An alternative model for the earliest evolution of vascular plants

Land plants comprise the bryophytes and the polysporangiophytes. All extant polysporangiophytes are vascular plants (tracheophytes), but to date, some basalmost polysporangiophytes (also called protracheophytes) are considered non-vascular. Protracheophytes include the Horneophytopsida and Aglaophyton/Teruelia. They are most generally considered phylogenetically intermediate between bryophytes and vascular plants and are therefore essential to elucidate the origins of current vascular floras. Here, we propose an alternative evolutionary framework for the earliest tracheophytes. The supporting evidence comes from the study of the Rhynie chert historical slides from the Natural History Museum of Lille (France). From this, we emphasize that Horneophyton has a particular type of tracheid characterized by narrow, irregular, annular and/or, possibly spiral wall thickenings of putative secondary origin, and hence that it cannot be considered non-vascular anymore. Accordingly, our phylogenetic analysis resolves Horneophyton and allies (i.e. Horneophytopsida) within tracheophytes, but as sister to eutracheophytes (i.e. extant vascular plants). Together, horneophytes and eutracheophytes form a new clade called herein supereutracheophytes. The thin, irregular, annular to helical thickenings of Horneophyton clearly point to a sequential acquisition of the characters of water-conducting cells. Because of their simple conducting cells and morphology, the horneophytophytes may be seen as the precursors of all extant vascular plant biodiversity. © 2019 Lethaia Foundation. Published by John Wiley & Sons Ltd