In most angiosperms, floral organs are acropetally initiated, i.e., from the perianth upwards to the gynoecium. The review surveys examples of deviation from this typical pattern that takes place in both oligomerous and polymerous whorled flowers. The plants displaying the same non-acropetal pattern of floral development are not necessarily closely related and thus similarities in their floral structure and development should be regarded as convergences. Vice versa, representatives of the same family often show different patterns of initiation of floral organs. Flowers with the same groundplan can demonstrate either typical acropetal or non-acropetal developmental pattern. In other words, evolution of patterns of floral development is relatively homoplastic. Presumably, the repeated transitions from acropetal to non-acropetal developmental patterns (and back) readily occurred in evolution and were of a saltational nature.
The phenomenon of heterochrony, or shifts in the relative timing of ontogenetic events, is important for understanding many aspects of plant evolution, including applied issues such as crop yield. In this paper, we review heterochronic shifts in the evolution of an important floral organ, the carpel. The carpels, being ovule-bearing organs, facilitate fertilisation, seed, and fruit formation. It is the carpel that provides the key character of flowering plants, angiospermy. In many angiosperms, a carpel has two zones: proximal ascidiate and distal plicate. When carpels are free (apocarpous gynoecium), the plicate zone has a ventral slit where carpel margins meet and fuse during ontogeny; the ascidiate zone is sac-like from inception and has no ventral slit. When carpels are united in a syncarpous gynoecium, a synascidiate zone has as many locules as carpels, whereas a symplicate zone is unilocular, at least early in ontogeny. In ontogeny, either the (syn)ascidiate or (sym)plicate zone is first to initiate. The two developmental patterns are called early and late peltation, respectively. In extreme cases, either the (sym)plicate or (syn)ascidiate zone is completely lacking. Here, we discuss the diversity of carpel structure and development in a well-defined clade of angiosperms, the monocotyledons. We conclude that the common ancestor of monocots had carpels with both zones and late peltation. This result was found irrespective of the use of the plastid or nuclear phylogeny. Early peltation generally correlates with ovules belonging to the (syn)ascidiate zone, whereas late peltation is found mostly in monocots with a fertile (sym)plicate zone.
The basalmost monocot genus Acorus is well-known for its use in traditional oriental medicine. It comprises the groups of A. calamus and A. gramineus. A recent study recognized three species in the latter group, A. gramineus, A. macrospadiceus, and A. tatarinowii. The material currently known as A. tatarinowii has been extensively studied as a source of various chemical compounds and for producing the first published genome of Acorus, which is important for understanding the origin and evolution of monocots. Using the data from morphology, anatomy, and biogeography, we argue that the type material of A. tatarinowii does not match the interpretation of the species name as adopted in the current literature and herbarium collections (to a taxon of the A. gramineus group from Southeast Asia) but rather belongs to the A. calamus group. Moreover, the name A. macrospadiceus also cannot be used because it was invalidly published. Under a narrow species concept, other appropriate species names should be found or proposed for the plants currently named A. tatarinowii and A. macrospadiceus. However, we discourage the use of a narrow species concept in the A. gramineus group as insufficiently justified and suggest recognizing a single polymorphic species, A. gramineus s.l., at least until a comprehensive taxonomic revision of the group is available. Apart from the presentation of our revised taxonomic framework, we update the geographical distributions of Acorus species in Vietnam, Laos, and Thailand.