The large brain and small postcanine teeth of modern humans are among our most distinctive features, and trends in their evolution are well studied within the hominin clade. Classic accounts hypothesize that larger brains and smaller teeth coevolved because behavioral changes associated with increased brain size allowed a subsequent dental reduction. However, recent studies have found mismatches between trends in brain enlargement and posterior tooth size reduction in some hominin species. We use a multiple-variance Brownian motion approach in association with evolutionary simulations to measure the tempo and mode of the evolution of endocranial and dental size and shape within the hominin clade. We show that hominin postcanine teeth have evolved at a relatively consistent neutral rate, whereas brain size evolved at comparatively more heterogeneous rates that cannot be explained by a neutral model, with rapid pulses in the branches leading to later Homo species. Brain reorganization shows evidence of elevated rates only much later in hominin evolution, suggesting that fast-evolving traits such as the acquisition of a globular shape may be the result of direct or indirect selection for functional or structural traits typical of modern humans.endocast | postcanine teeth | evolutionary rates | selection | paleoanthropology I n comparison with other hominins, modern humans are characterized by their large brain and small posterior teeth. These traits are among our most distinctive features, and trends in their evolution are well studied because of the phylogenetic and functional implications of variation in dental and cerebral anatomy (1-3). Brain expansion and postcanine reduction appear to follow parallel trends during hominin evolution, and classic views consider that an increase in brain size was linked to more complex behavior that included the manufacture and use of stone tools, which allowed a subsequent dental reduction. A shift toward a higher-quality diet during the evolution of early Homo also has been related to brain size increase and posterior tooth reduction (4, 5). However, it has been suggested recently that brain expansion in early Homo, as inferred from endocranial volume, substantially preceded dental reduction (6). It also has been noted that early in the Neanderthal lineage strong dental reduction preceded the additional brain expansion seen in the later "classic" Neanderthals (7). The suggestion that stone tool use and manufacture substantially predated the increase in brain size observed in early Homo (8) adds further complexity to this scenario.Recent developments in ancestral state reconstruction (9, 10) allow lineage-specific patterns of brain expansion and dental reduction to be quantified and compared. Unlike traditional approaches to ancestral state reconstruction that assume a neutral evolutionary scenario, which is likely unrealistic in most cases, we used a variable rate approach that estimates differences in evolutionary rates across different branches of a given phylogeny. We appli...