Understanding the pattern of speciation in a group of plants is critical for understanding its morphological evolution. Lepidium is the genus with the largest variation in floral structure in Brassicaceae, a family in which the floral ground plan is remarkably stable. However, flowers in more than half of Lepidium species have reduced stamen numbers, and most of these also have reduced petals. The species with reduced flowers are geographically biased, distributed mostly in the Americas and Australia͞ New Zealand. Previous phylogenetic studies using noncoding regions of chloroplast DNA and rDNA internal transcribed spacer were incongruent in most New World species relationships. These data, combined with the presence of many polyploid Lepidium species, implied a reticulate history of the genus but did not provide enough information to infer the evolutionary pattern of flower structures. To address this question more thoroughly, sequences of the first intron of a single copy nuclear gene, PISTILLATA, were determined from 43 species. Phylogenetic analysis of the PI intron suggests that many species in the New World have originated from allopolyploidization, and that this is correlated with floral reduction. Interspecific hybrids were generated to understand why allopolyploidization is associated with reduced flowers. The phenotypes of F1 flowers indicate allelic dominance of the absence of lateral stamens, suggesting that propagation of dominant alleles through interspecific hybridization could account for the abundance of the allopolyploid species without lateral stamens.T hrough the study of model plant species, it is relatively well understood how organ identity is regulated in flowers (1-3). However, the underlying genetic mechanisms that lead to the remarkable variation in flower structures in angiosperms are not known. One way to address this question is to analyze floral variation in closely related species. Knowledge of the genetic bases of phenotypic variation between related species for which we know the phylogeny should lead to an understanding of the number of genetic changes underlying phenotypic evolution and other events that lead to the dispersal of certain phenotype(s).The basic floral ground plan of Brassicaceae is well conserved, with four sepals, four petals, six stamens, and two carpels (Fig. 1A). However, within Lepidium L. (Ϸ175 species worldwide), more than half of the species have only two or four stamens (Fig.