Assessing the extent to which population subdivision during cladogenesis is necessary for long-term phenotypic evolution is of fundamental importance in a broad range of biological disciplines. Differentiating cladogenesis from anagenesis, defined as evolution within a species, has generally been hampered by dating precision, insufficient fossil data, and difficulties in establishing a direct link between morphological changes detectable in the fossil record and biological species. Here we quantify the relative frequencies of cladogenesis and anagenesis for macroperforate planktic Foraminifera, which arguably have the most complete fossil record currently available, to address this question. Analyzing this record in light of molecular evidence, while taking into account the precision of fossil dating techniques, we estimate that the fraction of speciation events attributable to anagenesis is <19% during the Cenozoic era (last 65 Myr) and <10% during the Neogene period (last 23 Myr). Our central conclusion-that cladogenesis is the predominant mode by which new planktic Foraminifera taxa become established at macroevolutionary time scales-differs markedly from the conclusion reached in a recent study based solely on fossil data. These disparate findings demonstrate that interpretations of macroevolutionary dynamics in the fossil record can be fundamentally altered in light of genetic evidence. anagenesis, which occurs within a species, and cladogenesis, which occurs during speciation and results in the subdivision of a species into two reproductively isolated, independently evolving taxa (1, 2). Distinguishing between these modes is essential to determining the role of population subdivision in evolutionary dynamics (3,4). This topic has received considerable attention in paleontology (1, 2), perhaps because the fossil record provides the only direct record of long-term morphological change (5), but its importance extends to other biological disciplines. For example, in population genetics, models only predict that phenotypic change is contingent upon or associated with the evolution of reproductive isolation under specific circumstances (3). Thus, a primary role for cladogenesis may highlight the importance of particular speciation modes (2), metapopulations dynamics (6), or adaptive peaks in fitness landscapes (7,8). In community ecology, both evolutionary modes may influence biodiversity, but the mechanisms differ: only cladogenesis results in an increase in diversity (9-11), but anagenesis may help maintain diversity if it results in niche differences that facilitate species coexistence (12, 13).Remarkably few studies have attempted to quantify the relative frequencies of anagenesis and cladogenesis for entire species assemblages (e.g., refs. 5, 14, 15). Addressing this question is challenging due to insufficient fossil data for most taxa and the difficulties in relating morphological change to genetic divergence and the evolution of reproductive isolation (16). Fossil data can only be used to distinguish anag...