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Osteoderms are mineralised structures consisting mainly of calcium phosphate and collagen. They form directly within the skin, with or without physical contact with the skeleton. Osteoderms, in some form, may be primitive for tetrapods as a whole, and are found in representatives of most major living lineages including turtles, crocodilians, lizards, armadillos, and some frogs, as well as extinct taxa ranging from early tetrapods to dinosaurs. However, their distribution in time and space raises questions about their evolution and homology in individual groups. Among lizards and their relatives, osteoderms may be completely absent; present only on the head or dorsum; or present all over the body in one of several arrangements, including non‐overlapping mineralised clusters, a continuous covering of overlapping plates, or as spicular mineralisations that thicken with age. This diversity makes lizards an excellent focal group in which to study osteoderm structure, function, development and evolution. In the past, the focus of researchers was primarily on the histological structure and/or the gross anatomy of individual osteoderms in a limited sample of taxa. Those studies demonstrated that lizard osteoderms are sometimes two‐layered structures, with a vitreous, avascular layer just below the epidermis and a deeper internal layer with abundant collagen within the deep dermis. However, there is considerable variation on this model, in terms of the arrangement of collagen fibres, presence of extra tissues, and/or a cancellous bone core bordered by cortices. Moreover, there is a lack of consensus on the contribution, if any, of osteoblasts in osteoderm development, despite research describing patterns of resorption and replacement that would suggest both osteoclast and osteoblast involvement. Key to this is information on development, but our understanding of the genetic and skeletogenic processes involved in osteoderm development and patterning remains minimal. The most common proposition for the presence of osteoderms is that they provide a protective armour. However, the large morphological and distributional diversity in lizard osteoderms raises the possibility that they may have other roles such as biomechanical reinforcement in response to ecological or functional constraints. If lizard osteoderms are primarily for defence, whether against predators or conspecifics, then this 'bony armour' might be predicted to have different structural and/or mechanical properties compared to other hard tissues (generally intended for support and locomotion). The cellular and biomineralisation mechanisms by which osteoderms are formed could also be different from those of other hard tissues, as reflected in their material composition and nanostructure. Material properties, especially the combination of malleability and resistance to impact, are of interest to the biomimetics and bioinspired material communities in the development of protective clothing and body armour. Currently, the literature on osteoderms is patchy and is distributed across a wide range of journals. Herein we present a synthesis of current knowledge on lizard osteoderm evolution and distribution, micro‐ and macrostructure, development, and function, with a view to stimulating further work. [ABSTRACT FROM AUTHOR]

Bone-producing osteoblasts and dentin-producing odontoblasts are closely related cell types, a result from their shared evolutionary history in the ancient dermal skeleton. In mammals, the two cell types can be distinguished based on histological characters and the cells' position in the pulp cavity or in the tripartite periodontal complex. Different from mammals, teleost fish feature a broad diversity in tooth attachment modes, ranging from fibrous attachment to firm ankylosis to the underlying bone. The connection between dentin and jaw bone is often mediated by a collar of mineralized tissue, a part of the dental unit that has been termed "bone of attachment". Its nature (bone, dentin, or an intermediate tissue type) is still debated. Likewise, there is a debate about the nature of the cells secreting this tissue: osteoblasts, odontoblasts, or yet another (intermediate) type of scleroblast. Here, we use expression of the P/Q rich secretory calcium-binding phosphoprotein 5 (scpp5) to characterize the cells lining the so-called bone of attachment in the zebrafish dentition. scpp5 is expressed in late cytodifferentiation stage odontoblasts but not in the cells depositing the "bone of attachment". nor in bona fide osteoblasts lining the supporting pharyngeal jaw bone. Together with the presence of the osteoblast marker Zns-5, and the absence of covering epithelium, this links the cells depositing the "bone of attachment" to osteoblasts rather than to odontoblasts. The presence of dentinal tubule-like cell extensions and the near absence of osteocytes, nevertheless distinguishes the "bone of attachment" from true bone. These results suggest that the "bone of attachment" in zebrafish has characters intermediate between bone and dentin, and, as a tissue, is better termed "dentinous bone". In other teleosts, the tissue may adopt different properties. The data furthermore support the view that these two tissues are part of a continuum of mineralized tissues. Expression of scpp5 can be a valuable tool to investigate how differentiation pathways diverge between osteoblasts and odontoblasts in teleost models and help resolving the evolutionary history of tooth attachment structures in actinopterygians. [ABSTRACT FROM AUTHOR]

