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.     

The structure of the osteoderms in the Gekko: Tarentola mauritanica

Histological and cytological analysis reveals that the osteoderms of Tarentola mauritanica are composed of an outer part superimposed on a basal region. The structure of both parts can be related to that of the surrounding dermis. The basal part of the osteoderms, inserted in the dense dermis, is made up of abundant closely packed collagen fibrils that orient the mineral deposit. The outer part, located in the superficial loose dermis, is crossed by few bundles of mineralized collagen fibrils arising from the basal part. These bundles connect the osteoderm to the overlying loose dermis. The outer superficial part is characterized by the presence of mineralized globules surrounding the mineralized collagen bundles. In these globules, the crystals are deposited on a microfibrillar matrix rich in acidic mucosubstances and composed of radially oriented, tangled microfilaments that lie among the collagen bundles. The two different mineralizing systems in the osteoderms of Tarentola mauritanica may reflect two different organic matrices. The mineral is deposited in a preexisting dermal tissue, as a "metaplastic ossification," and is another expression of the potential retained by the reptilian dermis to form mineralized structures.     

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.     

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.     

New material of Pachyarmatherium from the late Pleistocene of northeastern Brazil: insights into its morphology and systematics

A study of recently collected and other undescribed material of Pachyarmatherium brasiliense from the State of Pernambuco, Brazil, was performed, and its type material was revised, improving our understanding of this species and providing additional information on its affinities. Two fused osteoderms representing the posteriormost osteoderm row of the pectoral buckler are described for the first time in the Pachyarmatherium. Several additional specimens included buckler, border, and at least one caudal osteoderm. The relationships of Pachyarmatherium have been subject to different interpretations. We propose that the presence of some osteoderm features suggests a closer affinity with Dasypodinae/cf. Dasypodini armadillos. A key feature supporting this conclusion is the presence of complementary small figures in adjoining buckler osteoderms of Pachyarmatherium, indicating that a single scale covered portions of three contiguous osteoderms. Other features observed are related to the histological microstructure, which includes a central cortex characterized by a poorly developed cancellous bone and a deep cortex with mineralized collagen fiber bundles that are oriented roughly parallel to the deep osteoderm surface. Likewise, these histological features support its exclusion from glyptodonts and pampatheres, which show a different histological arrangement in their osteoderms.     

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.     

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.     

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.     

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.     

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.     

Osteoderm morphology and development in the nine-banded armadillo, Dasypus novemcinctus (Mammalia, Xenarthra, Cingulata)

Among modern mammals, armadillos (Xenarthra, Cingulata) are the only group that possesses osteoderms, bony inclusions within the integument. Along the body, osteoderms are organized into five discrete assemblages: the head, pectoral, banded, pelvic, and tail shields. The pectoral, banded, and pelvic shields articulate to form the carapace. We examined osteoderm skeletogenesis in the armadillo Dasypus novemcinctus using serial and whole-mount histochemistry. Compared with the rest of the skeleton, osteoderms have a delayed onset of development. Skeletogenesis begins as condensations of osteoblasts secreting osteoid, localized within the papillary layer of the dermis. Osteoderm formation is asynchronous both within each shield and across the body. The first osteoderms to mineralize are situated within the pectoral shield of the carapace, followed by elements within the banded, head, pelvic, and tail shields. In general, within each shield ossification begins craniomedially and proceeds caudally and laterally, except over the head, where the earliest elements form over the frontal and parietal bones. The absence of cartilage precursors indicates that osteoderms are dermal elements, possibly related to the all-encompassing vertebrate dermal skeleton (exoskeleton). The mode of development of D. novemcinctus osteoderms is unlike that described for squamate osteoderms, which arise via bone metaplasia, and instead is comparable with intramembranously derived elements of the skull.     

