Morpho Evo-Devo of the Gynoecium: Heterotopy, Redefinition of the Carpel, and a Topographic Approach

Since the 19th century, we have had countless debates, sometimes acrimonious, about the nature of the gynoecium. A pivotal question has been whether all angiosperms possess carpels or if some or all angiosperms are acarpellate. We can resolve these debates if we do not define the carpel as a closed megasporophyll but simply as an appendage that encloses the placenta or a single ovule. This redefinition may, however, lead to confusion because often it may not be clear whether the traditional (classical) definition of the carpel or the redefinition is implied. Therefore, a topographic approach is proposed that is compatible with the redefinition. According to this approach, gynoecia comprise one or more gynoecial appendages and placentas or single ovules that may be formed in different positions. Heterotopy refers to these different positions. In the context of evo-devo, which explores evolutionary changes in development, morpho evo-devo delves into spatial shifts of the placentas and ovules leading to heterotopy. Furthermore, it considers shifts in timing (heterochrony) and other processes leading to heteromorphy. Recognizing spatial shifting of the placentas or a single ovule and other evolutionary processes opens up new vistas in the search for the ancestor(s) of angiosperms and their gynoecia.