“…However, ladderanes do not preserve well (Rush et al, 2011) and, thus far, the oldest sedimentary record containing ladderanes extends only 140 kyr (Jaeschke et al, 2009). More recently, a unique bacteriohopanepolyol (BHP) biomarker for marine anammox 'Candidatus Scalindua profunda' was identified (Schwartz-Narbonne et al, 2020): a bacteriohopanetetrol isomer with unknown stereochemistry, BHT-x (Rush et al, 2014;Schwartz-Narbonne et al, 2020), has strongly depleted δ 13 C values in sediments consistent with fractionation associated with the reductive acetyl-CoA pathway used by anammox (Hemingway et al, 2018;Lengger et al, 2019), and has a distinct niche in the nitrite maximum of stratified water column settings (e.g., Matys et al, 2017). In comparison to ladderanes, BHPs preserve well and BHT has been detected in samples as old as ca.…”