The Convergent Evolution of Suspensory Posture and Locomotion in Tree Sloths

Recent phylogenetic analyses imply a distant relationship and long separated evolution of two-toed sloths (Choloepus) and three-toed sloths (Bradypus). No known fossil sloth is interpreted to have been suspensory. As a consequence, the suspensory posture and locomotion of the extant genera likely evolved convergently in both lineages, forming a new framework for the analysis of functional aspects of the locomotor apparatus of extant tree sloths. The suspensory posture and locomotion has altered functional demands from the phylogenetically plesiomorphic non-suspensory pronograde situation. Here, anatomical traits that have been argued to be of adaptive significance for quadrupedal suspensory locomotion are reviewed and the evolution of these traits is discussed in light of the new framework. Experimental data are largely limited to Choloepus, but help to deduce functional aspects of the anatomy in Bradypus as well. The most important adaptive traits are hands and feet modified into relatively rigid hook-like appendages, great mobility of all joints proximal to the midcarpal and transverse tarsal joints, relatively long arms with a relatively short scapula, a rounded thorax with a small diameter, a highly mobile sterno-clavicular articulation, and emphasis on powerful flexion in the proximal limb joints via advantageous lever arms. Despite these changes, patterns of limb kinematics remained conservative during the course of evolution in the lineages leading to extant tree sloths, and it is suggested here that this also applies to the pattern of neuromuscular control of limb movements during locomotion. Morphological ‘solutions’ to altered functional demands posed by inversed orientation of the body differ in the two genera of extant tree sloths, thereby corroborating the proposed diphyly. Convergent evolution in tree sloths may be attributed to functional constraints posed by fossorial adaptations in early Xenarthra that canalized sloths to adopt a suspensory posture and locomotion in the arboreal habitat.     

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.     

The Forelimb of Early Miocene Sloths (Mammalia, Xenarthra, Folivora): Morphometrics and Functional Implications for Substrate Preferences

Early Miocene sloths are represented by a diversity of forms ranging from 38 to 95?kg. Their forelimb bones differ in shape from those of their closest living relatives (less than 10?kg), Bradypus and Choloepus. Such differences in shape could be related to differences in substrate preference (arboreal, semiarboreal, or ground-dwelling) or substrate use (climbing, digging, etc.). In order to detect putative patterns related to substrate preference, 21 linear measurements were defined and taken on the forelimb bones. The sample was composed of 22 specimens of fossil sloths and 134 specimens of extant mammals (marsupials, xenarthrans, pangolins, rodents, primates, and carnivorans), including arboreal, semiarboreal, and ground-dwelling taxa. Principal Components Analyses were performed on logarithms of original measurements, while functional indexes (Index of Fossorial Ability, Brachial Index, and Distal Epiphyseal Index) were calculated on raw data. The first three PCs accounted for 93.8% of the cumulative variability. PC1 roughly represented size, while positive values of PC2 represented mechanical advantage for features related to digging habits. Fossil sloths were clearly separated from living ones, sharing a common morphospace with anteaters and other good diggers. Conversely, living sloths shared a morphospace with primates. Similar results were obtained for DEI and IFA, with fossil sloths showing similar values to extant digging mammals. These results suggest that fossil sloths have a different functional pattern of forelimb use than that of extant ones, probably more similar to vermilinguas and pangolins, including putative good digging capabilities and/or semiarboreal habits. Substrate use seems to be interfering in the analysis of substrate preference based on forelimb morphology.     

Limb kinematics during locomotion in the two-toed sloth (Choloepus didactylus,Xenarthra) and its implications for the evolution of the sloth locomotor apparatus

