Developmental genetic pathways involved in flower formation in model plants such as Arabidopsis and maize enable us to identify genes, gene families, and gene networks that are involved in the regulation of flower initiation, growth and differentiation. These genes can then function as “candidate genes” and their expression, function, and biochemical interactions can be explored in other lineages to determine if they provide a necessary and sufficient toolkit for the development of the flower. Likewise, a view to the fossil record can provide documentation of reproductive innovations occurring within gymnosperms or along the stem lineage leading to angiosperms, elucidating the transitions required for the evolution of the angiosperm flower from an ancestral reproductive strobilus. Here we discuss the origin and subsequent evolution in form of the flower, highlighting recent studies in paleobotany, morphology, evolution, and developmental genetics with the goal of outlining advances in our understanding of flower evolution.