Analysis of the spatio-temporal parameters of gaits in Dasypus novemcinctus (Xenarthra: Dasypodidae)

Armadillos comprise a particular group of armoured animals whose functional morphology of locomotion remains unclear. For the first time, the kinematic patterns of Dasypus novemcinctus are analysed. Eight specimens of nine-banded armadillos were studied at a research institute in São Paulo State, Brazil. The individuals were induced to cross a horizontal corridor and each gait performed during the time each of them was kept inside this structure was recorded to a detailed analysis posteriorly performed in a computer program. Four parameters regarding speed range were considered: stride frequency (Hz) (1/stride period), stride length (m), speed (ms⁻¹) and duty factor (%). A total of 89 strides have been analysed among symmetrical (60.6%) and asymmetrical gaits (39.4%), and six footfall patterns were here reported as follows: lateral sequences (symmetrical), transverse gallop, canter, bound, half-bound and crutch walk (asymmetrical). This kind of analysis implements our knowledge on the locomotory aspects of these animals, hence contributing to the improvement of our knowledge on this still poorly known group.     

A reappraisal of the phylogeny of Mylodontidae (Mammalia,Xenarthra) and the divergence of mylodontine and lestodontine sloths

The phylogeny of mylodontid sloths has recently been the subject of multiple studies. Contrasting hypotheses have been proposed, especially for the relationships among late Miocene–Pleistocene mylodontines and lestodontines. In this paper, a new and detailed phylogenetic analysis is conducted, after adding new characters and taxa previously unexplored from a phylogenetic point of view. New features derived from postcranial skeletal anatomy are added to previous studies based on craniodental evidence. In this way, the current reappraisal represents the first exhaustive phylogenetic study on the Mylodontidae that incorporates features coded for the entire skeleton. When available, multiple specimens of each bony element are observed for each operational taxonomic unit, in order to take into account intraspecific variation. The taxonomic sample of this study considers Mylodontinae at the specific level. However, many other Mylodontidae are considered, and their phylogenetic relationships tested. The taxonomic sample of this study is enriched with new taxa from central and northern South America, with the aim of compensating for the knowledge bias in favour of austral mylodontids, which have historically been more extensively studied than those from tropical latitudes. Special emphasis is given to the phylogenetic relationships of Mylodontinae, and particularly to the mylodontine and lestodontine sloths, that are recovered in the present study as monophyletic clades, and together form a larger monophyletic group. According to the present results, the Mylodontini–Lestodontini split occurred at the middle–late Miocene transition, giving rise to independent adaptive radiations across South and North America.     

Morphology of female genital tracts in Dasypodidae (Xenarthra,Mammalia): a comparative survey

Previous works about comparative spermatology in Dasypodidae determined that sperm morphology is a striking variable among genera. It was suggested that this sperm feature may be related to specific morphologies of the female reproductive tract. The present comparative study of the morphology of the female genital tract from seven species corresponding to six genera of Dasypodidae is aimed to determine the main similarities and differences between the species and to establish a possible correlation with the sperm shapes and sizes. Genital tracts were studied macroscopically and histologically. Dasypus hybridus has disk-shaped ovaries and the cortex occupies almost all the organ with a single oocyte in each follicle. Tolypeutes matacus, Chaetophractus villosus, Chaetophractus vellerosus, Zaedyus pichiy, Cabassous chacoensis and Clamyphorus truncatus possess ovoid and elongated ovaries, with both longitudinally polarized cortex and medulla, and the peculiar presence of several oocytes in the same follicle. D. hybridus and T. matacus have a simple pear-shaped uterus, but in the other species the uterus is pyramid shaped and bicornuate. The uterine cervix is very long in all studied species. Only T. matacus presents a true vagina as in most eutherian mammals; on the other hand, in the other species a urogenital sinus is observed. The structure of female reproductive tracts in Dasypodidae contains a mixture of assumedly primary and other derived features. According to the different morphologies of the regions analyzed, a classification of the female genital tracts in three groups can be performed (group 1: Dasypus; group 2: Tolypeutes; group 3: Chaetophractus, Zaedyus, Cabassous, Clamyphorus) and a correlation between each group and a specific sperm morphology can be established.     

Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence

This study is undertaken in order to evaluate specific hypotheses of relationship among extant and extinct sloths (Mammalia, Xenarthra, Tardigrada). Questions of particular interest include the relationship among the three traditional family groupings of extinct ground sloths and the monophyletic or diphyletic origin of the two genera of extant tree sloths. A computer‐based cladistic investigation of the phylogenetic relationships among 33 sloth genera is performed based upon 286 osteological characteristics of the skull, lower jaw, dentition and hyoid arch. Characters are polarized via comparisons with the following successive outgroups, all members of the supraordinal grouping Edentata: the Vermilingua, or anteaters; the Cingulata, or armadillos and glyptodonts; the Palaeanodonta; and the Pholidota, or pangolins. The results of the analysis strongly corroborate the diphyly of living tree sloths, with the three‐toed sloth Bradypus positioned as the sister‐taxon to all other sloths, and the two‐toed sloth Choloepus allied with extinct members of the family Megalonychidae. These results imply that the split between the two extant sloth genera is ancient, dating back perhaps as much as 40 Myr, and that the similarities between the two taxa, including their suspensory locomotor habits, present one of the most dramatic examples of convergent evolution known among mammals. The monophyly of the three traditional ground sloth families Megatheriidae, Megalonychidae and Mylodontidae is confirmed in the present study, and the late Miocene–Pleistocene nothrotheres are shown to form a clade. It is suggested that this latter clade merits recognition as a distinct family‐level grouping, the family Nothrotheriidae. The monophyly of the Megatherioidea, a clade including members of the families Megatheriidae, Megalonychidae and Nothrotheriidae, is also supported. Within Megatherioidea, the families Nothrotheriidae and Megatheriidae form a monophyletic group called the Megatheria. The relationships within the families Megatheriidae and Mylodontidae are fully and consistently resolved, although the hypothesized scheme of relationships among the late Miocene to Pleistocene members of the mylodontid subfamily Mylodontinae differ strongly from any proposed by previous authors. Within the family Megalonychidae, Choloepus is allied to a monophyletic grouping of West Indian sloths, although the relationships within this clade are not fully resolved. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 255–305.     

Feeding mechanics and dietary implications in the fossil sloth Neocnus (Mammalia: Xenarthra: Megalonychidae) from haiti

Haitian species of the extinct ground sloth genus Neocnus (Mammalia: Pilosa: Megalonychidae) have previously been hypothesized to have a much reduced jugal bone and a correspondingly reduced masseter musculature but a paucity of specimens has prevented further investigation of this hypothesis. Recent discovery of jugal bones belonging to Haitian specimens of Neocnus within the University of Florida Museum collections enables the element to be more accurately described. The discovery also makes it possible to explore mastication in these sloths. Osteological characters related to feeding were examined, along with comparative estimations of bite force with the extant tree sloths, Bradypus and Choloepus, and their known dietary habits as a means to infer aspects of the paleodiet of Neocnus. There is a significant difference in moment arm calculations for m. masseter between predicted and actual jugals, but the overall significance for bite force is lost and hampered by small sample size. Neocnus demonstrates a variety of characters that are similar to those of Bradypus and not to Choloepus, which is a close phylogenetic relative. The masticatory musculature of Neocnus enabled a chewing cycle emphasizing a grinding combination of mesiodistal and linguobuccal movements of the molariform dentition. The orientations of m. masseter and m. temporalis are estimated to produce relatively high bite force ratios that imply a masticatory system with stronger versus faster components. Because of the similarity of bite forces and jaw mechanics to those of Bradypus, in addition to a number of osteological adaptations indicative of herbivorous grazers (elevated mandibular condyle, large and complex masseter, and robust angular process), the Haitian forms of Neocnus are considered to have been selective feeders with a folivorous diet. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

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. J. Morphol. 271:1119–1133, 2010. © 2010 Wiley‐Liss, Inc.     

