Reduction, reorganization and stasis in the evolution of turtle shell elements

Abstract

Novel phenotypic configurations can profoundly alter the evolutionary trajectories of species. Although innovation can precede lengthy periods of evolutionary stasis, the potential for species to diversify further can be realized via modular changes across distinct levels of hierarchical organization. To test this expectation, we undertook anatomical network analyses to model the organization and composition of the turtle's shell. Our results suggest that stem turtles featured the greatest diversity in the number of skeletal (bones) and epidermal (scutes) shell elements. The shell subsequently underwent numerical simplification. Thus, the sum of potential connections (links) in shell networks has diminished in modern turtles. Some network system descriptors of complexity, integration and modularity covaried with the number of network components (nodes), which has remained evolutionarily stable since the Jurassic. We also demonstrated that shell reorganization might be feasible within modular subdivisions, particularly in modern turtles with simplified and less integrated network structures. We discuss how these findings align with previous studies on numerical simplification with enhanced skeletal specialization in the tetrapod skull. Altogether, our analyses expose the evolvability of the turtle's shell and bolster the foundation for further macroevolutionary comparisons of ancient and modern species.