Cone and ovule development in Cunninghamia and Taiwania (Cupressaceae sensu lato) and its significance for conifer evolution

We examined the early developmental stages of the seed cones and seeds of two conifer genera, Cunninghamia and Taiwania, using scanning electron microscope (SEM) images of freshly collected material. In recent similar studies, these two taxa were not described. The present paper aims to fill that gap. Both genera appear to have features crucial to the understanding of the evolution of the cupressaceous cone, characteristic of the families Cupressaceae and Taxodiaceae, and provide further evidence for the need to merge these families. These features are: the ovuliferous scale in Cunninghamia develops as a small lobe with each of three ovules; in Taiwania these lobes are absent, but a small ridge could be a vestige of them. In neither of these two genera does an ovuliferous scale develop to maturity and only limited intercalary growth transforms the bracts, of which only their width and final shape distinguishes them from sterile leaves. Thus, the bracts, not the ovuliferous scales, form the mature cone in these two genera. This trend is continued in more derived genera of Cupressaceae. Another key extant taxon that has helped to elucidate the evolution of this type of conifer cone is Sciadopitys; similar studies have already been done on this genus, and we compared our findings to them. We also considered certain fossil Mesozoic conifer cones, which shed further light on the evolution of the cupressaceous cone. The evidence from these various genera strongly indicates that recently reconstructed phylogenies of gymnosperms based on molecular evidence from extant taxa do not reflect the evolution that actually happened. Such studies need to take into account nonmolecular evidence, as detailed here.