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The Perseverance rover landed in Jezero crater, Mars, in February 2021. We used the Scanning Habitable Environments with Raman and Luminescence for Organics and Chemicals (SHERLOC) instrument to perform deep-ultraviolet Raman and fluorescence spectroscopy of three rocks within the crater. We identify evidence for two distinct ancient aqueous environments at different times. Reactions with liquid water formed carbonates in an olivine-rich igneous rock. A sulfate-perchlorate mixture is present in the rocks, which probably formed by later modifications of the rocks by brine. Fluorescence signatures consistent with aromatic organic compounds occur throughout these rocks and are preserved in minerals related to both aqueous environments.

The record of life during the Proterozoic is preserved by several different lithologies, but two in particular are linked both spatially and temporally: chert and carbonate. These lithologies capture a snapshot of dominantly peritidal environments during the Proterozoic. Early diagenetic chert preserves some of the most exceptional Proterozoic biosignatures in the form of microbial body fossils and mat textures. This fossiliferous and kerogenous chert formed in shallow marine environments, where chert nodules, layers, and lenses are often surrounded by and encased within carbonate deposits that themselves often contain kerogen and evidence of former microbial mats. Here, we review the record of biosignatures preserved in peritidal Proterozoic chert and chert-hosting carbonate and discuss this record in the context of experimental and environmental studies that have begun to shed light on the roles that microbes and organic compounds may have played in the formation of these deposits. Insights gained from these studies suggest temporal trends in microbial-environmental interactions and place new constraints on past environmental conditions, such as the concentration of silica in Proterozoic seawater, interactions among organic compounds and cations in seawater, and the influence of microbial physiology and biochemistry on selective preservation by silicification.

The preservation of organic biosignatures during the Proterozoic Eon required specific taphonomic windows that could entomb organic matter to preserve amorphous kerogen and even microbial body fossils before they could be extensively degraded. Some of the best examples of such preservation are found in early diagenetic chert that formed in peritidal environments. This chert contains discrete domains of amorphous kerogen and sometimes kerogenous microbial mat structures and microbial body fossils. Our understanding of how these exquisite microfossils were preserved and the balance between organic degradation and mineral formation has remained incomplete. Here, we present new insights into organic preservation in Proterozoic peritidal environments facilitated through interactions among organic matter, cations, and silica. Organic matter from Proterozoic peritidal environments is not preserved by micro- or cryptocrystalline quartz alone. Rather, preservation includes cation-rich nanoscopic phases containing magnesium, calcium, silica, and aluminum that pre-date chert emplacement and may provide nucleation sites for silica deposition and enable further chert development. Using scanning electron microscopy and elemental mapping with energy dispersive X-ray spectroscopy, we identify cation enrichment in Proterozoic organic matter and cation-rich nanoscopic phases that pre-date chert. We pair these analyses with precipitation experiments to investigate the role of cations in the precipitation of silica from seawater. Our findings suggest that organic preservation in peritidal environments required rapid formation of nanoscopic mineral phases through the interactions of organic matter with seawater. These organic-cation interactions likely laid the initial foundation for the preservation and entombment of biosignatures, paving the way for the development of the fossiliferous chert that now contains these biosignatures and preserves a record of Proterozoic life.

Enzymes catalyze key reactions within Earth’s life-sustaining biogeochemical cycles. Here, we use metaproteomics to examine the enzymatic capabilities of the microbial community (0.2 to 3 µm) along a 5,000-km-long, 1-km-deep transect in the central Pacific Ocean. Eighty-five percent of total protein abundance was of bacterial origin, with Archaea contributing 1.6%. Over 2,000 functional KEGG Ontology (KO) groups were identified, yet only 25 KO groups contributed over half of the protein abundance, simultaneously indicating abundant key functions and a long tail of diverse functions. Vertical attenuation of individual proteins displayed stratification of nutrient transport, carbon utilization, and environmental stress. The microbial community also varied along horizontal scales, shaped by environmental features specific to the oligotrophic North Pacific Subtropical Gyre, the oxygen-depleted Eastern Tropical North Pacific, and nutrient-rich equatorial upwelling. Some of the most abundant proteins were associated with nitrification and C1 metabolisms, with observed interactions between these pathways. The oxidoreductases nitrite oxidoreductase (NxrAB), nitrite reductase (NirK), ammonia monooxygenase (AmoABC), manganese oxidase (MnxG), formate dehydrogenase (FdoGH and FDH), and carbon monoxide dehydrogenase (CoxLM) displayed distributions indicative of biogeochemical status such as oxidative or nutritional stress, with the potential to be more sensitive than chemical sensors. Enzymes that mediate transformations of atmospheric gases like CO, CO 2 , NO, methanethiol, and methylamines were most abundant in the upwelling region. We identified hot spots of biochemical transformation in the central Pacific Ocean, highlighted previously understudied metabolic pathways in the environment, and provided rich empirical data for biogeochemical models critical for forecasting ecosystem response to climate change.

