"Ferns, encompassing approximately 12,000 species (PPG 1, 2016), represent only about 4% of diversity (Mehltreter, 2010); nonetheless, they provide important (Sharpe et al., 2010), and comprise a substantial portion of commercial ornamental plant production as well as horticultural economic development (Hoshizaki and Moran, 2001; Singh and Johari, 2018). Their evolutionary and paleobiological history is extensive beginning in the , approximately 390 million years ago (mya) with expanded diversity during the (65 mya), when s were becoming more dominant (Schneider et al., 2004; Schuettpelz and Pryer, 2009). Ferns were dominant flora in the , later losing space to s and angiosperms. Increasing forests provided suitable protective, shady environments where ferns flourished on the forest floor or as epiphytes on tree trunks and limbs (Watkins and Cardelús, 2012). Subsequently, over geological time spans, through adaptation and evolutionary radiation (Sessa, 2018), ferns proliferated to occupy a wide diversity of terrestrial environments; ranging from s and coastal locations to more open s, mountain terrains, and s. Consequently, their and life histories ... have commensurately expanded to include a rich panoply of different life forms, physiological adaptive strategies, and wide global habitats."