Archaea are ubiquitous in marine plankton, and fossil forms of archaeal tetraether membrane lipids in sedimentary rocks document their participation in marine biogeochemical cycles for >100 million years. Ribosomal RNA surveys have identified four major clades of planktonic archaea but, to date, tetraether lipids have been characterized in only one, the Marine Group I Thaumarchaeota. The membrane lipid composition of the other planktonic archaeal groups-all uncultured Euryarchaeota-is currently unknown. Using integrated nucleic acid and lipid analyses, we found that Marine Group II Euryarchaeota (MG-II) contributed significantly to the tetraether lipid pool in the North Pacific Subtropical Gyre at shallow to intermediate depths. Our data strongly suggested that MG-II also synthesize crenarchaeol, a tetraether lipid previously considered to be a unique biomarker for Thaumarchaeota. Metagenomic datasets spanning 5 y indicated that depth stratification of planktonic archaeal groups was a stable feature in the North Pacific Subtropical Gyre. The consistent prevalence of MG-II at depths where the bulk of exported organic matter originates, together with their ubiquitous distribution over diverse oceanic provinces, suggests that this clade is a significant source of tetraether lipids to marine sediments. Our results are relevant to archaeal lipid biomarker applications in the modern oceans and the interpretation of these compounds in the geologic record.euryarchaea | glycerol dialkyl glycerol tetraether | oceanography | environmental genomics | TEX 86 index

SignificanceCyanobacteria were responsible for the origin of oxygenic photosynthesis, and have since come to colonize almost every environment on Earth. Here we show that their ecological range is not limited by the presence of sunlight, but also extends down to the deep terrestrial biosphere. We report the presence of microbial communities dominated by cyanobacteria in the continental subsurface using microscopy, metagenomics, and antibody microarrays. These cyanobacteria were related to surface rock-dwelling lineages known for their high tolerance to environmental and nutritional stress. We discuss how these adaptations allow cyanobacteria to thrive in the dark underground, a lifestyle that might trace back to their nonphotosynthetic ancestors.

[2] Two commonly used proxies based on the distribution of glycerol dialkyl glycerol tetraethers (GDGTs) are the TEX 86 (TetraEther indeX of 86 carbon atoms) paleothermometer for sea surface temperature reconstructions and the BIT (Branched Isoprenoid Tetraether) index for reconstructing soil organic matter input to the ocean. An initial round-robin study of two sediment extracts, in which 15 laboratories participated, showed relatively consistent TEX 86 values (reproducibility 63-4 C when translated to temperature) but a large spread in BIT measurements (reproducibility 60.41 on a scale of 0-1). Here we report results of a second round-robin study with 35 laboratories in which three sediments, one sediment extract, and two mixtures of pure, isolated GDGTs were analyzed. The results for TEX 86 and BIT index showed improvement compared to the previous round-robin study. The reproducibility, indicating interlaboratory variation, of TEX 86 values ranged from 1.3 to 3.0 C when translated to temperature. These results are similar to those of other temperature proxies used in paleoceanography. Comparison of the results obtained from one of the three sediments showed that TEX 86 and BIT indices are not significantly affected by interlaboratory differences in sediment extraction techniques. BIT values of the sediments and extracts were at the extremes of the index with values close to 0 or 1, and showed good reproducibility (ranging from 0.013 to 0.042). However, the measured BIT values for the two GDGT mixtures, with known molar ratios of crenarchaeol and branched GDGTs, had intermediate BIT values and showed poor reproducibility and a large overestimation of the ''true'' (i.e., molarbased) BIT index. The latter is likely due to, among other factors, the higher mass spectrometric response of branched GDGTs compared to crenarchaeol, which also varies among mass spectrometers. Correction for this different mass spectrometric response showed a considerable improvement in the reproducibility of BIT index measurements among laboratories, as well as a substantially improved estimation of molar-based BIT values. This suggests that standard mixtures should be used in order to obtain consistent, and molar-based, BIT values.

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Abstract. Benthic foraminifera were cultured for five months at four temperatures (4, 7, 14 and 21 • C) to establish the temperature dependence of foraminiferal calcite δ 18 O and Mg/Ca. Two Bulimina species (B. aculeata and B. marginata) were most successful in terms of calcification, adding chambers at all four temperatures and reproducing at 7 and 14 • C. Foraminiferal δ 18 O values displayed ontogenetic variations, with lower values in younger individuals. The δ 18 O values of adult specimens decreased with increasing temperature in all but the 4 • C treatment, exhibiting a relationship consistent with previous δ 18 O paleotemperature calibration studies. Foraminiferal Mg/Ca values, determined by laser ablation inductively coupled plasma mass spectrometry, were broadly consistent with previous Mg/Ca calibration studies, but extremely high values in the 4 • C treatment and higher than predicted values at two of the other three temperatures make it challenging to interpret these results.

Serpentinization is a weathering process in which ultramafic rocks react with water, generating a range of products, including serpentine and other minerals, in addition to H 2 and low-molecular-weight hydrocarbons that are capable of sustaining microbial life. Lipid biomarker analyses of serpentinite-hosted ecosystems hold promise as tools for investigating microbial activity in ancient Earth environments and other terrestrial planets such as Mars because lipids have the potential for longer term preservation relative to DNA, proteins, and other more labile organic molecules. Here, we report the first lipid biomarker record of microbial activity in the mantle section of the Samail Ophiolite, in the Sultanate of Oman, a site undergoing active serpentinization. We detected isoprenoidal (archaeal) and branched (bacterial) glycerol dialkyl glycerol tetraether (GDGT) lipids, including those with 0-3 cyclopentane moieties, and crenarchaeol, an isoprenoidal GDGT containing four cyclopentane and one cyclohexane moieties, as well as monoether lipids and fatty acids indicative of sulfatereducing bacteria. Comparison of our geochemical data and 16S rRNA data from the Samail Ophiolite with those from other serpentinite-hosted sites identifies the existence of a common core serpentinization microbiome. In light of these findings, we also discuss the preservation potential of serpentinite lipid biomarker assemblages on Earth and Mars. Continuing investigations of the Samail Ophiolite and other terrestrial analogues will enhance our understanding of microbial habitability and diversity in serpentinite-hosted environments on Earth and elsewhere in the Solar System.

Abstract. Benthic foraminifera were cultured for five months at four temperatures (4, 7, 14 and 21 °C) to establish the temperature dependence of foraminiferal δ18O and Mg/Ca. Two Bulimina species (B. aculeata and B. marginata) were most successful, adding chambers at all four temperatures and reproducing at 7 and 14 °C. Foraminiferal δ18O values displayed ontogenetic variations, with lower values in younger individuals. The δ18O values of adult specimens decreased with increasing temperature in all but the 4 °C treatment, exhibiting a relationship consistent with previous δ18O paleotemperature calibration studies. Foraminiferal Mg/Ca values, determined by laser ablation inductively coupled plasma mass spectrometry, were broadly consistent with previous Mg/Ca calibration studies, but extremely high values in the 4 °C treatment and higher than expected values at two of the other three temperatures make it challenging to interpret these results.

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