Dataset and script: Nested modularity and integration unlock otolith diversity in sound-sensitive croaker fishes

Otoliths in bony fishes exhibit remarkable morphological diversity, closely linked to auditory and vestibular functions. In this study, we examined otolith shape using geometric morphometrics on 2D images from 135 species across 65 genera of Sciaenidae (croakers), a family of sound-producing fishes. Our analyses reveal that otolith morphology is governed by a hierarchical modular structure comprising nested modules that regulate the otolith contour, sulcus acusticus, ostium, cauda, and their subregions. Strong and differential morphological integration among modules contributes to morphological deviation across lineages, while the weak phylogenetic signal in modularity suggests that functional and developmental factors primarily drive otolith diversification. This repository includes: Landmark datasets and metadata used for geometric morphometric, modularity, integration, and phylogenetic analyses. MATLAB and R scripts for modularity, integration, and phylogenetic tests. DNA and phylogenetic files used for phylogenetic analysis. All materials are provided to ensure full reproducibility of the analyses presented in the manuscript. See README.md for more details.