Comparative floral ontogenesis among Persoonioideae including Bellendena (Proteaceae)

Floral ontogeny is described and compared in five species and four genera of the hypothetically basal proteaceous subfamily Persoonioideae sensu Johnson and Briggs. The hypotheses surrounding the origin of the peculiar proteaceous flower and homologous structures within the flowers are examined using ontogenetic morphological techniques. Ontogenetic evidence reveals that the proteaceous flower is simple, composed of four tepals, each tepal initiated successively with the lateral tepals being initiated first and second followed by the successive initiation of the sagittal tepals. Each of four stamens is initiated opposite a tepal in a similar sequence to tepal initiation. A single carpel develops terminally from the remaining floral meristem. In taxa of Persoonieae, nectaries are initiated from a broadened receptacle in alternistamenous sites after zonal growth beneath and between the tepals and stamens has begun. The nectaries are interpreted as secondary organs, not reduced homologues of a "lost" petal or stamen series. Developmental variation is present among the examined taxa in several forms including the development of a Vorlauferspitze (spine) on the upper portion of the tepals, adnation between the anthers and tepals, and formation of the carpel. In Placospermum the early formation of the carpel cleft extends to the floral receptacle and in the other taxa, the carpel cleft is distinctly above the receptacle. Different developmental pathways result in similar mature morphologies of the carpel in Persoonia falcata and Placospermum coriaceum. Bellendena montana is unique relative to the other taxa in having free stamens, a punctate stigma, reduced (not lost) floral bracts, and the floral and bract primordia are initiated from a common meristem. This study provides a foundation for future studies of the developmental basis of floral diversity within Proteaceae.