This study demonstrates the impact that integration of data from the fossil record can have on reconstructions of character evolution and establishes that body size evolution in sloths was complex, but dominated by trended walks towards the enormous sizes exhibited in some recently extinct forms.
Most of the mammalian diversity is known only from fossils, and only a few of these fossils are well preserved or abundant. This undersampling poses serious problems for understanding mammalian phenotypic evolution under a quantitative genetics framework, since this framework requires estimation of a group's additive genetic variance-covariance matrix (G matrix), which is impossible, and estimating a phenotypic variance-covariance matrix (P matrix) requires larger sample sizes than what is often available for extinct species. One alternative is to use Gor P matrices from extant taxa as surrogates for the extinct ones. Although there are reasons to believe this approach is usually safe, it has not been fully explored. By thoroughly determining the extant and some extinct Xenarthra (Mammalia) cranium P matrices, this study aims to explore the feasibility of using extant G or P matrices as surrogates for the extinct ones and to provide guidelines regarding the reliability of this strategy and the necessary sample sizes. Variance-covariance and correlation P matrices for 35 cranium traits from 16 xenarthran genera (12 extant and 4 extinct) were estimated and compared between genera. Results show xenarthran P-matrix structures are usually very similar if sample sizes are reasonable. This study and others developed with extant therian mammals suggest, in general, that using extant G or P matrices as an approximation to extinct ones is a valid approach. Nevertheless, the accuracy of this approach depends on sample size, selected traits, and the type of matrix being considered.
The presence of osteoderms in the skin of some extinct sloths and in cingulates (armadillos, pampatheres, and glyptodonts) has often been considered a pleisomorphic character of the Xenarthra. While osteoderms are known from the earliest cingulates, they are absent in most sloths including the two extant taxa and only appear late in their fossil record. Osteoderms are currently only reported from five genera of mylodonts and two megatheres, out of the over 100 currently recognized genera of sloths. Consequently, rather than a plesiomorphic character of the Xenarthra, which has been secondarily lost in sloths, it is more likely that osteoderms in sloths are the result of parallel evolution to the cingulates that independently evolved in one, possibly two different sloth clades.
Here we describe two new megalonychid sloths from the late Miocene of the Urumaco Formation (Falcón State, Venezuela), Urumacocnus urbanii gen. et. sp. nov. and Pattersonocnus diazgameroi gen. et sp. nov. The recovery of these distinct taxa greatly improves our understanding of sloth diversity in the late Miocene of northern South America. A phylogenetic analysis based on the combination of cranial and postcranial elements (particularly the femur) partially supports previous interpretations of the relationships of genera within the Megalonychidae, but also suggests the existence of a more complex set of subclades within the family in South America, North America and the Antilles. http://zoobank.org/urn:lsid:zoobank.org:pub:594532F8-9E25-4282-B1AD-9FC5484DFD91