Skull Shape, Masticatory Apparatus, and Diet of Vassallia and Holmesina (Mammalia: Xenarthra: Pampatheriidae): When Anatomy Constrains Destiny

The form and function of the masticatory apparatus of the fossil genera Vassallia and Holmesina are analyzed so that the possible dietary behaviors of these pampathere xenarthrans might be inferred. Analysis is based on comparisons of dental morphology and skeletal features (through RFTRA) associated with the masticatory musculature among the pampatheres, the extant dasypodids Euphractus and Dasypus, and the glyptodont Propalaeohoplophorus. A method is proposed for generating a moment arm of the massetericus independently of the muscle's line of action, which allows direct comparison among extant and fossil mammals. The masticatory apparatus of the pampatheres strongly resembles that of Euphractus among extant forms, but the development of muscular attachment sites indicates a more powerful musculature, particularly the massetericus; the taxa differ most markedly in dental morphology. Long moment arms about the jaw joint and large ratios of muscle to bite moments indicate forceful rather than quick movements. The various skeletal and dental features analyzed suggest that the masticatory apparatus of the pampatheres was more powerful and efficient in transverse chewing than in dasypodids and that they were primarily grazers consuming mainly coarse vegetation. These features, some shared with herbivorous ungulates, include wide, relatively flat mandibular condyles; condyles well dorsal to muscular insertion sites; expanded angular processes; unfused symphysis; a posteriorly extended tooth row; open-rooted teeth; mesial teeth that bear mainly transverse striations; distal teeth that are mesiodistally elongated, bear basined occlusal surfaces, and in Vassallia possess a central island of resistant dentine that acted as a functional analogue of an ectoloph; and teeth with a stepwise arrangement. The results of this study indicate that detailed analysis and comparison of morphology lead to useful predictions of behavior.     

The Phylogeny of the Myrmecophagidae (Mammalia, Xenarthra, Vermilingua) and the Relationship of Eurotamandua to the Vermilingua

A cladistic investigation of the phylogenetic relationships among the three extant anteater genera and the three undoubted extinct myrmecophagid genera is performed based upon osteological characteristics of the skull and postcranial skeleton. One hundred seven discrete morphological characters are analyzed using the computer program PAUP. Characters are polarized via comparison to the successive xenarthran outgroups Tardigrada (represented by the living sloth Bradypus) and Cingulata (represented by the recent armadillos Dasypus and Euphractus). The analysis results in a single most-parsimonious tree (TL = 190, CI = 0.699, RI = 0.713). The tree corroborates the monophyly of the subfamilies Cyclopinae and Myrmecophaginae, the former including the extant Cyclopes and the Pliocene genus Palaeomyrmidon. Within the Myrmecophaginae the Miocene genus Protamandua is the sister taxon to a clade including the remaining three genera. The recent Tamandua is in turn the sister taxon to the extant Myrmecophaga plus the Pliocene genus Neotamandua. Contrary to the suggestions of recent authors, weak support is provided for the taxonomic distinctiveness of the latter genus from the recent Myrmecophaga. The monophyly of the Myrmecophagidae is supported by 15 unequivocal synapomorphies. The monophyly of the Cyclopinae and Myrmecophaginae is supported by 3 and 13 unambiguous synapomorphies, respectively. The enigmatic Eocene genus Eurotamandua, from the Messel fauna of Germany, is coded for the 107 morphological characters above and included in two subsequent PAUP analyses. The palaeanodont Metacheiromys is also added to these two analyses as a nonxenarthran outgroup to test for the possibility that Eurotamandua lies outside the Xenarthra. In the first analysis, Eurotamandua is constrained a priori to membership in the Vermilingua. The single most-parsimonious tree (TL = 224, CI = 0.618) that results places Eurotamandua as the sister group to the remaining anteater genera, contra Storch and Habersetzer's (1991) assignment of Eurotamandua to the vermilinguan subfamily Myrmecophaginae. Eurotamandua shares six unequivocal synapomorphies with other anteaters, including the absence of teeth and the presence of a lateral tuberosity on the fifth metatarsal. The remaining vermilinguans are united by 11 unequivocal synapomorphies, plus an additional 10 ambiguous synapomorphies. In the second analysis, the position of Eurotamandua is unconstrained. The resulting single most-parsimonious tree (TL = 219, CI = 0.632) places Eurotamandua outside Vermilingua as the sister group to the Pilosa (Vermilingua plus Bradypus). The monophyly of this node is supported by four unambiguous synapomorphies in the unconstrained analysis. Further manipulation of this second analysis shows that placement of Eurotamandua as the sister group to the Xenarthra or to the Palaeanodonta adds three steps to the shortest tree but is more parsimonious than its placement as a sister group to the Vermilingua is the previous analysis. The addition of pangolins to the analysis does little to alter the major phylogenetic conclusions of the study. The allocation of Eurotamandua to the Xenarthra, but as a sister group to the Pilosa, is a novel arrangement which leaves open the biogeographic question of how a xenarthran reached Western Europe during the Eocene.     

