Patterns of Floral Structure and Orientation in Japonolirion , Narthecium, and Tofieldia

Floral evolution requires reassessment in basal monocots, including species formerly assigned to Melanthiaceae, in the light of recent developments in the molecular phylogenetics of monocots. We have investigated flowers of Tofieldia (Tofieldiaceae), Japonolirion (Petrosaviaceae), and Narthecium (Nartheciaceae). We confirm Engler's (1888) hypothesis that orientation of lateral flowers in monocots is dependent on presence and position of additional phyllomes on the pedicel. The type of floral orientation that occurs in Tofieldia is unusual for monocots, since the additional phyllomes are represented by calyculus scales rather than a bracteole, and the outer whorl tepals are initiated alternating with the calyculus scales. In Japonolirion and Narthecium, a bracteole is inserted in an adaxialtransverse or transverse position; either the outer median tepa! is adaxial or no single tepa! is inserted in the median position. In Tofieldia, the pedicel has a calyculus of an abaxial and two adaxialtransverse phyllomes; the outer median tepa! is adaxial. Additional phyllomes on the pedicel are not adaxial, in contrast to adaxial prophylls in the vegetative regions. The presence or absence of a bracteole or calyculus is taxonomically important. Tofieldia pusilla differs from the other species of Tofieldia examined in the absence of a flower-subtending bract, but the calyculus demonstrates some bract-like features in position, structure and development, which can be interpreted as a hybridization of developmental pathways. The abaxial calyculus scale of T. coccinea is delayed in development.