Osteoderm histology of the Pampatheriidae (Cingulata,Xenarthra, Mammalia): Implications for systematics,osteoderm growth,and biomechanical adaptation

Pampatheres are extinct, large‐bodied cingulates, which share morphological characters with both armadillos and glyptodonts but are considered to be more closely related to the latter. The osteoderm histology of six pampathere taxa was examined and compared to the histology of other cingulate osteoderms. This study investigates the development and functional adaptation of pampathere osteoderms as well as the phylogenetic relationships of the Pampatheriidae within the Cingulata. We found that pampathere osteoderms share a uniform histological organization based on a basic diploe‐like structure. After initial stages of intramembranous growth, metaplastic ossification, that is, the direct incorporation and mineralization of pre‐existing protein fibers, plays an important role in osteoderm development and provides information on various kinds of soft tissue otherwise not preserved. The latest stages of osteoderm growth are dominated by periosteal bone formation especially in the superficial cortex. Movable band osteoderms show regular arrangements of incorporated fibers that may increase the resistance of particularly weak areas against strain. The histological composition of pampathere osteoderms is plesiomorphic in its basic structure but shows a number of derived features. A unique array of Sharpey's fibers that are incorporated into the bone matrix at sutured osteoderm margins is interpreted as a synapomorphy of pampatheres. The arrangement of dermal fibers in the deep and superficial cortexes supports the close relationship between pampatheres and glyptodonts. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Dermal armour histology of aetosaurs (Archosauria: Pseudosuchia), from the Upper Triassic of Argentina and Brazil

Cerda, I.A. & Desojo, J.B. 2010: Dermal armour histology of aetosaurs (Archosauria: Pseudosuchia), from the Upper Triassic of Argentina and Brazil. Lethaia, Vol. 44, pp. 417–428. One of the most striking features documented in aetosaurs is the presence of an extensive bony armour composed of several osteoderms. Here, we analyse the bone microstructure of these elements in some South American Aetosaurinae aetosaurs, including Aetosauroides scagliai. In general terms, Aetosaurinae osteoderms are compact structures characterized by the presence of three tissue types: a basal cortex of poorly vascularized parallel‐fibred bone tissue, a core of highly vascularized fibro‐lamellar bone, and an external cortex of rather avascular lamellar bone tissue. Sharpey’s fibres are more visible at the internal core, toward the lateral margins and aligned parallel to the major axis of the dermal plate. No evidence of metaplastic origin is reported in the osteoderms, and we hypothesize an intramembranous ossification for these elements. The bone tissue distribution reveals that the development of the osteoderm in Aetosaurinae starts in a position located medial to the plate midpoint, and the main sites of active osteogenesis occur towards the lateral and medial edges of the plate. The osteoderm ornamentation is originated and maintained by a process of resorption and redeposition of the external cortex, which also includes preferential bone deposition in some particular sites. Given that no secondary reconstruction occurs in the osteoderms, growth marks are well preserved and they provide very important information regarding the relative age and growth pattern of Aetosaurinae aetosaurs. □Aetosauria, Aetosauroides, Archosauria, bone microstructure, integumentary skeleton, osteoderm.     

Do the ornamented osteoderms influence the heat conduction through the skin? A finite element analysis in Crocodylomorpha

In order to assess the implication of the crocodylomorph ornamented osteoderms on the skin conduction during basking, we have performed three dimensional modeling and finite element analyses on a sample which includes both extant dry bones and well-preserved fossils tracing back to the Early Jurassic. In purpose to reveal the possible implication of the superficial ornamentation on the osteoderm heat conduction, we repeated the simulation on an equivalent set of smoothed 3D-modeled osteoderms. The comparison of the results evidenced that the presence of the apical sculpture has no significant impact on the osteoderm global conduction. Furthermore, as we also aimed to assess the influence of the inner bone porosity on the osteoderm conduction, we modified the heat equation parameters so that the 3D-modeled osteoderms successively score the compact and the cancellous bone properties (i.e. mass density, heat capacity, thermal conductivity and thermal diffusivity). Finally, we repeated the analyses using the soft-dermis properties which lead to outline that neither the degree of porosity nor the presence of the osteoderms (in itself) significantly modifies the heat conduction through the crocodylomorph skin. Consequently, as hypothesized by previous authors, if the dermal shield happens to be involved into heat capture during basking for crocodylians, this process must mainly rely on a convective effect based on the osteoderm relative degree of vascularization. This last assumption could thus explain why the crocodylians which produce little metabolic heat would carry an entire vascularized osteoderm shield.     

