Patterns of dermal denticle loss in sharks.

As they grow, sharks both replace lost denticles and proliferate the number of denticles by developing new (de novo) denticles without prior denticle shedding. The loss and replacement of denticles has potential impacts on the energetic cost of maintaining the skin surface, the biomechanical functions of shark skin, as well as our ability to predict shark abundance from fossil denticle occurrence in sediment cores. Here, we seek to better understand patterns of denticle loss and to show how denticles are being replaced in mature sharks. We illustrate shark skin surfaces with missing denticles and quantify both within-species and between-species patterns of missing denticles using images from across regions of the body for two species and images at similar body regions for 16 species of sharks. Generally, sharks are missing similar numbers of denticles (0%-6%) between species and regions. However, there are exceptions: in the smooth dogfish, the nose region is missing significantly more denticles than most posterior-body and fin regions, and the common thresher shark is missing significantly more denticles than the smooth dogfish, leopard shark, angel shark, bonnethead, and gulper shark. Denticle regrowth starts with crown development and mineralization beneath the epidermis, followed by eruption of the crown, and finally the mineralization of the root. The pulp cavity of replacement denticles is initially large and surrounded by a thin shell of enameloid upon eruption of the denticle. After eruption of the denticle, the deposition of dentine continues internally after the denticle reaches its final position. Replacement of missing denticles, representing less than 6% of the skin surface at any one time, may not compromise hydrodynamic function, but by constantly updating the skin surface throughout life, sharks may reduce surface fouling and maintain a functional complex skin surface by repairing local damage to individual denticles.