Fig. 1. Modern snakes originated as burrowers, based on their inner ear morphology.

(A) Snake skulls in right lateral view, showing that the inner ear (orange) locates inside the braincase and opens to the stapes (blue) in the middle ear. Ear and skull models are not to scale. (B) Inner ear of Laticauda colubrina, an aquatic species. (C) Ptyas mucosa, terrestrial generalist. (D) Xenopeltis unicolor, a burrowing species. (E) Hypothetical ancestor of crown snakes, predicted as burrowing with 70.1% probability. (F) D. patagonica, predicted as burrowing with 93.4% probability. (G) Phylogeny of all snakes and lizards in this study, adapted from Gauthier et al. (8), Pyron et al. (19), and Yi and Norell (20).