The development of cephalic armor in the tokay gecko (Squamata: Gekkonidae: Gekko gecko)

Armored skin resulting from the presence of bony dermal structures, osteoderms, is an exceptional phenotype in gekkotans (geckos and flap-footed lizards) only known to occur in three genera: Geckolepis, Gekko, and Tarentola. The Tokay gecko (Gekko gecko LINNAEUS 1758) is among the best-studied geckos due to its large size and wide range of occurrence, and although cranial dermal bone development has previously been investigated, details of osteoderm development along a size gradient remain less well-known. Likewise, a comparative survey of additional species within the broader Gekko clade to determine the uniqueness of this trait has not yet been completed. Here, we studied a large sample of gekkotans (38 spp.), including 18 specimens of G. gecko, using X-rays and high-resolution computed tomography for visualizing and quantifying the dermal armor in situ. Results from this survey confirm the presence of osteoderms in a second species within this genus, Gekko reevesii GRAY 1831, which exhibits discordance in timing and pattern of osteoderm development when compared with its sister taxon, G. gecko. We discuss the developmental sequence of osteoderms in these two species and explore in detail the formation and functionality of these enigmatic dermal ossifications. Finally, we conducted a comparative analysis of endolymphatic sacs in a wide array of gekkotans to explore previous ideas regarding the role of osteoderms as calcium reservoirs. We found that G. gecko and other gecko species with osteoderms have highly enlarged endolymphatic sacs relative to their body size, when compared to species without osteoderms, which implies that these membranous structures might fulfill a major role of calcium storage even in species with osteoderms.     

Osteohistological characterization of notosuchian osteoderms: Evidence for an overlying thick leathery layer of skin

Osteoderms are mineralized structures embedded in the dermis, known for nonavian archosaurs, squamates, xenarthrans, and amphibians. Herein, we compared the osteoderm histology of Brazilian Notosuchia of Cretaceous age using three neosuchians for comparative purposes. Microanatomical analyses showed that most of them present a diploe structure similar to those of other pseudosuchians, lizards, and turtles. This structure contains two cortices (the external cortex composed of an outer and an inner layers, and the basal cortex) and a core in-between them. Notosuchian osteoderms show high bone compactness (>0.85) with varying degrees of cancellous bone in the core. The neosuchian Guarinisuchus shows the lowest bone compactness with a well-developed cancellous layer. From an ontogenetic perspective, most tissues are formed through periosteal ossification, although the mineralized tissues observed in baurusuchid LPRP/USP 0634 suggest a late metaplastic development. Histology suggests that the ossification center of notosuchian osteoderm is located at the keel. Interestingly, we identified Sharpey's fibers running perpendicularly to the outer layer of the external cortex in Armadillosuchus arrudai, Itasuchus jesuinoi, and Baurusuchidae (LPRP/USP 0642). This feature indicates a tight attachment within the dermis, and it is evidence for the presence of an overlying thick leathery layer of skin over these osteoderms. These data allow a better understanding of the osteohistological structure of crocodylomorph dermal bones, and highlight their structural diversity. We suggest that the vascular canals present in some sampled osteoderms connecting the inner layer of the external cortex and the core with the external surface may increase osteoderm surface and the capacity of heat transfer in terrestrial notosuchians.     

Biomechanics of the vertebrae and associated osteoderms of the Early Permian amphibian Cacops aspidephorus

Two series of osteoderms associated with the anterior three-quarters of the presacral vertebral column of the Early Permian temnospondylous amphibian Cacops aspidephorus have important implications for biomechanics of the axial skeleton. An internal series consists of an osteoderm fused to the distal tip of each neural spine. Lying dorsal to the internal series and overlapping each internal osteoderm is a second external series. The orientation of the zygapophyseal facets implies modest lateral flexion with limited coupled axial rotation of the column. However, the osteoderms restricted any possible lateral flexion through their inverted V-shape, strongly angled overlap between each external osteoderm and its neighbouring internal osteoderms, and the presence of a midsagittal flange on the ventral surface of each external osteoderm that fits into grooves on the anterior and posterior edges of the neighbouring internal osteoderms. This configuration allowed vertical flexion of the vertebral column with little lateral flexion. The rod-like nature of osteoderms with the anterior three-quarters of the presacral vertebrae suggests a restricted form of forward movement for Cacops unlike that of other early tetrapods.     