Microbial fossils preserved by early diagenetic chert provide a window into the Proterozoic biosphere, but seawater chemistry, microbial processes, and the interactions between microbes and the environment that contributed to this preservation are not well constrained. Here, we use fossilization experiments to explore the processes that preserve marine cyanobacterial biofilms by the precipitation of amorphous silica in a seawater medium that is analogous to Proterozoic seawater. These experiments demonstrate that the exceptional silicification of benthic marine cyanobacteria analogous to the oldest diagnostic cyanobacterial fossils requires interactions among extracellular polymeric substances (EPS), photosynthetically induced pH changes, magnesium cations (Mg2+ ), and >70 ppm silica.

Cyanobacteria and extracellular polymeric substances (EPS) in peritidal pustular microbial mats have a two-billion-year-old fossil record. To understand the composition, production, degradation, and potential role of EPS in modern analogous communities, we sampled pustular mats from Shark Bay, Australia and analyzed their EPS matrix. Biochemical and microscopic analyses identified sulfated organic compounds as major components of mat EPS. Sulfur was more abundant in the unmineralized regions with cyanobacteria and less prevalent in areas that contained fewer cyanobacteria and more carbonate precipitates. Sequencing and assembly of the pustular mat sample resulted in 83 high-quality metagenome-assembled genomes (MAGs). Metagenomic analyses confirmed cyanobacteria as the primary sources of these sulfated polysaccharides. Genes encoding for sulfatases, glycosyl hydrolases, and other enzymes with predicted roles in the degradation of sulfated polysaccharides were detected in the MAGs of numerous clades including Bacteroidetes, Chloroflexi, Hydrogenedentes, Myxococcota, Verrucomicrobia, and Planctomycetes. Measurable sulfatase activity in pustular mats and fresh cyanobacterial EPS confirmed the role of sulfatases in the degradation of sulfated EPS. These findings suggest that the synthesis, modification, and degradation of sulfated polysaccharides influence microbial interactions, carbon cycling, and biomineralization processes within peritidal pustular microbial mats.

Microbial exopolymeric substances (EPSs) form the organic, polysaccharide-rich matrix of marine microbial mats and can mediate the binding and precipitation of carbonate minerals therein. Here, we investigate the molecular ecology of carbohydrate degradation and production in pustular mats from Shark Bay, Western Australia, by analyzing 84 metagenome-assembled genomes (MAGs) and the composition of microbial communities enriched from a pustular mat on various polysaccharide substrates. The annotation of 4000 genes from hundreds of carbohydrate-active enzyme (CAZyme) families in the MAGs and mapping of polysaccharide-degrading CAZymes to their predicted substrates identify trends in the distribution and localization of degradation-associated CAZymes across different bacterial phyla. The compositions of microbial communities enriched on a range of polysaccharides inoculated with pustular mat material support the predicted trends. The combined metagenomic and experimental analyses reveal a widespread potential for EPS degradation among MAGs from Shark Bay pustular mats and suggest distinct roles for some phyla that are reported at high abundances in mats. Specifically, Bacteroidetes are likely to be primary degraders of polysaccharide EPSs, alongside Planctomycetes and a small subset of Alphaproteobacteria and Gammaproteobacteria. Planctomycetes, some Bacteroidetes, Verrucomicrobia, Myxococcota and Anaerolineae are also predicted to favor degradation of sulfated substrates, which are present in the EPS matrix of pustular mats. Large sets of functionally varied CAZymes without signal peptides tagging them for export implicate Anaerolineae and Verrucomicrobia in degrading the downstream products of primary EPS degradation.ImportanceModern marine microbial mats are rich in exopolymeric substances (EPSs) — complex, high molecular weight polymers secreted by bacteria — that mediate the formation of carbonate minerals and the preservation of microbial textures in mats. However, the organisms involved in EPS cycling in these mats have not been identified and the links between EPS degradation, carbonate precipitation, and microbial ecology in mats remain poorly understood. We define distinct roles in EPS cycling for many major microbial taxa that are both ubiquitous and abundant in pustular microbial mats from Shark Bay, Australia. The large genomic potential of these microbes for the modification and degradation of diverse extracellular organic polymers provides a blueprint for future studies aimed at quantifying and verifying the specific contributions of these microbes to EPS degradation, carbon cycling and carbonate precipitation.

