Mechanisms for the evolution of complex and diversely elaborated sepals in Iris identified by comparative analysis of...

Crested sepals, which have evolved at least five times in Iris, are adaxially elaborated with a sinuous and/or uneven median structure (crest) along the proximal-distal axis and sometimes with various lateral structures (ridges, crests, and linear protuberances) flanking the median crest. These structural elaborations are complex yet diverse in form, providing a good opportunity to investigate developmental mechanisms for the diversification of reproductive lateral organs.

Morphologies of the median and lateral structures at different developmental stages from selected crested sepals representing the major types of structural elaborations were recorded using scanning electron microscopy and light microscopy. Developmental (morphogenetic) events that contribute to changes in shape (e.g., sinuousness, unevenness) between consecutive stages were recorded. Developmental sequences-trajectories that consist of a series of developmental events-were compared in a phylogenetic context.

Three developmental events (development of outgrowths, greater expansion of the upper zone, and greater expansion of the lower zone), are shared across lineages, occur in the same developmental sequences, and are responsible for the changes in shape during the development of diverse structural elaborations. In addition, two novel developmental events and the development of trichomes on elaborate structures were observed within the core-crested clade.

Developmental sequences are conserved across independently evolved crested lineages. Heterochronic and heterotopic shifts of developmental events play the major role in the diversification of elaborations of crested sepals in Iris. The evolution of novel developmental events and the development of trichomes also contribute to the diversity.