Anatomical network analyses reveal evolutionary integration and modularity in the lizards skull.

The morphology of lizard skulls is highly diverse, and it is crucial to understand the factors that constrain and promote their evolution to understand how lizards thrive. The results of interactions between cranial bones reflecting these factors can be detected as integration and modularity, and the analysis of integration and modularity allows us to explore the underlying factors. In this study, the integration and modularity of the skulls of lizards and the outgroup tuatara are analyzed using a new method, Anatomical Network Analysis (AnNA), and the factors causing lizards morphological diversity are investigated by comparing them. The comparison of modular structures shows that lizard skulls have high integration and anisomerism, some differences but basically common modular patterns. In contrast, the tuatara shows a different modular pattern from lizards. In addition, the presence of the postorbital bar by jugal and postorbital (postorbitofrontal) also reflect various functional factors by maintaining low integration. The maintenance of basic structures due to basic functional requirements and changes in integration within the modules play a significant role in increasing the morphological diversity of the lizard skull and in the prosperity of the lizards.