Abstract: The fossil record documents two mutually exclusive macroevolutionary modes separated by the transitional Ediacaran Period. Despite the early appearance of crown eukaryotes and an at least partially oxygenated atmosphere, the pre-Ediacaran biosphere was populated almost exclusively by microscopic organisms exhibiting low diversity, no biogeographical partitioning and profound morphological/evolutionary stasis. By contrast, the post-Ediacaran biosphere is characterized by large diverse organisms, bioprovinciality and conspicuously dynamic macroevolution. The difference can be understood in terms of the unique escalatory coevolution accompanying the early Ediacaran introduction of eumetazoans, followed by their early Cambrian (Tommotian) expansion into the pelagic realm. Eumetazoans reinvented the rules of macroecology through their invention of multitrophic food webs, large body size, life-history trade-offs, ecological succession, biogeography, major increases in standing biomass, eukaryote-dominated phytoplankton and the potential for mass extinction. Both the pre-Ediacaran and the post-Ediacaran biospheres were inherently stable, but the former derived from the simplicity of superabundant microbes exposed to essentially static, physical environments, whereas the latter is based on eumetazoan-induced diversity and dynamic, biological environments. The c. 100-myr Ediacaran transition (extending to the base of the Tommotian) can be defined on evolutionary criteria, and might usefully be incorporated into the Phanerozoic.