How does complex social behavior evolve? What are the developmental building blocks of division of labor and specialization, the hallmarks of insect societies? Studies have revealed the developmental origins in the evolution of division of labor and specialization in foraging worker honeybees, the hallmarks of complex insect societies. Selective breeding for a single social trait, the amount of surplus pollen stored in the nest (pollen hoarding) revealed a phenotypic architecture of correlated traits at multiple levels of biological organization in facultatively sterile female worker honeybees. Verification of this phenotypic architecture in "wild-type" bees provided strong support for a "pollen foraging syndrome" that involves increased senso-motor responses, motor activity, associative learning, reproductive status, and rates of behavioral development, as well as foraging behavior. This set of traits guided further research into reproductive regulatory systems that were co-opted by natural selection during the evolution of social behavior. Division of labor, characterized by changes in the tasks performed by bees, as they age, is controlled by hormones linked to ovary development. Foraging specialization on nectar and pollen results also from different reproductive states of bees where nectar foragers engage in pre-reproductive behavior, foraging for nectar for self-maintenance, while pollen foragers perform foraging tasks associated with reproduction and maternal care, collecting protein.