Multicellular filaments from the ca. 1200-Ma Hunting Formation (Somerset Island, arctic Canada) are identified as bangiacean red algae on the basis of diagnostic cell-division patterns. As the oldest taxonomically resolved eukaryote on recordBangiomorpha pubescensn. gen. n. sp. provides a key datum point for constraining protistan phylogeny. Combined with an increasingly resolved record of other Proterozoic eukaryotes, these fossils mark the onset of a major protistan radiation near the Mesoproterozoic/Neoproterozoic boundary.Differential spore/gamete formation showsBangiomorpha pubescensto have been sexually reproducing, the oldest reported occurrence in the fossil record. Sex was critical for the subsequent success of eukaryotes, not so much for the advantages of genetic recombination, but because it allowed for complex multicellularity. The selective advantages of complex multicellularity are considered sufficient for it to have arisen immediately following the appearance of sexual reproduction. As such, the most reliable proxy for the first appearance of sex will be the first stratigraphic occurrence of complex multicellularity.Bangiomorpha pubescensis the first occurrence of complex multicellularity in the fossil record. A differentiated basal holdfast structure allowed for positive substrate attachment and thus the selective advantages of vertical orientation; i.e., an early example of ecological tiering. More generally, eukaryotic multicellularity is the innovation that established organismal morphology as a significant factor in the evolutionary process. As complex eukaryotes modified, and created entirely novel, environments, their inherent capacity for reciprocalmorphologicaladaptation, gave rise to the “biological environment” of directional evolution and “progress.” The evolution of sex, as a proximal cause of complex multicellularity, may thus account for the Mesoproterozoic/Neoproterozoic radiation of eukaryotes.