Morphogenesis of shell and scutes in the turtle Emydura macquarii

Formation of the scutes and dermis of the embryonic shell of the turtle Emydura macquarii was studied using light and electron microscopy. Shell morphogenesis begins at embryonic stage 15 and the shape of the shell is mostly completed by embryonic stage 19. The carapace anlagen arises as a thickening of the skin in the dorsal part of the mid-trunk region between the anterior and posterior limbs. This thickening extends ventro-laterally to form ridges at the margins of the carapace. Each ridge forms as a thick epidermal placode over a condensation of mesenchymal cells. The epidermis behind the advancing margins of the carapace is cuboidal or columnar but does not form placodes. The margins of the carapace expand rapidly in all directions. The plastron anlagen is derived from epidermal cells localised in the latero-ventral regions between the fore- and hind-limbs. Plastron placodes are present laterally, while the mid-ventral and central epidermis remains cuboidal or columnar but does not form placodes at embryonic stage 16. The plastron thickening rapidly moves from a latero-ventral position to a flat ventral position between embryonic stages 16 and 19. Dermal–epidermal anchoring complexes occur throughout placodes of both the carapace and plastron, but are rare in non-placode areas. The accumulation of a dense mesenchyme beneath the shell epidermis forms a dermal cushion that surrounds the body cavity. The superficial dermis close to the epidermis is made of mesenchymal fibroblasts at embryonic stage 19, although the inner-most areas contain bipolar fibroblasts and extracellular fibrils. Scutes with serrations at their borders form as invaginations of the epidermis into the dermis in the mid-dorsal areas of the embryo at embryonic stages 18–19. Dermal–epidermal anchoring complexes are located around the infoldings that form the scutes of the hinge region. The epidermis of the shell has 2–3 suprabasal cells at embryonic stages 19–22, and lacks keratinisation before embryonic stage 22 when it has 4–6 suprabasal layers with 2–3 external layers made of flat cells. The dermis thickens and has numerous collagen fibrils after embryonic stage 19. The formation of dermal bones begins at embryonic stage 18–19 in the plastron. Only small areas of the carapace near to the bridge have begun to form dermal bone at embryonic stage 19. Calcification begins at embryonic stage 19, but is still incomplete at embryonic stages 24–25.