The fin-to-limb transition was marked by the origin of digits and the loss of dermal fin rays. Paleontological research into this transformation has focused on the evolution of the endoskeleton, with little attention paid to fin ray structure and function. To address this knowledge gap, we study the dermal rays of the pectoral fins of 3 key tetrapodomorph taxa--Sauripterus taylori (Rhizodontida), Eusthenopteron foordi (Tristichopteridae), and Tiktaalik roseae (Elpistostegalia)--using computed tomography. These data show several trends in the lineage leading to digited forms, including the consolidation of fin rays (e.g., reduced segmentation and branching), reduction of the fin web, and unexpectedly, the evolution of asymmetry between dorsal and ventral hemitrichia. In Eusthenopteron, dorsal rays cover the preaxial endoskeleton slightly more than ventral rays. In Tiktaalik, dorsal rays fully cover the third and fourth mesomeres, while ventral rays are restricted distal to these elements, suggesting the presence of ventralized musculature at the fin tip analogous to a fleshy "palm." Asymmetry is also observed in cross-sectional areas of dorsal and ventral rays. Eusthenopteron dorsal rays are slightly larger than ventral rays; by contrast, Tiktaalik dorsal rays can be several times larger than ventral rays, and degree of asymmetry appears to be greater at larger sizes. Analysis of extant osteichthyans suggests that cross-sectional asymmetry in the dermal rays of paired fins is plesiomorphic to crown group osteichthyans. The evolution of dermal rays in crownward stem tetrapods reflects adaptation for a fin-supported elevated posture and resistance to substrate-based loading prior to the origin of digits. [ABSTRACT FROM AUTHOR]

A sample of late Viséan limestone from the Włodawa IG-4 borehole, east of Lublin, Poland, yielded a piece of a tooth and a few hundred well-preserved scales comparable to those of "Ctenacanthus" costellatus Traquair, 1884 from Glencartholm, Scotland, UK. Most of the scales are typical compound body scales of the ctenacanthid type. Their crowns are composed of several separate odontodes whose distal ends are turned backwards and bases are characterised by concave undersides. In the material, there are also sparse scales with similar crowns but with flat or convex bulbous bases, as well as ornamented plates and single, star-like denticles, probably from the head region. The taxonomic status of "Ctenacanthus" costellatus was analysed and a new generic name for that species, viz. Glencartius gen. nov., is proposed. [ABSTRACT FROM AUTHOR]

Cosmine is a combination of tissues (enamel, dentine, and bone) and a structure (pore-canal system) not found in living vertebrates. Long considered a synapomorphy of sarcopterygians (lobe-finned fishes), new discoveries and phylogenetic interpretations have shown that the assembly of cosmine is a deeper evolutionary event than previously thought and that cosmine constitutes a primitive feature of osteichthyans (bony fishes). A precursor of both cosmine and ganoine might have been present in the last common ancestor of sarcopterygians and actinopterygians (ray-finned fishes). However, despite numerous and thorough studies on cosmine since the late 19th century, its role and biological properties are still a mystery. Currently, the most consensual hypothesis considers cosmine as a metabolic regulator of mineral ions through intricated processes of deposition and resorption of dermal tissues. A new ontogenetic and phylogenetic scenario is emerging from microtomographic data revealing that a pore-canal system initially associated with multiple superimposed odontodes in stem osteichthyans and early sarcopterygians was reorganized and simplified in rhipidistians into a uniform cosmine sheet with a single layer of enamel overlying the dentine (eucosmine). In this review, I will summarize the current knowledge of cosmine origins, structure, development, and possible function based on previous research. I will also present new histological data on poorly known early sarcopterygians to highlight the importance of histological studies for upcoming analyses of sarcopterygian interrelationships. Future efforts to unravel the evolution of cosmine will need new virtual and three-dimensional approaches to paleohistology that will change our current framework of studying early vertebrate evolution and certainly revolutionize our understanding of this distinctive feature. [ABSTRACT FROM AUTHOR]