Dental microwear in the orthodentine of the Xenarthra (Mammalia) and its use in reconstructing the palaeodiet of extinct taxa: the case study of Nothrotheriops shastensis (Xenarthra,Tardigrada, Nothrotheriidae)

The utility of orthodentine microwear analysis as a proxy for dietary reconstruction in xenarthrans (tree sloths, armadillos) was quantitatively and statistically accessed via low‐magnification stereomicroscopy. Features such as number of scratches and pits, as well as presence of gouges, hypercoarse scratches, > four large pits, > four cross scratches, and fine, mixed or coarse scratch texture were recorded in 255 teeth from 20 extant xenarthran species. Feature patterns are consistent with scar formation through abrasional (tooth–food) and attritional (tooth–tooth) contact. Number of scratches is the most dietary diagnostic microwear variable for xenarthrans, with herbivorous sloths characterized by > ten scratches and nonherbivorous armadillos by < ten scratches. Discriminant function analysis differentiated arboreal folivores (sloths) and frugivore‐folivores (sloths) both from each other and from fossorial carnivore‐omnivores (armadillos) and insectivores (armadillos). Microwear patterns in carnivore‐omnivores and insectivores are difficult to distinguish between; armadillo microwear may reflect a fossorial lifestyle (grit consumption) rather than primary diet. Cabassous centralis is anomalous in its microwear signal relative to all other insectivores. To test the utility of orthodentine microwear analysis as an indicator of palaeodiet in extinct xenarthrans, microwear in the ground sloth Nothrotheriops shastensis was quantitatively and statistically compared to microwear in extant taxa. Microwear patterns in N. shastensis are most comparable to extant folivores based on scratch number and hierarchical cluster analysis. This strongly supports an herbivorous diet for N. shastensis that is corroborated by multiple independent lines of evidence. Thus, orthodentine microwear analysis can be used to reconstruct diet in extinct xenarthrans. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 201–222.     

Hibernation and daily torpor in an armadillo, the pichi (Zaedyus pichiy)

Hibernation and daily torpor are physiological strategies to cope with energetic challenges that occur in many mammalian and avian taxa, but no reliable information exists about daily torpor or hibernation for any xenarthran. Our objective was to determine whether the pichi (Zaedyus pichiy), a small armadillo (Xenarthra, Dasypodidae) that inhabits arid and semi-arid habitats in central and southern Argentina and Chile, enters shallow daily torpor or prolonged deep hibernation during winter when environmental temperature and food availability are low. We studied body temperature changes during winter in semi-captive pichis by means of temperature dataloggers implanted subcutaneously. All individuals entered hibernation, characterized by torpor events of 75 ± 20 h during which the subcutaneous temperature (T_sc) decreased to 14.6 ± 2.1 °C. These events were interrupted by periods of euthermia of 44 ± 38 h with a T_sc of 29.1 ± 0.7 °C. After the hibernation season, daily torpor bouts of 4 to 6 h occurred irregularly, with T_sc dropping to as low as 24.5 °C. We conclude that the pichi is a true hibernator and can enter daily torpor outside of the hibernation season.     

Husbandry of a pink fairy armadillo (Chlamyphorus truncatus): case study of a cryptic and little known species in captivity

Pink fairy armadillos, Chlamyphorus truncatus, are poorly known fossorial mammals that are endemic to central Argentina. These smallest of all extant armadillos are rarely observed in the field and extremely difficult to maintain under captive conditions. This case study describes the husbandry of a male pink fairy armadillo that was maintained in an artificial environment for 8 months. A stable, undisturbed environment consisting of abundant compact sand and hiding places on the surface was the key to its successful maintenance. The artificial diet consisted of a semiliquid mixture of ground cat food, insectivore diet, mashed banana, vitamins, and minerals. Any slight modification to its environment or diet triggered a stress response. Zoo Biol 30:225–231, 2011. © 2010 Wiley‐Liss, Inc.     

