“…Their temporal record spans minimally from Permian (Grimaldi and Engel 2005;Staniczek, 2007) to recent, with approximately 3500 extant species described so far (Sartori and Brittain, 2015). The phylogenetic position of the group is controversial and has been discussed on the basis of three hypothesis: (i) as closely related to Odonata (Palaeoptera hypothesis; Martynov, 1925;Crampton, 1924;Haas and Kukalov a-Peck, 2001;Kukalov a-Peck, 2008;Regier et al, 2010;Blanke et al, 2013;Thomas et al, 2013;Song et al, 2019), (ii) as closely related to Neoptera (Chiastomyaria hypothesis; Carle, 1982aCarle, , 1982bKjer, 2004;Yoshizawa and Johnson, 2005;Misof et al, 2007;Simon et al, 2009), and (iii) sister group of clade Odonata þ Neoptera (Metapterygota hypothesis; Staniczek, 2000;Wheeler et al, 2001;Ogden and Whiting, 2003;Grimaldi and Engel, 2005;Willkommen and H€ ornschemeyer, 2007;Zhang et al, 2008). Despite this long history of disagreement, there is a general consensus that mayflies constitute an early branch of winged insects (Grimaldi and Engel, 2005;Dominguez et al, 2006;Staniczek, 2007;Sartori and Brittain, 2015).…”