Scales of non-avian reptiles and their derivatives contain corneous beta proteins coded from genes localized in the Epidermal Differentiation Complex

The evolution of modern reptiles from basic reptilian ancestors gave rise to scaled vertebrates. Scales are of different types, and their corneous layer can shed frequently during the year in lepidosaurians (lizards, snakes), 1-2 times per year in the tuatara and in some freshwater turtle, irregularly in different parts of the body in crocodilians, or simply wore superficially in marine and terrestrial turtles. Lepidosaurians possess tuberculate, non-overlapped or variably overlapped scales with inter-scale (hinge) regions. The latter are hidden underneath the outer scale surface or may be more exposed in specific body areas. Hinge regions allow stretching during growth and movement so that the skin remains mechanically functional. Crocodilian and turtles feature flat and shield scales (scutes) with narrow inter-scale regions for stretching and growth. The epidermis of non-avian reptilian hinge regions is much thinner than the exposed outer surface of scales and is less cornified. Despite the thickness of the epidermis, scales are mainly composed of variably amount of Corneous Beta Proteins (CBPs) that are coded in a gene cluster known as EDC (Epidermal Differentiation Complex). These are small proteins, 100-200 amino acid long of 8-25 kDa, rich in glycine and cysteine but also in serine, proline and valine that participate to the formation of beta-sheets in the internal part of the protein, the beta-region. This region determines the further polymerization of CBPs in filamentous proteins that, together a network of Intermediate Filament Keratins (IFKs) and other minor epidermal proteins from the EDC make the variable pliable or inflexible corneous material of reptilian scales, claws and of turtle beak. The acquisition of scales and skin derivatives with different mechanical and material properties, mainly due to the evolution of reptile CBPs, is essential for the life and different adaptations of these vertebrates.