DEFH28 is a novel MADS-box gene from Antirrhinum majus. Phylogenetic reconstruction indicates that it belongs to the SQUA-subfamily of MADS-box genes. Based on its expression pattern and the phenotype of transgenic plants it is predicted that DEFH28 exerts a dual function during flower development, namely control of meristem identity and fruit development. Firstly, DEFH28 is expressed in the inflorescence apical meristem and might control, together with SQUAMOSA (SQUA), floral meristem identity in Antirrhinum. Also, DEFH28 is sufficient to switch inflorescence shoot meristem to a floral fate in transgenic Arabidopsis thaliana plants. Secondly, DEFH28 is expressed in carpel walls, where it may regulate carpel wall differentiation and fruit maturation. Support for this later role comes from overexpression of DEFH28 throughout the silique in transgenic Arabidopsis plants where it altered the identity of the replum and valve margin cells so that they adopted a valve cell identity. This late aspect of the DEFH28 function is identical to the FRUITFULL (FUL) function of Arabidopsis as demonstrated in gain-of-function plants. FUL, like DEFH28, belongs to the SQUA-subfamily of MADS-box genes. DEFH28 most likely represents the ortholog of FUL. Promoter analysis shows that the control mechanism conferring a carpel wall specific expression has been conserved between Antirrhinum and Arabidopsis during evolution. Although the overall flower development between Antirrhinum and Arabidopsis is very similar, their carpels mature into different types of fruits: capsules and siliques, respectively. Therefore, it is suggested that the role of DEFH28 in control of carpel wall differentiation reflects a conserved molecular mechanism integrated into two very different carpel developmental pathways.