RESEARCH ARTICLE INVITED SPECIAL ARTICLE For the Special Issue: The Tree of Death: The Role of Fossils in Resolving the Overall Pattern of Plant Phylogeny Origin of Equisetum: Evolution of horsetails (Equisetales) within the major euphyllophyte clade Sphenopsida Andrés Elgorriaga1,5 , Ignacio H. Escapa1, Gar W. Rothwell2,3, Alexandru M. F. Tomescu4, and N. Rubén Cúneo1 Manuscript received 3 December 2017; revision accepted 5 June PREMISE OF THE STUDY: Equisetum is the sole living representative of Sphenopsida, a 2018. clade with impressive species richness, a long fossil history dating back to the Devonian, 1 CONICET, Museo Paleontológico Egidio Feruglio, Trelew, and obscure relationships with other living pteridophytes. Based on molecular data, the Chubut 9100, Argentina crown group age of Equisetum is mid- Paleogene, although fossils with possible crown 2 Department of Botany and Plant Pathology, Oregon State synapomorphies appear in the Triassic. The most widely circulated hypothesis states that the University, Corvallis, OR 97331, USA lineage of Equisetum derives from calamitaceans, but no comprehensive phylogenetic studies 3 Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701, USA support the claim. Using a combined approach, we provide a comprehensive phylogenetic 4 Department of Biological Sciences, Humboldt State University, analysis of Equisetales, with special emphasis on the origin of genus Equisetum. Arcata, CA 95521, USA METHODS: We performed parsimony phylogenetic analyses to address relationships of 43 5 Author for correspondence (e-mail:
[email protected]) equisetalean species (15 extant, 28 extinct) using a combination of morphological and Citation: Elgorriaga, A., I. H. Escapa, G. W. Rothwell, A. M. F.