In order to gain insight into the function of the extant sloth locomotion and its evolution, we conducted a detailed videoradiographic analysis of two-toed sloth locomotion (Xenarthra: Choloepus didactylus). Both unrestrained as well as steady-state locomotion was analyzed. Spatio-temporal gait parameters, data on interlimb coordination, and limb kinematics are reported. Two-toed sloths displayed great variability in spatio-temporal gait parameters over the observed range of speeds. They increase speed by decreasing the durations of contact and swing phases, as well as by increasing step length. Gait utilization also varies with no strict gait sequence or interlimb timing evident in slow movements, but a tendency to employ diagonal sequence, diagonal couplet gaits in fast movements. In contrast, limb kinematics were highly conserved with respect to ‘normal’ pronograde locomotion. Limb element and joint angles at touch down and lift off, element and joint excursions, and contribution to body progression of individual elements are similar to those reported for non-cursorial mammals of small to medium size. Hands and feet are specialized to maintain firm connection to supports, and do not contribute to step length or progression. In so doing, the tarsometatarsus lost its role as an individual propulsive element during the evolution of suspensory locomotion. Conservative kinematic behavior of the remaining limb elements does not preclude that muscle recruitment and neuromuscular control for limb pro- and retraction are also conserved. The observed kinematic patterns of two-toed sloths improve our understanding of the convergent evolution of quadrupedal suspensory posture and locomotion in the two extant sloth lineages.     

The locomotion of Babakotia radofilai inferred from epiphyseal and diaphyseal morphology of the humerus and femur

Palaeopropithecids, or “sloth lemurs,” are a diverse clade of large‐bodied Malagasy subfossil primates characterized by their inferred suspensory positional behavior. The most recently discovered genus of the palaeopropithecids is Babakotia, and it has been described as more arboreal than Mesopropithecus, but less than Palaeopropithecus. In this article, the within‐bone and between‐bones articular and cross‐sectional diaphyseal proportions of the humerus and femur of Babakotia were compared to extant lemurs, Mesopropithecus and Palaeopropithecus in order to further understand its arboreal adaptations. Additionally, a sample of apes and sloths (Choloepus and Bradypus) are included as functional outgroups composed of suspensory adapted primates and non‐primates. Results show that Babakotia and Mesopropithecus both have high humeral/femoral shaft strength proportions, similar to extant great apes and sloths and indicative of forelimb suspensory behavior, with Babakotia more extreme in this regard. All three subfossil taxa have relatively large femoral heads, also associated with suspension in modern taxa. However, Babakotia and Mesopropithecus (but not Palaeopropithecus) have relatively small femoral head surface area to shaft strength proportions suggesting that hind‐limb positioning in these taxa during climbing and other behaviors was different than in extant great apes, involving less mobility. Knee and humeral articular dimensions relative to shaft strengths are small in Babakotia and Mesopropithecus, similar to those found in modern sloths and divergent from those in extant great apes and lemurs, suggesting more sloth‐like use of these joints during locomotion. Mesopropithecus and Babakotia are more similar to Choloepus in humerofemoral head and length proportions while Palaeopropithecus is more similar to Bradypus. These results provide further evidence of the suspensory adaptations of Babakotia and further highlight similarities to both extant suspensory primates and non‐primate slow arboreal climbers and hangers. J. Morphol. 277:1199–1218, 2016. © 2016 Wiley Periodicals, Inc.     

A Paleogeographic Overview of Tropical Fossil Sloths: Towards an Understanding of the Origin of Extant Suspensory Sloths?

Modern sloths are among the more characteristic mammals of South and Central American faunas. Recent discovery in four Paleogene, 22 Neogene, and dozens of Pleistocene fossiliferous localities in the tropics has revealed an unexpected paleobioversity constituted by some 81 fossil sloth species. Probably originating in southern South America near the Eocene/Oligocene transition, sloths were represented in the tropics during the late Oligocene by Pseudoglyptodon, Mylodontidae, and Megalonychidae. The latter occupied the West Indies between at least the late early Miocene and late Pleistocene, and two mylodontid clades, Octodontobradyinae and Urumacotheriinae, were characteristic of Amazonian localities from the Colhuehuapian and the Laventan periods, respectively, until the end of the Miocene. Megatheriinae and Nothrotheriidae appeared during the middle Miocene, colonizing the tropics and then North America, where Mylodontidae and Megalonychidae had already been present since the early late Miocene. Nothrotheriids are more abundant and diversified during the late Miocene in the tropics than in southern South America. Remains closely related to either of the modern sloths are absent from the fossil record, including those in the tropics. The characteristic suspensory posture of Bradypus and Choloepus appeared independently and likely after the Miocene epoch, and thus well after the hypothesized split suggested by molecular studies of the respective clades of these genera. Given their current widespread distribution in and reliance on the tropics, prospecting efforts for the direct fossil kin of suspensory sloths should concentrate on deposits in the Amazonian region, as this area has shown promise in producing fossil sloths.     

