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Evolution of the Turtle Body Plan by the Folding and Creation of New Muscle Connections

Shelling Turtles In almost all vertebrates, the shoulder girdle (scapula) lies outside the ribs. The turtle is unique in that the carapace, the dorsal part of the shell, which is formed from the ribs, encapsulates the scapula. To understand the origin of the turtle-specific body plan, Nagashima et al. (p. 193; see the cover; see the Perspective by Rieppel) compared chicken, mouse, and the Chinese soft shelled-turtle, Pelodiscus sinensis. Modern embryos were studied via whole-mount immunostaining, three-dimensional reconstructions, and with markers for early skeletal precursors and compared with previously reported fossils. Initially, embryos of the three animals share a common developmental pattern, one that is likely to have been shared with their last common ancestor. This pattern, however, is modified in the turtle by a specific folding of its body wall during embryogenesis. This folding preserves some of the connectivity between skeletal and muscle elements but also produces new connections. The turtle body plan, unique among amniotes, is based on the folding of an ancestral pattern during embryogenesis. The turtle shell offers a fascinating case study of vertebrate evolution, based on the modification of a common body plan. The carapace is formed from ribs, which encapsulate the scapula; this stands in contrast to the typical amniote body plan and serves as a key to understanding turtle evolution. Comparative analyses of musculoskeletal development between the Chinese soft-shelled turtle and other amniotes revealed that initial turtle development conforms to the amniote pattern; however, during embryogenesis, lateral rib growth results in a shift of elements. In addition, some limb muscles establish new turtle-specific attachments associated with carapace formation. We propose that the evolutionary origin of the turtle body plan results from heterotopy based on folding and novel connectivities.