Squamates constitute a major vertebrate radiation, representing almost one-third of all known amniotes. Although speciose and morphologically diverse, they remain poorly represented in developmental studies. Here, we present an embryonic staging table of in ovo development for the basal gekkotan Eublepharis macularius (the leopard gecko) and advocate this species as a laboratory-appropriate developmental model. E. macularius, is a hardy and tractable species of relatively large body size (with concomitantly relatively large eggs and embryos), that is widely available and easy to maintain and propagate. Additionally, E. macularius displays a body plan appropriate to the study of the plesiomorphic quadrupedal condition of early pentadactylous terrestrial amniotes. Although not unexpected, it is worth noting that the morphological events characterizing limb development in E. macularius are comparable with those described for the avian Gallus gallus. Therefore, E. macularius holds great promise as a model for developmental studies focusing on pentadactyly and the formation of digits. Furthermore, it is also attractive as a developmental model because it demonstrates temperature-dependent sex determination. The staging table presented herein is based on an all-female series and represents the entire 52 day in ovo period. Overall, embryogenesis of E. macularius is similar to that of other squamates in terms of developmental stage attained at the time of oviposition, patterns of limb and pharyngeal arch development, and features of the appearance of scalation and pigmentation, indicative of a conserved developmental program.
Five nominal elements comprise the circumorbital series of bones in gekkotans: prefrontal, postfrontal, postorbital, jugal, and lacrimal. Determination of the homology of two of these, the postfrontal and postorbital, has been particularly problematic. Two conflicting hypothesis exist relating to these: either the postorbital is lost and the postfrontal remains or they fuse during development to form a combined element, the postorbitofrontal. Such a combined element apparently occurs in at least some members of all lizard clades. There is, however, no direct developmental evidence that supports either theory. To overcome that, we investigate the sequence and pattern of ossification in the circumorbital region in a developmental series of the Leopard gecko. We posit that both the postfrontal and postorbital appear during development. Contrary to previous predictions they neither fuses to each other, nor do either degenerate. Instead, the postfrontal shifts anteriorly and fuses with the frontal to become indistinguishable from it by the time of hatching, and the postorbital persists as a robust independent element bounding the frontoparietal suture. These observations accord, in part, with both hypotheses of homology of these elements and result in the recognition of a new pattern, placing in doubt the existence of the composite postorbitofrontal. The phylogenetic implications of these findings may prove to be far reaching if similar and conserved patterns of development are encountered in other clades.
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