Growth, development, and systematics of ferns: Does Botrychium s.l. (Ophioglossales) really produce secondary xylem?1

Developmental morphology and anatomy of Botrychium s.l. were studied to clarify rhizome ontogeny and patterns of tissue maturation that can be used to test the hypothesis that ferns of the Ophioglossales may represent living progymnosperms. Serial anatomical sections of the rhizomes of B. virginianum and B. dissectum reveal that apical meristematic activity and vascular tissue maturation occur over an extended period of several years and then stop. Most of the xylem consists of radial rows of tracheids and interspersed ray-like xylem parenchyma cells that are similar in these respects to secondary xylem, but pits occur on all tracheid walls as is characteristic of primary xylem. No vascular cambium is initiated in mature primary tissues, nor is there secondary phloem. Radial rows of xylem cells are produced by the direct continuation of divisions that begin at the shoot apical meristem, forming a cylinder of radially aligned procambial cells before the differentiation of protoxylem. Continuing divisions over a period of several years increase the number of thin-walled cells and tracheids in each radial row back to about one internode behind where the current year's frond trace diverges from the rhizome stele. At more proximal levels of the rhizome, procambial cell divisions cease and there is no additional tracheid differentiation. These data reveal that the rhizome matures over an exceedingly long period of several years, but that growth is ultimately determinate, thus supporting hypotheses that the Ophioglossales is more closely related to other groups of living ferns than to progymnosperms and seed plants.