Invasion of new adaptive zones retains telltale signs of directional selection at macroevolutionary scales in mammals

Directional selection is often viewed as a transient force in macroevolution, with its signal eroded over time by stabilizing and fluctuating selection. Yet, transitions into new adaptive zones are predicted to impose strong and sustained selective pressures that may leave a detectable signature even across deep timescales. We test this prediction by comparing the rates of multivariate skull morphological evolution required to traverse the boundaries between adaptive zones against genetic drift expectations. Our dataset includes 11,793 specimens spanning 231 species from 12 mammalian clades, each containing unique ecological transitions into new adaptive zones. Using a quantitative genetics framework, we estimated the phenotypic distances between ancestral and derived adaptive zones and contrasted them with null expectations under genetic drift. While a few adaptive zone invasions (e.g., marsupials and rodents) are consistent with drift, most exhibit substantially elevated rates of evolution. These results suggest that directional selection has recurrently shaped mammalian cranial evolution during major ecological shifts. We propose that adaptive zone transitions represent evolutionary contexts in which adaptation leaves a persistent macroevolutionary signal, challenging the prevailing view that long-term patterns are dominated by static forces.