To disentangle the mechanisms producing the biological diversity seen in nature, ecologists increasingly seek to integrate ecology and evolution (Jetz et al., 2012;McGill et al., 2019;Wiens & Donoghue, 2004). Mapping traits onto phylogenies is essential for such integration, as mapping traits reveals the rates and tempo of evolution of behavioral, morphological, and ecological characteristics (Bollback, 2006). Knowledge of trait evolution has often been applied to evaluate the evolutionary mechanisms producing, for example, the appearance of ecological innovations and the bursts behind evo-