Studies in the Alismataceae. XII. Floral organogenesis in Damasonium alisma and Baldellia ranunculoides , and comparisons with Butomus umbellatus

Floral organogenesis of Damasonium alisma Mill. occurs at first in an alternating trimerous pattern typical of Alismataceae, with the formation of three sepals, then three bulges, corresponding to the CA (common perianth–androecium) primordia described in some other Alismataceae, alternating with the sepals. A petal is initiated on each bulge, and a pair of stamens is initiated either on or close to it. Three carpels are initiated in positions alternating with the petals and stamen pairs, and three further carpels then arise above and between the first three. At maturity and in fruit the carpels lie in a whorled arrangement. Floral development in Baldellia ranunculoides (L.) Parl. is identical up to the initiation of the six stamens. After this, six carpel primordia are formed alternating with the stamens, and further carpel primordia arise alternating with those previously formed. In Butomus, up to the initiation of the first six stamens, the general plan of development resembles that of the two Alismataceae. Three further whorls of organs arise in alternation: a whorl of three stamens arises over the stamen pairs followed by two whorls each of three carpel primordia. It is argued that the trimerous appearance of the whorl of sepals (or outer perianth in Butomus) arises de novo and represents a genuine expression of trimery. However, most of the subsequent features of development in these flowers can be seen as arising from phyllotactic mechanisms that cause new primordia to arise between and above pre-existing ones. Consequently the appearance of trimerous or hexamerous whorls above the first whorl of perianth does not represent a fundamental feature of development. The nature of variations in the positional relationships of inner perianth, stamen, and carpel primordia in various Alismataceae and Butomus strengthen the case that there is a significant developmental association between inner perianth members and associated pairs of stamens, which may be connected with the evolution of the flowers from pseudanthial structures.Key words: Baldellia, Butomus, Damasonium, Alismatidae, flower, organogenesis.