Osteoderm histology of Proterochampsia and Doswelliidae (Reptilia: Archosauriformes) and their evolutionary and paleobiological implications

Postcranial osteoderms are commonly developed in the major lineages of Archosauriformes, including forms such as proterochampsids and doswelliids. Here, we survey the histology of osteoderms of the doswelliids Archeopelta arborensis and Tarjadia ruthae, and the proterochampsids Chanaresuchus bonapartei and Pseudochampsa ischigualastensis to understand better the morphogenesis of these skeletal elements. Whereas, the Doswelliid osteoderms possess a trilaminar organization, in which two cortices (external and basal) can be differentiated from an internal core of cancellous bone, these elements are compact structures in proterochampsids. The osteoderms of P. ischigualastensis are avascular and they consist entirely of parallel‐fibered bone. Conversely, the osteoderms of C. bonapartei are well vascularized structures composed of zones of woven‐fibered bone and annuli of parallel‐fibered bone. The rather simple microstructure observed in P. ischigualastensis osteoderms suggests that these elements grew at a constant, low rate. Compared with proterochampsids, doswelliid osteoderms possess a more complex histology, which appears to be linked to variations in the growth rate during the osteoderm formation and also to the development of the external ornamentation. A comparison of our findings with the results of earlier studies on other archosauriforms (phytosaurs and pseudosuchians) reveals that the general osteoderm histology of doswelliids bears a closer resemblance to that of phytosaurs and pseudosuchians than the proterochampsid osteoderm microstructure. If all archosauriform osteoderms are homologous structures, the closer resemblance of doswellid osteoderm microstructures to that of phytosaurs and pseudosuchians is in agreement with the hypothesis that doswellids are more closely related to archosaurs than proterochampsids. J. Morphol. 276:385–402, 2015. © 2015 Wiley Periodicals, Inc.     

Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods

Buchwitz, M., Witzmann, F., Voigt, S. and Golubev, V. 2012. Osteoderm microstructure indicates the presence of a crocodylian‐like trunk bracing system in a group of armoured basal tetrapods. —Acta Zoologica (Stockholm) 93 : 260–280. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel‐fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel‐fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio‐ or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian‐like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.     

A survey of osteoderm histology and ornamentation among Crocodylomorpha: A new proxy to infer lifestyle?

Osteoderms of eight extant and extinct species of crocodylomorphs are studied histologically and morphologically. Most osteoderms display the typical “crocodilian” structure with a woven-fibered matrix surrounded by an upper and a lower parallel fibered matrix. The dorsal ornamentation of those specimens consists of a pit-and-ridge structure, with corresponding remodeling mechanisms. However, an osteoderm of Iberosuchus, studied here for the first time, differs in being nearly devoid of ornamentation; moreover, it shows strong bundles of straight Sharpey's fibers perpendicular to the surface in its lateral and dorsal walls, along with a rough plywood-like structure in its basal plate. This suggests that this osteoderm was more deeply anchored within the dermis than the other osteoderms studied hitherto. This peculiar structure might have been linked to a terrestrial ecology and a specific thermoregulation strategy. Some other notosuchians in our sample do not exhibit ornamentation on their osteoderms, as opposed to neosuchians. Considering current interpretations of osteoderm function(s) in crocodilians, our observations are discussed in reference to possible ecophysiological peculiarities of Notosuchia in general, and Iberosuchus in particular.     

