Turtle morphology is unlike that of any other vertebrate. The uniqueness of the turtle's bodyplan is attributed to the manner in which the turtle's ribs are ensnared within its hard upper shell. The exact embryological and genetic mechanisms underpinning this peculiar anatomical structure are still a matter of debate, but biologists agree that the evolution of the turtle shell lies in the embryonic development of the turtle.
Two main elements characterize the skeletal morphology of turtles: the carapace and the plastron. For a turtle, the carapacial ridge begins in the embryo as a bulge posterior to the limbs but on both sides of the body. Such outgrowths are the first indication of shell development in turtle embryos. While the exact mechanisms underpinning the formation of the carapacial ridge are still not entirely known, some biologists argue that understanding these embryonic mechanisms is pivotal to explaining both the development of turtles and their evolutionary history.
Ann Campbell Burke examines the development and evolution of vertebrates, in particular, turtles. Her Harvard University experiments, described in Development of the Turtle Carapace: Implications for the Evolution of a Novel Bauplan, were published in 1989. Burke used molecular techniques to investigate the developmental mechanisms responsible for the formation of the turtle shell. Burke's work with turtle embryos has provided empirical evidence for the hypothesis that the evolutionary origins of turtle morphology depend on changes in the embryonic and developmental mechanisms underpinning shell production.
In "Behavioral Thermoregulation by Turtle Embryos," published in Proceedings of the National Academy of Sciences in April, 2011, Wei-Guo Du, Bo Zhao, Ye Chen, and Richard Shine report that turtle embryos can move towards warmer temperatures within the egg when presented with a small, 0.8 degrees Celsius gradient. This behavioral thermoregulation may benefit the embryo's fitness by accelerating the rate of development enough to decrease the incubation period by up to four and a half days. Embryos are generally thought to have little control over their surroundings. This study revealed that embryos may be able to control their developmental environment by modifying their behavior.
The sex of a reptile embryo partly results from the production of sex hormones during development, and one process to produce those hormones depends on the temperature of the embryo's environment. The production of sex hormones can result solely from genetics or from genetics in combination with the influence of environmental factors. In genotypic sex determination, also called genetic or chromosomal sex determination, an organism's genes determine which hormones are produced. Non-genetic sex determination occurs when the sex of an organism can be altered during a sensitive period of development due to external factors such as temperature, humidity, or social interactions. Temperature-dependent sex determination (TSD), where the temperature of the embryo's environment influences its sex development, is a widespread non-genetic process of sex determination among vertebrates, including reptiles. All crocodilians, most turtles, many fish, and some lizards exhibit TSD.