@ARTICLE{TreeBASE2Ref17138, author = {Kathleen M. Pryer and Harald Schneider and Alan R. Smith and Raymond Cranfill and Paul G. Wolf and Jeffrey S. Hunt and S. D. Sipes}, title = {Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants.}, year = {2001}, keywords = {}, doi = {10.1038/35054555}, url = {}, pmid = {}, journal = {Nature}, volume = {409}, number = {}, pages = {618--622}, abstract = {Most of the 470-million-year history of plants on land belongs to bryophytes, pteridophytes, and gymnosperms, which eventually yielded to the remarkable ecological dominance by angiosperms 90 Myr ago. Our knowledge of angiosperm phylogeny, particularly the branching order of the earliest lineages, recently took an impressive step forward based on the concurrence of multigene sequence analyses. However, reconstructing relationships for all major lineages of vascular plants that diverged since the Devonian has remained a daunting challenge. Here we report on phylogenetic analyses of combined data from morphology and one nuclear and three plastid genes for 35 representatives from all major lineages of land plants. We demonstrate that there are three monophyletic groups of extant vascular plants: (1) lycophytes, (2) seed plants, and (3) a novel clade including equisetophytes (horsetails), psilotophytes (whisk ferns), and all eusporangiate and leptosporangiate ferns. Our maximum likelihood analysis shows with unambiguous support that horsetails and ferns together are the closest relatives to seed plants. This refutes the prevailing view that they are transitional evolutionary grades between bryophytes and seed plants and has important implications for our understanding of plant development and evolution.} }

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