Constraint and Allometric Diversification in a Simplified Neck: Shape Evolution of the Atlas in Hyloidea (Anura)

The anuran atlas represents one of the most structurally simplified cervical systems among tetrapods, consisting of a single vertebra that must simultaneously support the head, permit mobility, and withstand mechanical loads generated during locomotion and feeding. This unique configuration provides a compelling framework for investigating how structural constraint, allometry, and ecology interact to shape morphological evolution. Here, we examine atlas shape diversification across Hyloidea using geometric morphometrics and phylogenetic comparative methods on 421 specimens representing 133 species. We quantified shape variations in dorsal and ventral views and tested the relative contributions of body size, phylogenetic history, microhabitat, and locomotor mode to atlas morphology. Atlas shape exhibited significant phylogenetic signals and was predominantly structured by allometric scaling, with the atlas size (centroid size) explaining a significant proportion of variation in both anatomical views, particularly ventrally. Morphological differences were concentrated in atlas width and relative proportions, indicating that diversification occurs primarily through size-dependent remodeling rather than through major structural reorganization. In contrast, ecological variables explained little independent variation in atlas shape, although microhabitat modulated allometric trajectories in dorsal and ventral views. Together, these results indicate that atlas evolution in hyloid frogs is governed chiefly by size-related biomechanical demands operating within a phylogenetically structured and developmentally constrained system. More broadly, our findings suggest that extreme cervical simplification channels morphological diversification toward allometric modification, limiting the extent to which ecological specialization can drive anatomical divergence.