Floral development and anatomy of Pentadiplandra (Pentadiplandraceae):: a key genus in the identification of floral morphological trends in the core Brassicales

The development and anatomy of leaves and flowers have been investigated in the monotypic species Pentadiplandra brazzeana to discuss its position relative to the core Brassicales and to clarify floral evolutionary trends within the order. Young leaves are flanked by two large stipules that shrivel at anthesis. Young flowers develop as elliptical primordia on a flattened raceme. Sepal initiation is sequential, starting with the lateral sepals and ending with three median sepals. The petals and diplostemonous androecium arise as regular whorls. Five antesepalous carpels develop into a saccate gynoecium with axile placentation. Petals develop a cushion-like basal appendage covering the extrastaminal nectary. Several ontogenetic and anatomical data of the flower support the basal position of Pentadiplandra in the evolution of the core Brassicales and for the derivation of the order from pentamerous diplostemonous ancestors. A derivation of the tetramerous Capparaceae and Brassicaceae as well as hexamerous Tovariaceae is discussed. A number of synapomorphies are identified: sequential sepal initiation with laterals preceding median sepals, retarded petal growth, a (andro)gynophore, the extrastaminal. nectary, and stipulate leaves. The peculiar androecium of the Brassicaceae is caused by the strong median compression of the flower, leading to the loss and displacement of median stamens. The derivation of a parietal placenta from an axile placenta is supported. Evidence supports a close link with either Tovariaceae or Capparaceae, concomitant with macromolecular results.