Mature cone scale morphology in Pinaceae is varied but divides into two major groups (Fig. 2), each further divided into smaller subgroups, separable by differences in the shape and growth of the ovuliferous scale (hereafter referred to as the scale) and the position of the seeds on it, and the size and form of the bract scale (hereafter, the bract). These sub-groups correlate well with the different groupings indicated by the seed morphology and are keyed out below, with brief descriptions of the cones, scales and bracts, concentrating on significant or poorly described features.