Osteoderms are bony deposits that form within the dermal layer of skin in vertebrates. Within geckos, osteoderms are uncommon, only described inGekko geckoand the genusTarentola. A previous report of osteoderms in the Afro-Malagasy gekkonid genusGeckolepishas been questioned due to the extreme skin fragility and regional integumentary loss within the group. We re-evaluated the integument anatomy ofGeckolepis maculatausing µCT scanning, histology, and clearing and staining to verify the presence or absence of osteoderms and, if present, to characterise and compare their morphology to the osteoderms ofGekko geckoandTarentola, as well as osteoderm diversity within squamates. Osteoderms were confirmed present inGeckolepis maculata; however, these osteoderms are compound, imbricating, squamous elements that are morphologically similar to the dermal ossifications observed in scincids and gerrhosaurids, whileG. geckoandTarentolapossess plate-like and granular osteoderms. Our results suggest that osteoderms have independently evolved at least three times within the Gekkota and that epidermal anatomy may be a better predictor of osteoderm morphology than ancestral history. Further research is required to investigate the regenerative capability ofGeckolepisosteoderms following skin autotomy. [ABSTRACT FROM AUTHOR]

During fin morphogenesis, several mesenchyme condensations occur to give rise to the dermal skeleton. Although each of them seems to create distinctive and unique structures, they all follow the premises of the same morphogenetic principle. Holmgren's principle of delamination was first proposed to describe the morphogenesis of skeletal elements of the cranium, but Jarvik extended it to the development of the fin exoskeleton. Since then, some cellular or molecular explanations, such as the “flypaper” model (Thorogood et al.), or the evolutionary description by Moss, have tried to clarify this topic. In this article, we review new data from zebrafish studies to meet these criteria described by Holmgren and other authors. The variety of cell lineages involved in these skeletogenic condensations sheds light on an open discussion of the contributions of mesoderm- versus neural crest-derived cell lineages to the development of the head and trunk skeleton. Moreover, we discuss emerging molecular studies that are disclosing conserved regulatory mechanisms for dermal skeletogenesis and similarities during fin development and regeneration, which may have important implications in the potential use of the zebrafish fin as a model for regenerative medicine. [ABSTRACT FROM AUTHOR]

Background Adaptation to a new environment can be facilitated by co-inheritance of a suite of phenotypes that are all advantageous in the new habitat. Although experimental evidence demonstrates that multiple phenotypes often map to the same genomic regions, it is challenging to determine whether phenotypes are associated due to pleiotropic effects of a single gene or to multiple tightly linked genes. In the threespine stickleback fish (Gasterosteus aculeatus), multiple phenotypes are associated with a genomic region around the gene Ectodysplasin (Eda), but only the presence of bony lateral plates has been conclusively shown to be caused by Eda. Results Here, we ask whether pleiotropy or linkage is responsible for the association between lateral plates and the distribution of the neuromasts of the lateral line. We first identify a strong correlation between plate appearance and changes in the spatial distribution of neuromasts through development. We then use an Eda transgene to induce the formation of ectopic plates in low plated fish, which also results in alterations to neuromast distribution. Our results also show that other loci may modify the effects of Eda on plate formation and neuromast distribution. Conclusions Together, these results demonstrate that Eda has pleiotropic effects on at least two phenotypes, highlighting the role of pleiotropy in the genetic basis of adaptation. [ABSTRACT FROM AUTHOR]

The characteristic and morphologically variable pectoral-fin spine of catfishes (order Siluriformes) has been well-investigated based on later developmental stages (juveniles and adults) but information on the earliest life stages are lacking. Here, we document the ontogeny of pectoral-fin spines in four siluroid (Ictalurus punctatus, Noturus gyrinus, Silurus glanis and Akysis vespa) and two loricarioid catfishes (Corydoras panda and Ancistrus sp.). To further our understanding of pectoral-fin spine development, we also examined adult and juvenile specimens representing 41 of the currently 43 recognized families of catfishes. Development of the pectoral-fin spine is similar in all catfishes and resembles the development of a typical soft fin ray. Fusion between hemitrichia of the anteriormost lepidotrichium occurs proximally first, forming the spine proper, with growth of the spine occurring through the subsequent fusion of developing distal hemitrichial segments that comprise the spurious ray. The variation of pectoral-fin spine morphology observed is largely attributed to the presence/absence of five traits, which either develop as part of the hemitrichial segments that are added to the distal tip of the spine during growth (distal rami, anterior/posterior serrae) or develop independent of these segments (denticuli and odontodes)., (© 2019 Wiley Periodicals, Inc.)

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., (© 2015 Wiley Periodicals, Inc.)