Thoracic member (pectoral girdle and forelimb) bones of <Emphasis Type="Italic">Mylodon darwinii</Emphasis> Owen (Xenarthra,Mylodontidae) from the Late Pleistocene of Central Argentina and their phylogenetic implications

A nearly complete skeleton, including most of the thoracic member bones of the sloth Mylodon darwinii, have been found in Upper Pleistocene strata from Anisacate River, Argentina. The thoracic member bones resemble their homologues in Glossotherium robustum, Paramylodon harlani, and Mylodonopsis ibseni in the following traits: (1) the olecranon is mediolaterally compressed; (2) the radius has an acute styloid process; (3) the radial diaphysis medial border is straight for two thirds of its length; (4) the radial shaft medial border forms an angle with the medial border of the styloid process. The radius presents a distinctive, mostly proximally facing articular circumference. The unfused epiphyses and feeble muscle attachment ridges indicate a sub-adult ontogenetic stage. Deviation of the olecranon and weak M. teres major origin and insertion, suggest a low fossorial specialization. The structure of the thoracic limb bones does not support climbing habits, because pronation-related features are reduced and the humeral head is not prominent. The structure of the radius suggests graviportal adaptations: the proximal head is mediolaterally expanded and the diaphysis straight. A phylogenetic analysis adding thoracic member characters recovers M. darwinii as part of a clade that includes Glossotherium robustum and Paramylodon harlani, but excludes Lestodon armatus. This contrasts with the results of previous analyses focusing on the head skeleton, highlighting the relevance of sampling postcranial characters in phylogenetic analyses of mylodontine sloths.     

Gradual adaptation of bone structure to aquatic lifestyle in extinct sloths from Peru

Non-pathological densification (osteosclerosis) and swelling (pachyostosis) of bones are the main modifications affecting the skeleton of land vertebrates (tetrapods) that returned to water. However, a precise temporal calibration of the acquisition of such adaptations is still wanting. Here, we assess the timing of such acquisition using the aquatic sloth Thalassocnus, from the Neogene of the Pisco Formation, Peru. This genus is represented by five species occurring in successive vertebrate-bearing horizons of distinct ages. It yields the most detailed data about the gradual acquisition of aquatic adaptations among tetrapods, in displaying increasing osteosclerosis and pachyostosis through time. Such modifications, reflecting a shift in the habitat from terrestrial to aquatic, occurred over a short geological time span (ca 4 Myr). Otherwise, the bones of terrestrial pilosans (sloths and anteaters) are much more compact than the mean mammalian condition, which suggests that the osteosclerosis of Thalassocnus may represent an exaptation.     

Premaxillae of the Extinct Megalonychid Sloths <Emphasis Type="Italic">Acratocnus</Emphasis>, <Emphasis Type="Italic">Neocnus</Emphasis>, and <Emphasis Type="Italic">Megalonyx</Emphasis>, and their Phylogenetic Implications (Mammalia,Xenarthra)

In most folivorans, the premaxilla is loosely attached to the maxilla, so that it is often missing in otherwise very well-preserved fossil skulls. Despite its infrequent preservation in sloths, the premaxilla has been shown to have phylogenetically significant variation among the taxa that do preserve the element. In the family Megalonychidae, the premaxilla is known only in the early taxon Eucholoeops (Santacrucian South American Land Mammal Age [SALMA]), the extant two-toed sloth Choloepus, and the North American Neogene taxon Megalonyx, the last described only in an unpublished Master’s thesis. We report here the discovery of the premaxilla in two genera of extinct megalonychids, Neocnus and Acratocnus. These small bodied, semiarboreal megalonychid sloths are endemic to the islands of the Greater Antilles. Though the presence of sloths in the Caribbean dates at least to the late Oligocene, the best known taxa derive from Pleistocene to Holocene cave deposits in Puerto Rico, Hispaniola, and Cuba. We also describe the premaxilla in two species of Megalonyx from North America, the Blancan North American Land Mammal Age (NALMA) M. leptostomus and Rancholabrean NALMA M. jeffersonii. These species show a progressive reorientation of the premaxilla within Megalonyx from a primitive horizontal element to a nearly vertical element, and some significant changes in the anatomy of the incisive foramen. Morphological evidence suggests that a broadened, plate-like premaxilla constitutes a synapomorphy for the entire clade Megalonychidae. Furthermore, although Eucholoeops retains a short anterior process of the premaxilla like that of megatherioid sloths, this process is lacking in the other megalonychids, suggesting that the loss of this process may unite late Miocene to Recent megalonychids.     

