Biomarker evidence for nitrogen-fixing cyanobacterial blooms in a brackish surface layer in the Nile River plume during sapropel deposition

Sapropels are organic-rich sediment layers deposited in the eastern Mediterranean Sea during precession minima, resulting from an increase in export productivity and/or preservation. Increased freshwater delivery from the African continent resulted in stratification, causing deepwater anoxia, while nutrient input stimulated productivity, presumably at the deep chlorophyll maximum. Previous studies have suggested that during sapropel deposition, nitrogen fixation was widespread in the highly stratified surface waters, and that cyanobacteria symbiotic with diatoms (diatom-diazotroph associations, DDAs) were responsible. Here we analyzed sapropel S5 sediments for heterocyst glycolipids (HGs) from three locations in the eastern Mediterranean. HG biomarkers can differentiate between those heterocystous cyanobacteria that are free living (found predominately in freshwater or brackish environments) and those that are from DDAs (found in marine settings). In our primary core, from a location which would have been influenced by the Nile River outflow, we detected a HG with a pentose (C5) head group specific for DDAs. However, HGs with a hexose (C6) head group, specific to free-living cyanobacteria, were present in substantially (up to 60x) higher concentration. These data suggest that at our study location, free-living cyanobacteria were the dominant diazotrophs, rather than DDAs. The C6 HGs increased substantially at the onset of sapropel S5 deposition, suggesting that substantial seasonal cyanobacterial blooms were associated with a brackish surface layer flowing from the Nile into the eastern Mediterranean. Two additional S5 sapropels were analyzed, one also from the Nile delta region and one from the region between Libya and southwestern Crete. Overall, comparison of the HG distribution in the three S5 sapropels provides evidence that all three locations were initially influenced by surface salinities that were sufficiently low to support free-living heterocystous cyanobacteria. While free-living heterocystous cyanobacteria continued to outnumber DDAs during sapropel deposition at the two Nile-influenced sites, DDAs, indicators of persistent marine salinities, were the dominant diazotrophs in the upper part of the sapropel at the more westerly site. These results indicate that N2 fixation by free-living cyanobacteria offers an important additional mechanism to stimulate productivity in regions with strong river discharge during sapropel deposition