The Late Heavy Bombardment (LHB) and other late accretion impactor scenarios are often invoked as habitability constraints on the Hadean/Eoarchean Earth. These hypotheses either describe an “impact frustration” where life would not arise until high impact fluxes abated, or “impact bottlenecks” with Bacteria and Archaea representing surviving lineages that subsequently diversified. Phylogenomics studies using relaxed molecular clocks have frequently used these early impact fluxes, especially the LHB, as older-bound constraints on extant life’s early diversification. However, the intensity, timing, and sterilization potential of these scenarios is poorly constrained, and lacks consensus. We propose inverting this hypothesis testing, evaluating late accretion impact hypotheses using molecular clocks that do not presuppose impact frustration or bottlenecks as constraints. However, in the absence of these constraints, previous studies lack the precision to discriminate between these hypotheses. Our recently developed molecular clock approach, using horizontal gene transfers as “cross cutting events” between lineages, overcomes this limitation, and provides sufficient precision to test the proposed biological impact of specific planetary hypotheses such as the LHB. Using this methodology, we show that major bacterial groups likely diversified between 3.75 and 3.55 Ga, with the last common ancestor of extant Bacteria likely existing shortly after 3.8 Ga. These ages are consistent with the LHB impact bottleneck hypothesis, wherein bacteria and archaea represent survivors of early Archean cataclysms that extinguished most primordial biodiversity ~3.9 Ga. Future work extending this methodology to Archaea can potentially provide an independent test of the impact bottleneck hypothesis.

The discovery of hydrated magnesium carbonates in a paleolake in Jezero Crater, Mars, has inspired the study of carbonates associated with lacustrine systems on Earth. Lake Salda, Turkey, contains hydromagnesite-dominated microbialites in a mafic-rock hosted terrain. This alkaline lacustrine system provides an excellent natural laboratory in which to characterize various depositional environments and biosignature preserving potential of microbialites and other magnesium carbonate deposits. Here we aim to describe the textures, mineral composition, and macroscopic morphology of hydromagnesite structures in Lake Salda. All microbialites are covered by a thick, sticky organic-rich layer dominated by diatoms and cyanobacteria. A close association of contemporary hydromagnesite precipitation with organic-rich layers in microbialites indicates biologically mediated precipitation. Microbialites around the lake exhibit a range of morphologies, including finger-like, mini columnar, domical, and structureless. The recently exposed microbialites commonly contain colloform and cerebroid structures with fine, crude lamination, vuggy and globular textures. Layered microscopic textures preserve evidence of partially mineralized filamentous bacteria that trapped detrital grains and also contain globules embedded in extracellular polymeric substances, and dense hydromagnesite. The differences in mineralogy, density, and organic preservation potential within these textures likely correspond to the depositional environment. For example, the globular textures are closely associated with the purple layers in microbialites, including the layers in the recently sampled microbialites from the deeper part of the lake (ca. 10 m). These layers may form in the presence of anoxygenic purple sulfur bacteria. Our preliminary results suggest that the textures and macroscopic structures of hydromagnesite microbialites may record and reflect relationships among carbonate minerals, environmental factors, and microbial community composition.

Silicification was a major mode of fossilization in Proterozoic peritidal environments, but marine silica concentrations and the chemical and biological mechanisms that drove microbial silicification and formation of early diagenetic chert in these environments remain poorly constrained. Here, we use taphonomy experiments to demonstrate that photosynthetically active cyanobacteria that are morphologically analogous to the oldest cyanobacterial fossil, Eoentophysalis, mediate the formation of magnesium-rich amorphous silica in seawater that is undersaturated with respect to silica. These results show that microbes in Proterozoic tidal environments may have mediated their own silicification at lower silica concentrations than previously assumed.