The bone histology of osteoderms in temnospondyl amphibians and in the chroniosuchian Bystrowiella

Witzmann, F. and Soler-Gijón, R. 2010. The bone histology of osteoderms in temnospondyl amphibians and in the chroniosuchian Bystrowiella . — Acta Zoologica (Stockholm) 91 : 96–114
Bone histology of osteoderms in the armoured temnospondyl Peltobatrachus , plagiosaurids ( Gerrothorax , Plagiosuchus ) and dissorophids ( Aspidosaurus , Cacops , Platyhystrix ), as well as in the chroniosuchian Bystrowiella , is studied. The massive osteoderms of Peltobatrachus and Gerrothorax consist of homogeneous parallel-fibred bone, whereas in dissorophids, a lightly built, trabecular middle region is mantled by a thin cortex that is composed of a plywood-type structure. In Bystrowiella and Plagiosuchus , the osteoderms consist to a large degree of interwoven primary fibres and have cell lacunae that bear stumpy canaliculi. The differences in the histological structure of dissorophids and plagiosaurids suggest an iterative evolution of osteoderms. Furthermore, histology in Plagiosuchus indicates a metaplastic development of the osteoderms, whereas the osteoderms of Gerrothorax represent periosteal ossifications as in dissorophids. This suggests a convergent origin of osteoderms also within plagiosaurids. The extensive armour in Gerrothorax probably constituted a calcium reservoir, indicated by cyclical resorption events preserved in the external cortex and interpreted as a physiological response to periodic changes in salinity of the aquatic environment. In contrast, the unique osteoderm structure of dissorophids provides maximum stability and minimum bone mass, and is coherent with the interpretation that the osteoderms served to strengthen the vertebral column during terrestrial locomotion.     

Micromorphology of osteoderms in Dasypodidae (Cingulata,Mammalia): Characterization and 3D-reconstructions

Osteoderms are present in a variety of extinct and extant vertebrates, but among mammals, the presence of osteoderms is essentially restricted to armadillos (Cingulata, Dasypodidae). Osteoderms have been proposed to exhibit a variety of functionalities in Dasypodidae, mainly protection and thermoregulation, and they have been considered as one of the synapomorphies of this group. In this study, we use high-resolution microcomputed tomography to describe the osteoderm micromorphology of several extant species of Dasypodidae in a comparative context. This study allowed the identification, 3D-reconstruction and volume quantification of different internal structures of osteoderms as well as their interrelations. This detailed characterization of the internal osteoderm morphology was compared in a phylogenetic context to assess the evolutionary trends of the species involved. This enables the identification of distinctive patterns for the most widely recognized clades, the Dasypodinae and Euphractinae with a morphological homogeneity in the microstructure of their osteoderms, in comparison with Tolypeutinae where it has not been possible to establish a common morphological pattern. The most important features for linage differentiation is the degree of compaction of the osteoderms, the number of cavities and the development of hairs. It is likely that the differential development of the various structures occurred as adaptive response to climate changes.     

Bone microstructures and mode of skeletogenesis in osteoderms of three pareiasaur taxa from the Permian of South Africa

The extinct parareptilian clade of pareiasaurs was in the past often presented to constitute a morphocline from larger, less armoured forms to smaller, well armoured forms, indicating that the osteoderm cover became an increasingly prominent aspect in the post‐cranial skeleton of these animals. Here, we describe microanatomical and microstructural aspects of osteoderms of the three pareiasaur taxa Bradysaurus, Pareiasaurus and Anthodon from the Permian of South Africa. A generalized mode of osteoderm formation, consistent with intramembraneous skeletogenesis, is hypothesized to be present in all pareiasaurs. Few characters are shared between pareiasaur dermal armour and turtle shell bones and osteoderms. Otherwise, there is strong evidence from microanatomy and histology (i.e. absence of structures that formed via metaplasia of dermal tissue) that indicates nonhomology between pareiasaur dermal armour and the armour of living eureptiles. Analysis with bone profiler revealed no clear connection between bone compactness and lifestyle in the amniote osteoderm sample.     

