A new gnetalean macrofossil from the Early Cretaceous and its evolutionary significance

Modern gnetophytes consist of three families, i.e. the Ephedraceae, the Gnetaceae, and the Welwitschiaceae, which show diversified morphology. The basal group Ephedraceae possesses an extremely reduced female cone with only the uppermost pair/whorl being fertile, while both the Gnetaceae and the Welwitschiaceae bear female cones/spikes with multiple whorls of fertile bracts. Here, we describe a new ephedroid macrofossil, Liaoningia decussata Yang et Lin, gen. et sp. nov., from the Lower Cretaceous Yixian Formation of western Liaoning in China. This species has typical ephedroid morphology, e.g. the dichasial branching shoot system, swollen nodes, internodes having many fine longitudinal striations, and decussate phyllotaxy, lengthy linear leaves possessing two parallel veins, and female cones consisting of paired bracts. This new genus is similar to the Ephedraceae in its vegetative organs, but differs from the latter by the female spikes with multiple pairs of fertile bracts. In this respect Liaoningia is similar to both the Gnetaceae and the Welwitschiaceae, but distinctly differs from the latter two genera in its vegetative organs. As a result, our new genus shows intermediate morphology between the Ephedaceae and the other two families of the gnetophytes, and is probably a missing link in the evolutionary process from the Ephedraceae to the clade including the Gnetaceae and the Welwitschiaceae.