© 2020 American Chemical Society. Methanogenic archaea have been shown to reduce iron from ferric [Fe(III)] to ferrous [Fe(II)] state, but minerals that form during iron reduction by different methanogens remain to be characterized. Here, we show that zerovalent iron (ZVI) minerals, ferrite [α-Fe(0)] and austenite [γ-Fe(0)], appear in the X-ray diffraction spectra minutes after the addition of ferrihydrite to the cultures of a methanogenic archaeon, Methanosarcina barkeri (M. barkeri). M. barkeri cells and redox-active, nonenzymatic soluble organic compounds in organic-rich spent culture supernatants can promote the formation of ZVI; the latter compounds also likely stabilize ZVI. Methanogenic microbes that inhabit organic- and Fe(III)-rich anaerobic environments may similarly reduce Fe(III) to Fe(II) and ZVI, with implications for the preservation of paleomagnetic signals during sediment diagenesis and potential applications in the protection of iron metals against corrosion and in the green synthesis of ZVI.

The NASA Mars 2020 Perseverance rover mission will collect a suite of scientifically compelling samples for return to Earth. On the basis of orbital data, the Mars 2020 science team identified two notional sample caches to study (1) the geology of Jezero crater, collected during the prime mission and (2) the ancient crust outside of Jezero crater, collected during a possible extended mission. Jezero crater geology consists of well-preserved, Early Hesperian to Late Noachian deltaic and lacustrine deposits sourced from a river system that drained Noachian terrain. The crater floor comprises at least two distinct units of sedimentary or volcanic origin whose relationship to the deltaic deposits is presently unclear. Remotely-sensed data reveal signatures of carbonate+olivine and clay minerals within crater floor and crater margin units. Samples from within Jezero that comprise the prime mission notional sample collection thus include: crater floor units; fine- and coarse-grained delta facies, the former with potential to preserve organic matter and/or biosignatures, the latter to possibly constrain the type and timing of sediment deposition; chemical sediments with the potential to preserve biosignatures; a sample of crater rim bedrock; and at least one sample of regolith. The region of southern Nili Planum, directly outside the western rim of Jezero crater, is geologically distinct from Jezero crater and contains diverse Early or even Pre-Noachian lithologies, that may contain records of early planetary differentiation, magnetism, paleoclimate and habitability. The notional sample collection from this region will include: layered and other basement rocks; megabreccias, which may represent blocks of (pre-)Noachian crust; basement-hosted hydrothermal fracture fill; olivine+carbonate rocks that are regionally significant and may be related to units within Jezero crater; and mafic cap unit rocks. The samples described are notional and may change with ongoing surface investigations. However, the samples we anticipate collecting align well with community priorities for Mars exploration, addressing geologic diversity, potential ancient biologic activity on Mars, planetary evolution, volatiles, and human health hazards. Many other Mars 2020 team members were involved in this planning.

Perseverance’s Mastcam-Z instrument provides high-resolution stereo and multispectral images with a unique combination of spatial resolution, spatial coverage, and wavelength coverage along the rover’s traverse in Jezero crater, Mars. Images reveal rocks consistent with an igneous (including volcanic and/or volcaniclastic) and/or impactite origin and limited aqueous alteration, including polygonally fractured rocks with weathered coatings; massive boulder-forming bedrock consisting of mafic silicates, ferric oxides, and/or iron-bearing alteration minerals; and coarsely layered outcrops dominated by olivine. Pyroxene dominates the iron-bearing mineralogy in the fine-grained regolith, while olivine dominates the coarse-grained regolith. Solar and atmospheric imaging observations show significant intra- and intersol variations in dust optical depth and water ice clouds, as well as unique examples of boundary layer vortex action from both natural (dust devil) and Ingenuity helicopter–induced dust lifting. High-resolution stereo imaging also provides geologic context for rover operations, other instrument observations, and sample selection, characterization, and confirmation.

Perseverance images of a delta on Mars The Perseverance rover landed in Jezero crater, Mars, in February 2021. Earlier orbital images showed that the crater contains an ancient river delta that was deposited by water flowing into a lake billions of years ago. Mangold et al . analyzed rover images taken shortly after landing that show distant cliff faces at the edge of the delta. The exposed stratigraphy and sizes of boulders allowed them to determine the past lake level and water discharge rates. An initially steady flow transitioned into intermittent floods as the planet dried out. This history of the delta’s geology provides context for the rest of the mission and improves our understanding of Mars’ ancient climate. —KTS