Histological variability in fossil and recent alligatoroid osteoderms: Systematic and functional implications

Statements about morphological variation in extinct taxa often suffer from insufficient sampling that can be remedied by taking advantage of larger sample sizes provided by related, extant taxa. This analysis quantitatively and qualitatively examines histological and morphological variation of osteoderms from extant and extinct alligatoroid specimens. Statistically significant differences were correlated with changes in osteoderm size and shape. These differences are independent of position on the body, taxonomy, or evolution. Histological variation in alligatoroid osteoderms is due to morphological constraints on the elements themselves, and not taxonomic differences. This has implications for the recognition of histological characters in the osteoderms of extinct archosaur groups that lack extant representatives. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Comparative paleohistology in osteoderms of Pleistocene Panochthus sp. Burmeister, 1886 and Neuryurus sp. (Xenarthra,Glyptodontidae)

Xenarthran osteoderms are integumentary bones with high fossilization potential presenting a high degree of morphological and histological diversity. Here, new data on the osteoderms histology of two glyptodonts, Panochthus and Neuryurus are presented. The poor spatial organization of the mineralized fibers and a large trabecular area in the middle zone identified in Neuryurus indicate a different bone pattern than the one found in Panochthus, which is mainly characterized by a middle zone with less spongiosa. Through the Bone Profiler program, the degree of compactness of the specimens was obtained, with about 70% for Neuryurus sp. and approximately 90% for Panochthus sp., showing the difference in bone pattern. These values confirm the visible difference in the histological patterns of these taxa, especially in the middle zone. This work demonstrates the microstructural variation studied in osteoderms and shows the importance of paleohistology as a starting point for a better understanding of extinct taxa.     

Palaeohistology and external microanatomy of rauisuchian osteoderms (Archosauria: Pseudosuchia)

Abstract: The presence of postcranial dermal armour is plesiomorphic for Archosauria. Here, we survey the external microanatomy and histology of postcranial osteoderms (i.e. dorsal paramedian and caudal osteoderms) of rauisuchians, a widely distributed assemblage of extinct predatory pseudosuchians from the Triassic. The osteoderms of eight rauisuchian taxa were found to be rather compact bones, which usually lack significant bone remodelling or large areas of cancellous bone. The presence of highly vascularized woven or fibrolamellar bone tissue deposited in the core areas indicates higher growth rates during earlier life stages, whereas a more compact parallel‐fibred bone matrix indicates reduced growth rates in later development. This pattern of change corroborates earlier studies on long bone histology. With the exception of a bone tissue found in the sample of Batrachotomus kupferzellensis, which might be the result of metaplastic ossification, the general mode of skeletogenesis is comparable with intramembraneous ossification. The lack of cancellous bone tissue and remodelling processes associated with bone ornamentation, as well as the predominantly intramembraneous mode of ossification, indicates that rauisuchian osteoderm formation differs profoundly from that of the osteoderms of the only extant pseudosuchian lineage, the crocodylians.     

Ontogenetic histology of Stegosaurus plates and spikes

Abstract: The dinosaur Stegosaurus is characterized by osteoderms of alternating plates and terminal paired spikes. Previous studies have described the histological features and possible functions of these osteoderms. However, ontogenetic changes are poorly documented. In this study, the ontogenetic changes of the osteoderms are examined using eight different ontogenetic skeletons (a juvenile, a subadult, a young adult, and five old adults based on the cortical histology of their body skeletons). The juvenile plate and subadult spike show thin cortex and thick cancellous bone. The young adult plates have an extensive vascular network, which is also seen in old adults. Old adult spikes are different from old adult plates in having a thick cortex and a large axial channel. The cortical histology, in both plates and spikes, show well‐vascularized bone tissue consisting of dense mineralized fibres in young adult forms. In old adult forms, the bone tissues in the spikes become more compact and are extensively remodelled. This might contribute to the structural reinforcement of the spikes. The plates in old adult forms also show extensive remodelling and lines of arrested growth, but only limited signs of compaction. The timing for acquisition of features seen in old adults is different between plates (an extensive vascular network in the young adult) and spikes (a thick cortex with a large axial channel in old adults). The result suggests that the timing for plate and spike functions is different. The extensive vascular networks seen in large plates suggest their function is for display and/or thermoregulation. The thick cortical bone of spikes of old adults suggests that spikes acquire a weapon function for defence ontogenetically late.     

