“…Fossilized cyanobacteria with convincing heterocysts have been found in the 400-million-year (Ma)-old Rhynie Chert of Scotland (Croft & George, 1959), but most reports of older fossil heterocysts (e.g., Figure 6; Schopf, 1968;Licari & Cloud, 1968Cloud, 1976;Awramik & Barghoorn, 1977;Nagy, 1978) are disputed, so cannot reliably be used for interpreting ancient pN 2 . Time calibrations based on integrated phylogenetic and phenotypic data suggest that heterocystous cyanobacteria evolved 2,450-2,100 Ma ago (Tomitani et al, 2006), but possibly more recently (Uyeda, Harmon, & Blank, 2016). Alternatively, heterocystous cyanobacteria may have evolved well after other cyanobacterial assemblages, as has been proposed following phylogenetic analyses of single 1990), although oxygenic "whiffs" may have appeared in the Archean (Anbar et al, 2007).…”