Numerous permineralised, complex medullosan pollen organs are described from Middle and Upper Pennsylvanian sediments of North America. Together with a reinvestigation of type specimens for the previously described species, these provide the basis for fully characterising internal structure and external morphology, and for recognising structural homologies among all medullosan synangia. All medullosan pollen organs, simple and complex, can be interpreted as consisting of sporangial tubes that are embedded in parenchymatous and sclerenchymatous ground tissue, and that have vascular bundles within the parenchymatous tissue. The sporangial tubes are arranged in a single series, usually a ring, with parenchymatous ground tissues and vascular bundles to the outside of the ring and sclerenchymatous ground tissue to the inside of the ring. In the larger, complex forms the ring is plicately folded and there may be a centrally-placed hollow in the distal surface. Differences in morphology, in the folding pattern of the rings, in the structure (or absence) of a distal hollow, and in architecture of the major vascular system are employed to delimit genera. Size and shape of the organ, histological features, numbers of sporangia, and presence or absence of lacunae are used to delineate species. Currently recognised genera are Dolerotheca Halle, Sullitheca Stidd et al., Stewartiotheca Eggert and Rothwell, and Bernaultia gen. nov.