Morphological constraints in the digging apparatus of pocket gophers (Mammalia: Geomyidae)

The digging apparatus of pocket gophers offers a unique opportunity to examine morphological constraints within a historical context because relationships among extant taxa are well resolved and the features enhancing digging performance are relatively well understood. Structural and functional considerations suggest that the muscles associated with tooth- and claw-digging in pocket gophers are subjected to contrasting levels of morphological constraints. To assess this hypothesis, we analysed the bones and muscles of the jaws and forelimbs in four genera comprising five species of pocket gophers. Morphometric analyses were performed on 12 osteological measurements selected to reflect overall skull size, variation in rostral shape and procumbency, differences in overall length of the forelimbs and processes relating to the function of lever systems used in claw-digging. In addition, dissections were made of the jaw, hyoid, neck and all of the forelimb muscles excluding the intrinsic muscles of the manus. Results of our morphometric analyses corroborate the recent suggestion that pocket gophers encompass a wide range of morphological variation extending from claw-diggers to tooth-diggers. Myologically, however, we found structural variation only in the forelimb muscles, some of which may be advantageous for digging. No changes in jaw, neck and hyoid muscles, other than differences in muscle mass or those concordant with differences in rostral shape, were noted. These results support our hypothesis that constrasting levels of morphological constraint exist between the jaw and forelimb muscles of pocket gophers. We present a discussion of the structural and functional constraints on jaws and forelimbs in gophers as well as an analysis of historical constraints acting on this group, and perhaps on mammals in general.     

Reconstruction of the cranial musculature and masticatory function of the Pleistocene panamerican ground sloth Eremotherium laurillardi (Mammalia,Xenarthra, Megatheriidae)

Cranial musculature, dental function and mandibular movement patterns in Eremotherium laurillardi were reconstructed from the examination of crania and dentitions. Size, shape and pattern of muscle divisions were reconstructed from the examination of bony rugosities indicating muscle attachments. Details of masticatory muscle structure and function were based on dissections of the tree sloths Bradypus and Choloepus. Among sloths, masticatory muscles in E. laurillardi demonstrate a different synergist–antagonist pattern, reflecting greater emphasis on mediolateral mandibular movements. Eight cranial character complexes (anterior facial, zygomatic arch, superficial masseter, deep masseter–zygomaticomandibularis, pterygoid, temporal, occipital and occlusal) determined by interrelated contributions of each component made to group functions were identified. An elongate anterior face and predental spout in E. laurillardi allowed protrusion of a long narrow tongue at small degrees of gape, reflecting a probably ancestral xenarthran condition. Gape minimisation, in conjunction with the mediolaterally directed masticatory stroke in E. laurillardi, was a unique solution to increase masticatory efficiency by permitting molariform tooth shearing surfaces to remain in or near occlusion for a greater percentage of each chewing cycle.     

Myology of the Feeding Apparatus of Myrmecophagid Anteaters (Xenarthra: Myrmecophagidae)

The musculoskeletal feeding apparatus of anteaters in the family Myrmecophagidae (Eutheria: Xenarthra) is described, compared among the three extant genera (Tamandua, Myrmecophaga, Cyclopes), and interpreted in a phylogenetic framework. Character polarities are assessed with reference to other xenarthrans, eutherians, and didelphid marsupials. Xenarthrans are widely regarded as basal eutherians, and this is reflected in the apparent retention of plesiomorphic character states in jaw and pharyngeal musculature. Jaw closing muscles are architecturally simple, the stylohyoideus is absent, the stylopharyngeus is robust and architecturally complex, and the superior pharyngeal constrictor is weak. At the same time, the highly specialized trophic ecology of myrmecophagids is reflected in derived features of the jaw, tongue, and palatal musculature. The sternomandibularis is present, the tongue is largely composed of a sternog-lossus with no attachments to the hyoid apparatus, other glossus muscles are modified and do not enter the tongue, and the mylohyoideus and stylopharyngeus contribute to the soft palate, while other palatal muscles vary among the myrmecophagid genera. Feeding apparatus mycology provides further support for myrmecophagid monophyly. Documentation of the morphological transformations that lead to the myrmecophagid condition is hampered by incomplete data on feeding apparatus structure in nonmyrmecophagid xenarthrans (sloths and armadillos) but a tentative character mapping onto an independently derived phylogeny is offered.     

