In vertebrates, olfaction is mediated by several families of G protein-coupled receptors (GPCRs) including odorant receptors (ORs). In this study, we investigated the antiquity of OR genes by searching for amino acid motifs found in chordate ORs among the protein predictions from 12 nonchordate species. Our search uncovered a novel group of genes in the cnidarian Nematostella vectensis. Phylogenetic analysis that included representatives from the other major lineages of rhodopsin-like GPCRs showed that the cnidarian genes, the cephalochordate and vertebrate ORs, and a family of genes from the echinoderm, Strongylocentrotus purpuratus, form a monophyletic clade. The taxonomic distribution of these genes indicates that the formation of this clade and therefore the diversification of the rhodopsin-like GPCR family began at least 700 million years ago, prior to the divergence of cnidarians and bilaterians. ORs and other rhodopsin-like GPCRs have roles in cell migration, axon guidance, and neurite growth; therefore, duplication and divergence in this family may have played a key role in the evolution of cell type diversity (including the emergence of complex nervous systems) and in the evolution of metazoan body plan diversity.