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.