Osteoderm microstructure of “rauisuchian” archosaurs from South America

In this contribution we analyze and discuss the microanatomy and histology of postcranial osteoderms of a number of “rauisuchians” from different localities of South America (Argentina and Brazil). The studied sample includes osteoderms of Fasolasuchus tenax, Prestosuchus chiniquensis, Saurosuchus galilei and an undetermined rauisuchian from Brazil. The bone microanatomy of the osteoderms is variable: whereas some specimens have a rather compact structure, others show a diploe architecture with a central cancellous core bordered by two compact cortices. Both external and basal cortices are mainly composed of poorly vascularized, fine and coarse parallel-fibred bone and networks of interwoven and mineralized fiber bundles. The internal region of the non-remodeled specimens consists of a well-vascularized core in which the intrinsic fibers exhibit important variations (even in the same specimen), ranging from coarse, parallel-fibred to woven-fibred bone tissues. Lines of arrested growth (LAGs) are well recorded in both basal and external cortices. Differences in the bone microstructure (compact vs. diploe) could be related to the age, sex and reproductive status of the sampled individuals. Hence, age estimation based on the count of LAGs in rauisuchian osteoderms appears to be reliable only in the early stages of ontogeny. The bone microstructure suggests that rauisuchian osteoderms were originated through a mechanism that involves both intramembranous and metaplastic ossifications.     

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.     

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.     

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.     

Novel data on aetosaur (Archosauria,Pseudosuchia) osteoderm microanatomy and histology: palaeobiological implications

One of the most striking features of aetosaurs is the possession of an extensive bony armour composed of dorsal, ventral and appendicular osteoderms. With the purpose of establishing the main histological changes during ontogeny and the degree of histological variation within the armour, we analysed the bone histology of dorsal (paramedian and lateral), ventral and appendicular osteoderms from different taxa from the Late Triassic of South America, including Aetosauroides scagliai, Aetobarbakinoides brasiliensis and Neoaetosauroides engaeus. Histological data support an intramembranous origin for osteoderms. Nevertheless, evidence for metaplastic ossification (i.e. structural fibres) at advanced ontogenetic stages, in at least some elements, is also present. A variant type of parallel fibred bone, which we have named ‘crossed parallel fibred bone’, is characterized for aetosaurs. In this pseudosuchian group, osteoderms exhibit very important microstructural changes during ontogeny, which can be useful for determining ontogenetic stages from isolated elements. Histological data suggest a relatively early onset of sexual maturity among aetosaurs. Microanatomical analysis from different taxa reveal that having high values of compactness is the plesiomorphic condition for Aetosauria. The notably increased compactness of the osteoderms does not appear to be related to size, ontogeny, sex or reproductive status of the individuals. Although a high degree of compactness of osteoderms and other bones has been considered as evidence for an aquatic lifestyle in vertebrates, such an inference contradicts the current concept of a fully terrestrial lifestyle in aetosaurs.     

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.     

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.     

An enamel-like tissue,osteodermine, on the osteoderms of a fossil anguid (Glyptosaurinae) lizard

The Glyptosaurinae, a fossil clade of anguid lizards, have robust osteoderms, with a granular ornamentation consisting of tubercles. In this study, the structural and histological features of these osteoderms are described in order to reconstruct their developmental pattern and further document the possible homology that could exist between vertebrate integumentary skeletons. Glyptosaurine osteoderms display a diploe architecture and an unusually complex structure that includes four tissue types: an intensely remodeled core of woven-fibered bone, a thick basal layer of lamellar bone, a peripheral ring exhibiting histological features intermediate between these two tissues and containing dense bundles of long Sharpey fibers, and a superficial layer made of a monorefringent, acellular and highly mineralized material, different from bone, and comparable in many respects to hypermineralized tissues such as ganoine, enameloids and enamel. We call this tissue osteodermine. The growth pattern of glyptosaurine osteoderms is likely to have involved first metaplasia, at an early developmental stage, then appositional growth due to osteoblast activity. The superficial layer that is well developed at the tubercle level must have resulted from epidermal and dermal contributions, a conclusion that would support previous hypotheses on the role of epidermal-dermal interactions in the formation of squamate osteoderms.