Morphological integration and serial homology: A case study of the cranium and anterior vertebrae in salamanders

Serial homology or the repetition of equivalent developmental units and their derivatives is a phenomenon encountered in a variety of organisms, with the vertebrate axial skeleton as one of the most notable examples. Serially homologous structures can be viewed as an appropriate model system for studying morphological integration and modularity, due to the strong impact of development on their covariation. Here, we explored the pattern of morphological integration of the cranium and the first three serially homologous structures (atlas, first, and second trunk vertebrae) in salamandrid salamanders, using micro‐CT scanning and three‐dimensional geometric morphometrics. We explored the integration between structures at static and evolutionary levels. Effects of allometry on patterns of modularity were also taken into account. At the static level (within species), we analyzed inter‐individual variation in shape to detect functional modules and intra‐individual variation to detect developmental modules. Significant integration (based on inter‐individual variation) among all structures was detected and allometry is shown to be an important integrating factor. The pattern of intra‐individual, asymmetric variation indicates statistically significant developmental integration between the cranium and the atlas and between the first two trunk vertebrae. At the evolutionary level (among species), the cranium, atlas, and trunk vertebrae separate as different modules. Our results show that morphological integration at the evolutionary level coincides with morphological and functional differentiation of the axial skeleton, allowing the more or less independent evolutionary changes of the cranial skeleton and the vertebral column, regardless of the relatively strong integration at the static level. The observed patterns of morphological integration differ across levels, indicating different impacts of developmental and phylogenetic constraints and functional demands. This study explored how development, function, and evolution affect the patterns of morphological integration of serially homologous structures. The strength of morphological integration between cranium, atlas, first and second trunk vertebrae in salamandrids was quantified at multiple levels. Patterns of integration between adjacent and non‐adjacent elements and their strength differed at the static (on symmetric and asymmetric component of shape variation) and evolutionary levels. Our results indicate diff