The extent to which nutrients structure microbial communities in permanently stratified lakes is not well understood. This study characterized microbial communities from the anoxic layers of the meromictic and sulfidic Fayetteville Green Lake (FGL), NY, and investigated the roles of organic electron donors and terminal electron acceptors in shaping microbial community structure and interactions. Bacterial communities from the permanently stratified layer below the chemocline (monimolimnion) and from enrichment cultures inoculated by lake sediments were analyzed using 16S rRNA gene sequencing. Results showed that anoxygenic phototrophs dominated microbial communities in the upper monimolimnion (21 m), which harbored little diversity, whereas the most diverse communities resided at the bottom of the lake (~52 m). Organic electron donors explained 54% of the variation in the microbial community structure in aphotic cultures enriched on an array of organic electron donors and different inorganic electron acceptors. Electron acceptors only explained 10% of the variation, but were stronger drivers of community assembly in enrichment cultures supplemented with acetate or butyrate compared to the cultures amended by chitin, lignin or cellulose. We identified a range of habitat generalists and habitat specialists in both the water column and enrichment samples using Levin’s index. Network analyses of interactions among microbial groups revealed Chlorobi and sulfate reducers as central to microbial interactions in the upper monimolimnion, while Syntrophaceae and other fermenting organisms were more important in the lower monimolimnion. The presence of photosynthetic microbes and communities that degrade chitin and cellulose much below the chemocline supported the downward transport of microbes, organic matter and oxidants from the surface and the chemocline. Collectively, our data suggest niche partitioning of bacterial communities by interactions that depend on the availability of different organic electron donors and terminal electron acceptors. Thus, light, as well as the diversity and availability of chemical resources drive community structure and function in FGL, and likely in other stratified, meromictic lakes.

Methanogenic archaea have been shown to reduce iron from ferric [Fe(III)] to ferrous [Fe(II)] state, but minerals that form during iron reduction by different methanogens remain to be characterized. Here, we show that zerovalent iron (ZVI) minerals, ferrite [α-Fe(0)] and austenite [γ-Fe(0)], appear in the X-ray diffraction spectra minutes after the addition of ferrihydrite to the cultures of a methanogenic archaeon

Sediment resuspension and sediment erodibility were investigated in Hamelin Pool (Western Australia), which is one of the few modern carbonate settings where stromatolites and ooids are actively forming. For this purpose, the first time series of wave height and water turbidity were recorded at a ~2.5 m deep subtidal location in Hamelin Pool from July 2017 to March 2018. Sediment grain size analyses indicate that this environment is nearly free of mud, so the optical turbidity sensor was able to detect the intra-wave modulation of sand resuspension. This allowed us to use the standard deviation of the turbidity within a burst – as opposed to the mean turbidity – as a proxy for sand resuspension. This approach was further validated by wave tank experiments, which also helped to identify the occurrence of biofouling and discard the affected data. Although the wave regime was similar throughout the year, sand resuspension changed with seasons. Sediment erodibility, defined as the sand resuspension normalized by the bed shear stress, was higher during the Austral winter and spring, and lower in summer and autumn. This decrease in erodibility coincided with an increase in temperature and solar irradiance. These two environmental factors likely modulate the growth of benthic microbes, which in turn affects sediment dynamics. This seasonal modulation of sediment dynamics may influence additional processes, including the delivery of sand grains to stromatolites and the precipitation of aragonite around ooids and in stromatolite laminae.

The extent of oxygenated environments on the early Earth was much lower than today, and cyanobacteria were critical players in Earth's shift from widespread anoxia to oxygenated surface environments. Extant cyanobacteria that aggregate into cones, tufts and ridges are used to understand the long record of photosynthesis and microbe-mineral interactions during times when oxygen was much lower, i.e., the Archean and the Proterozoic. To better understand the metabolic versatility and physiological properties of these organisms, we examined publicly available genomes of cyanobacteria from modern terrestrial hydrothermal systems and a newly sequenced genome of a cyanobacterium isolated from conical and ridged microbialites that grow in occasionally sulfidic hydrothermal springs in Yellowstone National Park, USA. Phylogenomic analyses reveal that cyanobacteria from globally distributed terrestrial and shallow marine hydrothermal systems form a monophyletic clade within the Cyanobacteria phylum. Comparative genomics of this clade reveals the genetic capacity for oxygenic photosynthesis that uses photosystems I and II, and anoxygenic photosynthesis that uses a putative sulfide quinone reductase to oxidize sulfide and bypass photosystem II. Surprisingly large proportions of the newly sequenced genome from Yellowstone National Park are also dedicated to secondary metabolite production (15.1-15.6%), of which ∼6% can be attributed to antibiotic production and resistance genes. All this may be advantageous to benthic, mat-forming photosynthesizers that have to compete for light and nutrients in sporadically or permanently sulfidic environments, and may have also improved the tolerance of ancient counterparts of these cyanobacteria to sulfidic conditions in benthic communities that colonized the coastal margins in the Archean and the Proterozoic.

