Two major classes of genes directing flower development have so far been described: early activated genes regulating meristem identity and later acting genes controlling organ identity. Here, we show that the fimbriata (fim) gene acts between these two classes in a sequence of gene activation. The fim gene, originally described in 1930, was cloned by transposon tagging from Antirrhinum majus and encodes a product with no detectable homology to other proteins. Mutations in fim result in partial homeotic transformations of floral organs and in reduced determinacy of the meristem. Expression and function of fim depends on the activity of meristem identity genes, and fim in turn controls the spatial and temporal expression of organ identity genes. The pattern of fim expression defines a new domain of the floral meristem that changes with time in a complementary manner to those of the meristem identity gene floricaula and the organ identity gene plena.