PHYLOGENETIC DIVERSIFICATION OF EQUISETUM (EQUISETALES) AS INFERRED FROM LOWER CRETACEOUS SPECIES OF BRITISH COLUMBIA, CANADA

Three types of anatomically preserved vegetative shoots with features that characterize crown group Equisetum have been discovered in Lower Cretaceous deposits (approximate to 136 Ma) of British Columbia, Canada, suggesting the genus is much older than currently believed. Specimens include two types of aerial shoots described as E. haukeanum sp. nov. and E. vancouverense sp. nov. and one type of subterranean rhizome. Shoots are 1-2 mm in diameter, jointed, and in cross section have fluted stems with a hollow pith. Distinctive patterns of cortical sclerenchyma and different ridge morphologies characterize each shoot morphotype. Nodes display irregular branching, highly fused leaf sheaths, and a nodal diaphragm. The aerial stein morphospecies have vallecular canals on alternating radii with carinal canals of an equisetostele surrounded by only a few tracheids. No secondary tissues are produced. Bands of surficial stomata flank the furrows of one morphospecies. Rhizomes and aerial shoots are of a similar size, suggesting that the plants were equivalent in stature to the smallest living Equisetum species. These fossils augment our understanding of evolutionary transformations that led front Paleozoic Archaeocalamitaceae and Calamitaceae to crown group Equisetaceae, suggesting that the initial diversification of Equisetum began far earlier than suggested by molecular-clock-based estimates.