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The relative complexity of floral structures, their relative timing of evolution, and the number and size of fossils required to accurately reconstruct their evolution, are affected by the evolutionary history of the lineage under study. Small changes in the evolution of a trait can have significant effects on the inference of its evolution. For example, it is thought that the disparity of extant angiosperms is significantly underestimated because of the contribution of Amborella, the only living representative of this group30. Additionally, a lack of rigorous methods for inferring ancestral states in the absence of fossil representatives affects the inferences made from analysis of extant taxa. We are using rigorous methods, and the inferred ancestral flower is robust to varying assumptions on the putative ancestral state of additional characters (for example, the sexual system). Our inference of the four key innovations in floral morphology across the angiosperm phylogeny is also robust. However, we note that alternative interpretations of the ancestral flower and of the evolution of this trait are possible (see Supplementary Discussion).
We find that the common ancestor of all extant angiosperms had a gynoecium of more than five carpels, with four to five spirally arranged superior carpels and one inferior carpel. We infer that at least four carpels are necessary for the ancestral gynoecium in order to allow for the differentiation of a central style and a gynostegium. Evolutionary transitions to novel bilateral symmetry in the ancestral flower and the lateral petals of Amborella, which may have been important precursors to the functional differentiation of the labellum, are not considered in this study.
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