Phenotypic traits are often integrated into evolutionary modules: sets of organismal parts that evolve together. In social insect colonies, the concepts of integration and modularity apply to sets of traits both within and among functionally and phenotypically differentiated castes. On macroevolutionary timescales, patterns of integration and modularity within and across castes can be clues to the selective and ecological factors shaping their evolution and diversification. We develop a set of hypotheses describing contrasting patterns of worker integration and apply this framework in a broad (246 species) comparative analysis of major and minor worker evolution in the hyperdiverse ant genus Pheidole. Using geometric morphometrics in a phylogenetic framework, we inferred fast and tightly integrated evolution of mesosoma shape between major and minor workers, but slower and more independent evolution of head shape between the two worker castes. Thus, Pheidole workers are evolving as a mixture of intracaste and intercaste integration and rate heterogeneity. The decoupling of homologous traits across worker castes may represent an important process facilitating the rise of social complexity.

27Phenotypic traits are often integrated into evolutionary modules: sets of organismal parts that 28 evolve together. In social insect colonies the concepts of integration and modularity apply to sets 29 of traits both within and among functionally and phenotypically differentiated castes. On 30 macroevolutionary timescales, patterns of integration and modularity within and across castes 31 can be clues to the selective and ecological factors shaping their evolution and diversification. 32We develop a set of hypotheses describing contrasting patterns of worker integration and apply 33 this framework in a broad (246 species) comparative analysis of major and minor worker 34 evolution in the hyperdiverse ant genus Pheidole. Using geometric morphometrics in a 35 phylogenetic framework, we inferred fast and tightly integrated evolution of mesosoma shape 36 between major and minor workers, but slower and more independent evolution of head shape 37 between the two worker castes. Thus, Pheidole workers are evolving as a mixture of intra-and 38 inter-caste integration and rate heterogeneity. The decoupling of homologous traits across worker 39 castes may represent an important process facilitating the rise of social complexity. 40 41