Flower structure and development in Pennantiaceae: uncovering diversity of pseudomonomerous gynoecia in the basal grade of the order Apiales

Pseudomonomerous gynoecia with three (or four) carpels are unknown in the species-rich core group of Apiales, but this condition is shared by three species-poor families (Pennantiaceae, Torricelliaceae, Griseliniaceae) that form the basal grade of the order. Testing a hypothesis on the ancestral nature of carpel dimorphism in Apiales requires comparative data for all three lineages in this grade. We provide the first detailed description of flowers, including floral vasculature and gynoecium development, in a member of Pennantiaceae (Pennantia corymbosa). In contrast to many other Apiales, the inflorescence of Pennantia is paniculate and therefore has an unstable number of phyllomes in axes terminated by flowers. All phyllomes in the inflorescence are shifted onto lateral branches they subtend exhibiting recaulescence, a pattern that has not been reported elsewhere in Apiales. Plants are dioecious with functionally unisexual flowers. There are normally five stamens alternating with five petals. Anthers are present and produce pollen in stamens of male as well as female flowers, but ventral microsporangia are reduced in some anthers of female flowers. Anther morphology sometimes varies even among stamens of the same flower. Two types of synthecal anthers are recorded. Pollen dimorphism is confirmed: inaperturate pollen produced by stamens of female flowers supposedly acts as the only reward for pollinators in the absence of nectaries. The gynoecium of the female flower is syncarpous and pseudomonomerous: only one of three carpels is fertile. The gynoecium is initiated as three carpel primordia (future stigmas). One of them is smaller than the other two and occupies an alternistaminal (and antepetalous) position. The two large carpel primordia are located in the radii of stamens that are generally smaller (early in development) than the three other stamens. The carpel dimorphism is maintained at anthesis. The carpel with the smaller stigma is fertile, and those with larger stigmas are sterile. The carpels are congenitally united below the stigmas. The ovary is superior, unilocular (vs. inferior and plurilocular in Torricelliaceae and Griseliniaceae) and usually uniovulate with pendent ovule(s) inserted at the cross-zone level of the fertile carpel. As in most other Apiales, the short symplicate zone is sealed by postgenital fusion at anthesis and forms an internal compitum. The fertile carpel of the members of the basal grade of Apiales investigated so far is uniformly arranged in a petal radius. This is consistent with the idea that pseudomonomery is associated with stable patterns of flower groundplan in Apiales. Our data do not provide any clear structural or developmental evidence of independent origins of carpel dimorphism in Pennantiaceae, Torricelliaceae and Griseliniaceae.