Developmental mechanisms underlying differential claw expression in the autopodia of geckos

BACKGROUND: The limb and autopodium are frequently employed to study pattern formation during embryonic development, providing insights into how cells give rise to complex anatomical structures. With regard to the differentiation of structures at the distal tips of digits, geckos constitute an attractive clade, because within their ranks they exhibit multiple independent occurrences of claw loss and reduction, these being linked to the development of adhesive pads. The developmental patterns that lead to claw loss, however, remain undescribed. Among geckos, Tarentola is a genus characterized by large claws on digits III and IV of the manus and pes, with digits I, II, and V bearing only vestigial claws, or lacking them entirely. The variable expression of claws on different digits provides the opportunity to investigate the processes leading to claw reduction and loss within a single species. RESULTS: Here, we document the embryonic developmental dynamics that lead to this intraspecifically variable pattern, focusing on the cellular processes of proliferation and cell death. We find that claws initially develop on all digits of all autopodia, but, later in development, those of digits I, II, and V regress, leading to the adult condition in which robust claws are evident only on digits III and IV. Early apoptotic activity at the digit tips, followed by apoptosis of the claw primordium, premature ossification of the terminal phalanges, and later differential proliferative activity are collectively responsible for claw regression in particular digits. CONCLUSIONS: Claw reduction and loss in Tarentola result from differential intensities of apoptosis and cellular proliferation in different digits, and these processes have already had some effect before visible signs of claw development are evident. The differential processes persist through later developmental stages. Variable expression of iteratively homologous structures between digits within autopodia makes claw reduction and loss in Tarentola an excellent vehicle for exploring the developmental mechanisms that lead to evolutionary reduction and loss of structures. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1186/s13227-015-0003-9) contains supplementary material, which is available to authorized users

Eggshell morphology and gekkotan life-history evolution

Eggshell structure is related to fundamental aspects of embryonic development (via water and gas exchange), adult ecology and behavior (via nest site selection), and demography (via effects on survival). We compared life-history characteristics between gekkotans that lay rigid- versus parchment- shelled eggs to determine if evolutionary shifts in eggshell structure are associated with life-history evolution. Ancestral gekkotans laid parchment-shelled eggs, with rigid-shelled eggs evolving later. Clutch size in oviparous gekkotans is fixed at one or two eggs, and this characteristic eliminates an egg size versus clutch size tradeoff as a life-history strategy. We found that species laying rigid-shelled eggs exhibit (1) smaller eggs relative to adult body size, (2) smaller hatchlings relative to the size of the egg, (3) earlier embryonic stage at oviposition, (4) longer incubation periods, and (5) smaller adult body sizes than species laying parchment-shelled eggs. These patterns hold when accounting for phylogenetic relatedness, and are not explained by geographic distributions of climate and habitat. In general, our data support the hypothesis that the spherical shapes of rigid-shelled eggs limit their size (volume), which in turn has restricted hatchling size and adult body size. In contrast, while parchment-shelled eggs are similarly constrained in width, elongate shapes allow egg sizes, and hence hatchling sizes, to increase relative to adult body sizes. Finally, the evolution of rigid-shelled eggs may have allowed gekkotans to become so successful; over 1,000 species lay rigid-shelled eggs, as compared to about 200 species that lay eggs exhibiting the ancestral parchment-shelled condition