A mechanical link model of two-toed sloths: no pendular mechanics during suspensory locomotion

Sloths are morphologically specialized in suspensory quadrupedal locomotion and posture. During steady-state locomotion they utilize a trot-like footfall sequence. Contrasting the growing amount of published accounts of the functional morphology and kinematics of sloth locomotion, no study concerned with the dynamics of their quadrupedal suspensory locomotion has been conducted. Brachiating primates have been shown to travel at low mechanical costs using pendular mechanics, but this is associated with considerable dynamic forces exerted onto the support. To test whether sloth locomotion can be described by simple connected pendulum mechanics, we analyzed the dynamics of sloth locomotion with use of a mechanical segment link model. The model integrates the body segment parameters and is driven by kinematic data with both segment parameters and kinematic data obtained from the same sloth individual. No simple pendular mechanics were present. We then used the model to carry out an inverse dynamic analysis. The analysis allowed us to estimate net limb joint torques and substrate reaction forces during the contact phases. Predominant flexing limb joint torque profiles in the shoulder, elbow, hip, and knee are in stark contrast to published dominant extensor torques in the limb joints of pronograde quadrupedal mammals. This dissimilarity likely reflects the inverse orientation of the sloth towards the gravity vector. Nevertheless, scapular pivot and shoulder seem to provide the strongest torque for progression as expected based on unchanged basic kinematic pattern previously described. Our model predicts that sloths actively reduce the dynamical forces and moments that are transmitted onto the support. We conclude that these findings reflect the need to reduce the risk of breaking supports because in this case sloths would likely be unable to react quickly enough to prevent potentially lethal falls. To achieve this, sloths seem to avoid the dynamical consequences of effective pendular mechanics.     

A new nothrotheriid xenarthran from the early Pliocene of Pomata‐Ayte (Bolivia): new insights into the caniniform–molariform transition in sloths

Tardigrade xenarthrans are today represented only by the two tree sloth genera Bradypus and Choloepus, which inhabit the Neotropical rainforests and are characterized by their slowness and suspensory locomotion. Sloths have been recognized in South America since the early Oligocene. This monophyletic group is represented by five clades traditionally recognized as families: Bradypodidae, Megalonychidae, Mylodontidae (?), Megatheriidae (?) and Nothrotheriidae (?). A new nothrotheriid ground sloth represented by a dentary and several postcranial elements, Aymaratherium jeani gen. nov. , sp. nov. , from the early Pliocene locality of Pomata‐Ayte (Bolivia) is reported. This small‐ to medium‐sized species is characterized especially by its dentition and several postcranial features. It exhibits several convergences with the ‘aquatic’ nothrotheriid sloth Thalassocnus and the giant megatheriid ground sloth Megatherium (M.) americanum, and is interpreted as a selective feeder, with good pronation and supination movements. The tricuspid caniniform teeth of Aymaratherium may represent a transitional stage between the caniniform anterior teeth of basal megatherioids and basal nothrotheriids (1/1C‐4/3M as in Hapalops or Mionothropus) and the molariform anterior teeth of megatheriids (5/4M, e.g. Megatherium). To highlight the phylogenetic position of this new taxon among nothrotheriid sloths, we performed a cladistic assessment of the available dental and postcranial evidence. Our results, derived from a TNT treatment of a data matrix largely based on a published phylogenetic data set, indicate that Aymaratherium is either sister taxon to Mionothropus or sister to the clade Nothrotheriini within Nothrotheriinae. They further support the monophyly of both the Nothrotheriinae and the Nothrotheriini, as suggested previously by several authors.     

