The utility of maintaining distinct macroevolutionary and microevolutionary theory has long been debated. Although population and quantitative genetics provide an extensive list of microevolutionary forces that might explain macroevolutionary trends, studies of these processes are temporally limited and may fail to fully explain macroevolutionary patterns. To understand the relationship between the macroevolutionary pattern and microevolutionary forces, we must first understand how different populations respond to a given novel environment over hundreds or even thousands of generations. This study details the tempo of fitness gain over 2000 generations in four replicate lineages from each of five different ancestral Escherichia coli clones. Adaptive tempo was measured in the evolved lineages and ancestry was a significant source of variation in that tempo. Microevolutionary theory suggests that adaptive tempo should be proportional to the distance from an optimum phenotype. Demographic fitness measures allowed estimation of the ancestral distance from an optimum in the present study. Ancestral distance from an optimum was significantly related to adaptive tempo but it did not account for all of the observed variation. This suggests the existence of both ancestor and clade specific constraints. Understanding the role of such constraints is critical to both microevolutionary and macroevolutionary theory. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 403–411.