Exoskeleton and Systematics: A Historical Problem in the Classification of Glyptodonts

The glyptodonts (Mammalia: Cingulata) are characterized by an exoskeleton comprising most notably an armored tail and an immobile dorsal carapace formed by a large number of osteoderms. In 1889, Florentino Ameghino published the first phylogenetic scenario for the glyptodonts, based on the sequential application of two transformation series related to the morphology of the tail armor and carapace osteoderms. From the early to mid 1900s, several authors used Ameghino’s transformation series subordinated to a model of evolution in which derived glyptodont groups had arisen independently from separate pre-middle Miocene ancestors. This approach, in which the morphological states of Ameghino’s series were considered analogous rather than homologous, provided different phylogenetic scenarios and the paraphyletic classification still in use. Two recent cladistic analyses based on cranial and postcranial (including caudal tube) characters support the monophyly of glyptodonts and suggest novel intra-clade relationships. However, neither analysis included the classic osteoderm characters used by earlier authors. Therefore, we propose new osteoderm and carapace characters and evaluate their performance in a new cladistic analysis. We found that: a) some osteoderm characters used by earlier authors to support ancestor-descendent hypotheses are in fact fully homoplastic autapomorphies (e.g., multiplication of the number of rows of peripheral figures); b) characters previously believed to have originated independently in several groups (e.g., presence of caudal tube) are synapomorphies at a wider hierarchical level; c) some ancestor–descendant pre-cladistic hypotheses are incompatible with the topology and synapomorphy distribution obtained; and d) there is no reason to favor exoskeletal characters in glyptodont systematics.     

Armored geckos: A histological investigation of osteoderm development in Tarentola (Phyllodactylidae) and Gekko (Gekkonidae) with comments on their regeneration and inferred function

Osteoderms are bone‐rich organs found in the dermis of many scleroglossan lizards sensu lato, but are only known for two genera of gekkotans (geckos): Tarentola and Gekko. Here, we investigate their sequence of appearance, mode of development, structural diversity and ability to regenerate following tail loss. Osteoderms were present in all species of Tarentola sampled (Tarentola annularis, T. mauritanica, T. americana, T. crombei, T. chazaliae) as well as Gekko gecko, but not G. smithii. Gekkotan osteoderms first appear within the integument dorsal to the frontal bone or within the supraocular scales. They then manifest as mineralized structures in other positions across the head. In Tarentola and G. gecko, discontinuous clusters subsequently form dorsal to the pelvis/base of the tail, and then dorsal to the pectoral apparatus. Gekkotan osteoderm formation begins once the dermis is fully formed. Early bone deposition appears to involve populations of fibroblast‐like cells, which are gradually replaced by more rounded osteoblasts. In T. annularis and T. mauritanica, an additional skeletal tissue is deposited across the superficial surface of the osteoderm. This tissue is vitreous, avascular, cell‐poor, lacks intrinsic collagen, and is herein identified as osteodermine. We also report that following tail loss, both T. annularis and T. mauritanica are capable of regenerating osteoderms, including osteodermine, in the regenerated part of the tail. We propose that osteoderms serve roles in defense against combative prey and intraspecific aggression, along with anti‐predation functions. J. Morphol. 276:1345–1357, 2015. © 2015 Wiley Periodicals, Inc.     

Osteoderm morphology in recent and fossil euphractine xenarthrans

The presence of osteoderms within the integument, forming a carapace, is one of the most distinctive features of armadillos with the external morphology of these elements forming the basis of most systematic schemes. This is especially true for fossil taxa, where these elements are most frequent in the palaeontological record. A detailed study of osteoderms from the cephalic shield and different regions of the dorsal armour of Chaetophractus villosus (Euphractinae, Xenarthra) was made and compared to those of the extant genus Dasypus (Dasypodinae, Xenarthra), and the extinct genus ?Eutatus. Three distinct histological zones were recognized: outer and inner zones are thin, formed by regular compact bone, the middle zone is thicker, with large cavities that contain mainly adipose tissue, hair follicles, and sweat and sebaceous glands. The internal structure of ?Eutatus (also a member of Euphractinae) osteoderms is close to that of C. villosus, consistent with the notion that these taxa are phylogenetically closely related. In contrast, Dasypus shows marked differences. Dasypus shows hair follicles associated with both gland types (sweat and sebaceous) and connected to foramina on the external surface. Although not observed in adult C. villosus, it has been documented during embryonic development, only to atrophy later in ontogeny. Furthermore, the presence of red bone marrow is rare in C. villosus, but widespread in Dasypus novemcinctus osteoderms. These results suggest an early split of both subfamilies and support the hypothesis that the Euphractinae are more derived than the Dasypodinae.