Grazing in a New Late Oligocene Mylodontid Sloth and a Mylodontid Radiation as a Component of the Eocene-Oligocene Faunal Turnover and the Early Spread of Grasslands/Savannas in South America

We describe a new taxon of mylodontid sloth from the late Oligocene (Deseadan South American Land Mammal “age”), Salla Beds of Bolivia. This taxon, Paroctodontotherium calleorum, new genus and species, is one of the oldest known sloths, but it is surprisingly derived. It is referable to the Mylodontidae and, with just a little doubt, to the Mylodontinae. It shares a number of derived characteristics with other mylodontids and even mylodontines. These include: a relatively low temporomandibular joint; a relatively short zygomatic process of the squamosal; an elongated, narrow braincase; anteriorly diverging toothrows; broad muzzle; and greatly enlarged external nares. The relative width of the muzzle of Paroctodontotherium is as great as any Pleistocene mylodontid except the giant grazer, Lestodon. We review and critique methods of estimating diets of extinct sloths and propose a hypothesis in regard to the feeding ecology of Paroctodontotherium. Based upon its broad muzzle, the degree of tooth wear, and its presence in a habitat dominated by hypsodont herbivores, we propose that Paroctodontotherium was a bulk feeder that foraged near ground level. Grasses were likely a major component of its diet. The addition of this new taxon, along with other recently discovered taxa, illustrates that late Oligocene sloths had much greater diversity than recognized just a decade ago. This diversity is evident in species richness, variations in body sizes, dental morphologies, and means of locomotion. We regard this relatively sudden sloth radiation as a significant component of the Eocene-Oligocene faunal turnover and was related to the development of more open habitats of post-Eocene South America.     

A peculiar climbing Megalonychidae from the Pleistocene of Peru and its implication for sloth history

The Xenarthra, particularly the Tardigrada, are with the Notoungulata and Marsupialia among the most diversified South American mammals. Lujanian South American Land Mammal Age localities from the coastal Piedra Escrita site and Andean Casa del Diablo Cave, Peru, have yielded three specimens of the Megalonychidae Diabolotherium nordenskioldi gen. nov. This singular fossil sloth exhibits a peculiar mosaic of cranial and postcranial characters. Some are considered convergent with those of other sloths (e.g. 5/4 quadrangular teeth, characteristic of Megatheriidae), whereas others clearly indicate climbing capabilities distinct from the suspensory mode of extant sloths. The arboreal mode of life of D. nordenskioldi is suggested by considerable mobility of the elbow, hip, and ankle joints, a posteriorly convex ulna with an olecranon shorter than in fossorial taxa, a radial notch that faces more anteriorly than in other fossil sloths and forms an obtuse angle with the coronoid process (which increases the range of pronation–supination), a proximodistally compressed scaphoid, and a wide range of digital flexion. D. nordenskioldi underscores the great adaptability of Tardigrada: an arboreally adapted form is now added to the already known terrestrial, subarboreal, and aquatic (marine and freshwater) fossil sloths. A preliminary phylogenetic analysis of the Tardigrada confirmed the monophyly of Megatherioidea, Nothrotheriidae, Megatheriidae, and Megalonychidae, in which Diabolotherium is strongly nested.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 179–235.     

Paleoenvironment of Dryopithecus brancoi at Rudabánya, Hungary: evidence from dental meso- and micro-wear analyses of large vegetarian mammals

The environment of the hominoid Dryopithecus brancoi at Rudabánya (Late Miocene of Hungary) is reconstructed here using the dietary traits of fossil ruminants and equids. Two independent approaches, dental micro- and meso-wear analyses, are applied to a sample of 73 specimens representing three ruminants: Miotragocerus sp. (Bovidae), Lucentia aff. pierensis (Cervidae), Micromeryx flourensianus (Moschidae), and one equid, Hippotherium intrans (Equidae). The combination of meso- and micro-wear signatures provides both long- and short-term dietary signals, and through comparisons with extant species, the feeding styles of the fossil species are reconstructed. Both approaches categorize the cervid as an intermediate feeder engaged in both browsing and grazing. The bovid Miotragocerus sp. is depicted as a traditional browser. Although the dental meso-wear pattern of the moschid has affinities with intermediate feeders, its dental micro-wear pattern also indicates significant intake of fruits and seeds. Hippotherium intrans was not a grazer and its dental micro-wear pattern significantly differs from that of living browsers, which may suggest that the fossil equid was engaged both in grazing and browsing. However, the lack of extant equids which are pure browsers prevents any definitive judgment on the feeding habits of Hippotherium. Based on these dietary findings, the Rudabánya paleoenvironment is reconstructed as a dense forest. The presence of two intermediate feeders indicates some clearings within this forest; however the absence of grazers suggests that these clearings were most likely confined. To demonstrate the ecological diversity among the late Miocene hominoids in Europe, the diet and habitat of Dryopithecus brancoi and Ouranopithecus macedoniensis (Greece) are compared.     

