Adaptation is a fundamental process by which populations evolve to grow more fit in their environments. Recent studies are starting to show us that commensal microbes can evolve on short timescales of days and months, suggesting that ecological changes are not the only means by which microbes in complex natural populations respond to selection pressures. However, we still lack a complete understanding of the tempo and mode of adaptation in microbiomes given the many complex forces that natural populations experience, which include ecological pressures, changes in population size, spatial structure, and fluctuations in selection pressures. Advances in modeling complex populations and scenarios will allow us to understand adaptation not only in microbiomes but also more generically in other natural populations that experience similar complexities.