Compound pollen cone in a Paleozoic conifer

A rich fossil biota from a Pennsylvanian age deposit of eastern North America contains numerous vegetative and fertile specimens that conform to a single species of primitive walchian conifers. Among the specimens is a compound pollen cone that comprises closely spaced, helically arranged, leaf-like bracts with axillary dwarf shoots. The specimen looks superficially similar to an ultimate vegetative conifer shoot, but there are small appendages in the axil of each bract that represent the fertile dwarf shoots. Dwarf shoots consist of an axis that bears sterile scales and sporophylls with erect pollen sacs. Pollen found in the sacs is monosaccate and conforms to the sporae dispersae genus Potonicisporites Bhardwaj. This cone is a compound shoot system that is morphologically equivalent to the ovulate cones of conifers and to the pollen cones of Paleozoic cordaitaleans and modern gnetophytes. Therefore, it is fundamentally different from the simple pollen cones of other fossil and modern conifers. Discovery of this specimen unexpectedly supports molecular studies that predict a close relationship between Coniferales and Gnetales, and provides fossil evidence to help reconcile the discordant phylogenetic hypotheses of seed plant systematics that have been developed from morphological and molecular data.