“…In some cases, they occurred on most, if not all, lower epidermal cells, while in others they appeared only on the subsidiary or neighbour cells of the stomata, overarching the pore. As pointed out by Kerp & Barthel (1993), however, the presence of papillae can be a very variable feature, even within a single species, and great care must be taken when trying to use them as a diagnostic character; Zodrow et al (2010), for instance, found that in Macroneuropteris macrophylla there were more and larger papillae in the basal part of the frond than in more proximal parts. In most species the papillae were short, conical objects, the only exception being Neuralethopteris schlehanii where there was a gradation from short, conical to elongate papillae; the latter were in effect unicellular hairs.…”