Ungual phalanges analysis in Pleistocene ground sloths (Xenarthra,Folivora)

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.     

New ontogenetic evidence on the septomaxilla ofTamandua andCholoepus (Mammalia,Xenarthra), with a reevaluation of the homology of the mammalian septomaxilla

An intramembranous ossification at the anterior end of the cartilaginous nasal capsule is described for the first time in prenatal specimens of the anteaterTamandua and the slothCholoepus and redescribed in prenatal specimens of the armadillosDasypus andZaedyus. From comparisons of this bone with the septomaxilla of monotremes and various Mesozoic mammals, it is concluded that (1) the bone inTamandua andCholoepus is homologous with the central part (processus ascendens) of the bone inDasypus, Zaedyus, and other armadillos and (2) the xenarthran processus ascendens, in turn, is homologous with the central part of the septomaxilla of monotremes and various Mesozoic mammals. Therefore, the bone in question in xenarthrans is a true septomaxilla. It is further concluded that the armadillo septomaxilla has two neomorphic components: a lamina palatina beneath the cartilaginous nasal floor and a processus intrafenestralis extending rostrally into the nasal fossa.     

A new species of Megalonychidae (Mammalia, Xenarthra) from the Quaternary of Poço Azul (Bahia, Brazil)

The new genus and species Ahytherium aureum (Mammalia, Xenarthra, Megalonychidae) from the Quaternary of Poço Azul (Bahia, Brazil) is described. It is the first Brazilian megalonychid known from reasonably complete and well-preserved remains. Purported Brazilian megalonychids described in the past, such as Ocnopus gracilis and Xenocnus cearesis, are noted as belonging to other sloth clades, and the acceptance by past paleontologists of the existence of ‘strange’ megalonychids in Brazil is shown to be erroneous. Ahytherium aureum, in fact, exhibits typical megalonychid morphology. It differs from other known members of Megalonychidae in several characters, including a markedly shortened, but high rostral region, with dorsally inflated frontals, wide zygomatic processes of the frontal, narrow, blade-like and anterolaterally oriented lacrimals, curved, slender and oval caniniforms, gracile humerus with less developed deltopectoral shelf, and relatively distal position of the greater trochanter of the femur. A second specimen from São Paolo state is tentatively assigned to the new genus and species.     

First Neogene skulls of Doedicurinae (Xenarthra,Glyptodontidae): morphology and phylogenetic implications

Among Glyptodontidae, Doedicurinae (late Miocene–early Holocene) includes the glyptodonts with the largest size and latest records. Doedicurinae is mainly characterised by a smooth surface of the osteoderms with large foramina, and a particular morphology of the caudal tube. All taxa except one (Doedicurus clavicaudatus) have been recognised and characterised on the basis of remains of caudal tubes and/or dorsal carapaces. This situation produced an evident overestimation of the real diversity of this group, and a taxonomic revision is needed. In fact, no Neogene skulls were known. We present and describe the first two Neogene skulls belonging to Doedicurinae (cf. Eleutherocercus antiquus). The materials come from the El Polvorín and Chapadmalal Formations, in the surroundings of Olavarría and Mar del Plata localities, respectively (Buenos Aires province, Argentina). A cladistic analysis was carried out in order to situate these materials among Glyptodontidae and inferring new synapomorphies at skull level in Doedicurinae. Cf. Eleutherocercus antiquus clusters with the Pleistocene species Doedicurus clavicaudatus showing three unambiguous synapomorphies, which in turn represents the first skull synapomorphies for Doedicurinae. Finally, the presence of cf. Eleutherocercus antiquus in the El Polvorín and Chapadmalal Formations suggests that the stratigraphic distribution of this species could include the Montehermosan–Chapadmalalan interval.     

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.