Arendt, D.,Musser, J. M.,Baker, C. V. H.,Bergman, A.,Cepko, C.,Erwin, D. H.,Pavlicev, M.,Schlosser, G.,Widder, S.,Laubichler, M. D.,Wagner, G. P.

Cell types are the basic building blocks of multicellular organisms and are extensively diversified in animals. Despite recent advances in characterizing cell types, classification schemes remain ambiguous. We propose an evolutionary definition of a cell type that allows cell types to be delineated and compared within and between species. Key to cell type identity are evolutionary changes in the ‘core regulatory complex’ (CoRC) of transcription factors, that make emergent sister cell types distinct, enable their independent evolution and regulate cell type-specific traits termed apomeres. We discuss the distinction between developmental and evolutionary lineages, and present a roadmap for future research.