Feeding Ecology in Oligocene Mylodontoid Sloths (Mammalia,Xenarthra) as Revealed by Orthodentine Microwear Analysis

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.     

A new ground sloth (Mammalia: Xenarthra) from the Quaternary of Brazil

The record of South-American Pleistocene Megalonychidae is scarce. Of the species described for intertropical Brazil, including Megalonyx sp., Ocnopus gracilis, Valgipes deformis, Xenocnus cearensis and Ahytherium aureum, only the last, recently described, is valid. The new megalonychid species described here was recovered from the same locality as Ah. aureum. The latter is apparently more closely linked to the North-American Pleistocene forms whereas Australonyx aquae may be more closely related to the Antillean sloths. The fossil remains of extant taxa recovered in association with the new sloth species suggest that the region, currently within the Caatinga biome, was a mosaic of the Atlantic Forest and Savannah biomes during the final stages of the Pleistocene.     

Cretophengodidae,a new Cretaceous beetle family,sheds light on the evolution of bioluminescence

Bioluminescent beetles of the superfamily Elateroidea (fireflies, fire beetles, glow-worms) are the most speciose group of terrestrial light-producing animals. The evolution of bioluminescence in elateroids is associated with unusual morphological modifications, such as soft-bodiedness and neoteny, but the fragmentary nature of the fossil record discloses little about the origin of these adaptations. We report the discovery of a new bioluminescent elateroid beetle family from the mid-Cretaceous of northern Myanmar (ca 99 Ma), Cretophengodidae fam. nov. Cretophengodes azari gen. et sp. nov. belongs to the bioluminescent lampyroid clade, and would appear to represent a transitional fossil linking the soft-bodied Phengodidae + Rhagophthalmidae clade and hard-bodied elateroids. The fossil male possesses a light organ on the abdomen which presumably served a defensive function, documenting a Cretaceous radiation of bioluminescent beetles coinciding with the diversification of major insectivore groups such as frogs and stem-group birds. The discovery adds a key branch to the elateroid tree of life and sheds light on the evolution of soft-bodiedness and the historical biogeography of elateroid beetles.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.     

Bipedalism and quadrupedalism in Megatheriurn: an attempt at biomechanical reconstruction

Biomechanical reconstruction is increasingly being applied to the study of the mode of life of fossil animals. Different footprints from the fossil mammal Megatherium sp., the giant ground sloth, seem to indicate that it was able to use either bipedal or quadrupedal locomotion. By means of the estimation of the body mass of the type of the species Megatherium americanum , and using the published tracks, different mechanical parameters, such as speed, Froude number, indicators of athletic ability and bending and resistance moments of the vertebral column were calculated in both bipedal and quadrupedal conditions. Results on leg parameters are not conclusive as to the kind of locomotion to which Megatherium sp. was better adapted, but the calculations on the moments of resistance of the vertebral column and on the bending moment at breaking of the femur seem to indicate that Megatherium sp. presented adaptations to bipedalism. MEGATHERIUM, mammals, legs, vertebral column, locomotion, biomechanics, reconstruction .     

Ossified meniscus and cyamo-fabella in some fossil sloths: a morpho-functional interpretation

An articulated skeleton of Thalassocnus natans (Xenarthra : Nothrotheriidae) and a review of some other fossil sloths provide new information on sesamoid bones located at the knee joint. A sesamoid bone and an ossified meniscus have been identified at this joint. The cyamo-fabella (posterior sesamoid of the tibio-femoral articulation) of T. natans may act like a pulley through which the tendon of the m. gastrocnemius would have passed. The ossified meniscus, which is also present in Megatheriidae and Mylodontidae, could be related to the pedolateral stance and a large rotation of the knee joint during locomotion.