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The turtle shell is composed of dorsal armor (carapace) and ventral armor (plastron) covered by a keratinized epithelium. There are two epithelial appendages of the turtle shell: scutes (large epidermal shields separated by furrows and forming a unique mosaic) and tubercles (numerous small epidermal bumps located on the carapaces of some species). In our perspective, we take a synthetic, comparative approach to consider the homology and evolution of these integumental appendages. Scutes have been more intensively studied, as they are autapomorphic for turtles and can be diagnostic taxonomically. Their pattern of tessellation is stable phylogenetically, but labile in the individual. We discuss the history of developmental investigations of these structures and hypotheses of evolutionary and anomalous variation. In our estimation, the scutes of the turtle shell are an evolutionary novelty, whereas the tubercles found on the shells of some turtles are homologous to reptilian scales. J. Exp. Zool. (Mol. Dev. Evol.) 324B: 221–229, 2015. © 2015 Wiley Periodicals, Inc.

Generally, the bony armor of reptiles consists of separate osteoderms which develop in the dermis independently from the internal skeleton. Such a dermal armor is characteristic of ancient pareiasaurs that are closely related to Testudines. On the basis of this fact it has been assumed that the turtle shell is built up as the fusion of the osteoderms with some elements of the internal skeleton. However, this concept is not confirmed by morphogenetic data. During turtle ontogeny the reduction of the trunk myomeres leads to sinking of the vertebrae and ribs into the dermis. The neural and costal plates of the dorsal disk form as the outgrowths of these endoskeletal bones on inside the dermis. There are no independent superficial ossicles over the general plastral bony primordia in the ventral region of the body. Each plastral plate develops only from a single primordium like the clavicles, interclavicle and gastralia of other reptiles. Most likely, Testudines progressed in a unique evolutionary direction. Their bony shell is mainly the result of modification and consolidation of internal skeletal elements. The real osteoderms develop only on the body margins as connections between the dorsal and the ventral discs.

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Turtle horny shell has a scute pattern, which is conservative through evolution and across species. The discovery of epidermal placodes as the scute primordia and their strict topographical association to the somites of the turtle embryo suggested a new interpretation of the developmental mechanism of the scute pattern. Here, we tested the hypothesis that horny scutes develop from a mosaic of placodes corresponding exactly to the paths of myoseptae, with vertebral and pleural scutes developing staggered in adjacent segments, and marginal scutes developing in every segment. This scheme predicts little variation in marginals and suggests intercalary supernumerary scutes as potential variations for the vertebral and pleural rows. We examined spatial and numerical variations of the horny shell in 655 newly hatched olive ridley sea turtle, Lepidochelys olivacea, which is known to have a highly variable horny shell. In total, 120 patterns of carapacial scutes and 10 patterns of scutes on plastron, differing in the number and position of scutes were found. The number of vertebral scutes varied from 4 to 10. Variations with five, six and seven vertebrals occurred with the greatest and nearly equally frequency (31.5% on average). Pleural scutes were from 5 to 10 at one or both sides, and the typical symmetric pattern for sea turtles with five pairs of pleurals was only seen in ca. 12% of specimens. In contrast, the majority of the specimens (92.7%) had just 13 pairs of marginals, showing a stable normal pattern. Similarly, on plastron the horny scutes were conservative, too; about 85% of specimens standardly had six pairs of plastral scutes and all specimens had four pairs of inframarginals. Despite a high level of variation of vertebral and pleural scutes in olive ridley turtle, all patterns fall into the theoretical spectrum of possible variants predicted by the segment-dependent model of development of the turtle horny shell. Therefore, the results of our analysis support the existence of direct morphogenetic correlation between the number and distribution of normal and supernumerary scutes and metamere organization of the turtle embryo.

The postembryonic development of the turtle carapace was studied in the aquatic Еmys orbicularis and the terrestrial Тestudo graeca. Differences in the structure of the bony shell in aquatic and terrestrial turtles were shown to be associated with varying degrees of development of epidermal derivatives, namely, the thickness of the scutes and the depth of horny furrows. Sinking of the horny furrows into the dermis causes local changes in the structure of the collagen matrix, which might precondition the acceleration of the ossification. Aquatic turtles possess a relatively thin horny cover, whose derivatives are either weakly developed or altogether absent and thus make no noticeable impact on the growth dynamics of bony plates. Carapace plates of these turtles outgrow more or less evenly around the periphery, which results in uniform costals, relatively narrow and partly reduced neurals, and broad peripherals extending beyond the marginal scutes. In terrestrial turtles (Testudinidae), horny structures are much more developed and exert a considerable impact on the growth of bony elements. As a result, bony plates outgrow unevenly in the dermis, expanding fast in the zones under the horny furrows and slowly outside these zones. This determines the basic features of the testudinid carapace: alternately cuneate shape of costals, an alternation of broad octagonal and narrow tetragonal neurals, and the limitation of the growth of peripherals by pleuro‐marginal furrows. The evolutionary significance of morphogenetic and constructional differences in the turtle carapace, and the association of these differences with the turtle habitats are discussed.

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