Dolomite enabled the preservation of fine microbial textures in some Archean and Proterozoic marine microbialites, but has rarely done so during the Phanerozoic. Here, we report precipitation of dolomite in anoxygenic photosynthetic biofilms grown under chemical conditions relevant for Archean seawater. Ordered dolomite nucleates primarily on the surfaces of photosynthetic cells when manganese(II) is present, and nanocrystals of disordered dolomite form on exopolymeric substances in microbial cultures grown either in the dark or without manganese. Dolomite nucleation and maturation on different surfaces in photosynthetically active cultures amended with 0.1-1 mM manganese(II) enables the preservation of biofilm textures at scales larger than individual microbial cells. This provides a new model for the preservation of microbial textures by dolomite before the oxygenation of the oceanic photic zone., National Science Foundation (Award DMR-1419807)

Siliciclastic sediments of the Ediacaran Period contain exceptionally preserved fossils of macroscopic organisms, including three-dimensional casts and molds commonly found in sandstones and siltstones and some two-dimensional compressions reported in shales. The sporadic and variable associations of these exceptionally preserved macroscopic fossils with pyrite, clay minerals, and microbial fossils and textures complicate our understanding of fossilization processes. This hinders inferences about the evolutionary histories, tissue types, original morphologies, and lifestyles of the enigmatic Ediacara biota. Here, we investigate the delayed decay of scallop muscles buried in quartz sand or kaolinite for 45 days. This process occurs in the presence of microbial activity in mixed redox environments, but in the absence of thick, sealing microbial mats. Microbial processes that mediate organic decay and release the highest concentrations of silica and Fe(II) into the pore fluids are associated with the most extensive tissue decay. Delayed decay and the preservation of thick muscles in sand are associated with less intense microbial iron reduction and the precipitation of iron oxides and iron sulfides that contain Fe(II) or Fe(III). In contrast, muscles buried in kaolinite are coated only by, NASA Astrobiology Institute (Grant NNA13AA90A), Simons Foundation (Grants 327126 and 344707), American Chemical Society (Award 54498-ND8)

The phylum Cyanobacteria includes free-living bacteria and plastids, the descendants of cyanobacteria that were engulfed by the ancestral lineage of the major photosynthetic eukaryotic group Archaeplastida. Endosymbiotic events that followed this primary endosymbiosis spread plastids across diverse eukaryotic groups. The remnants of the ancestral cyanobacterial genome present in all modern plastids, enable the placement of plastids within Cyanobacteria using sequence-based phylogenetic analyses. To date, such phylogenetic studies have produced conflicting results and two competing hypotheses: (1) plastids diverge relatively recently in cyanobacterial evolution and are most closely related to nitrogen-fixing cyanobacteria, or (2) plastids diverge early in the evolutionary history of cyanobacteria, before the divergence of most cyanobacterial lineages. Here, we use phylogenetic analysis of ribosomal proteins from an expanded data set of cyanobacterial and representative plastid genomes to infer a deep placement for the divergence of the plastid ancestor lineage. We recover plastids as sister to Gloeomargarita and show that the group diverges from other cyanobacterial groups before Pseudanabaena, a previously unreported placement. The tree topologies and phylogenetic distances in our study have implications for future molecular clock studies that aim to model accurate divergence times, especially with respect to groups containing fossil calibrations. The newly sequenced cyanobacterial groups included here will also enable the use of novel cyanobacterial microfossil calibrations.

Ooids are sedimentary grains that are distributed widely in the geologic record. Their formation is still actively debated, which limits our understanding of the significance and meaning of these grains in Earth's history. Central questions include the role played by microbes in the formation of ooids and the sources of ubiquitous organic matter within ooid cortices. To address these issues, we investigated the microbial community composition and associated lipids in modern oolitic sands at Pigeon Cay on Cat Island, The Bahamas. Surface samples were taken along a transect from the shallow, turbulent surf zone to calmer, deeper water. Grains transitioned from shiny and abraded ooids in the surf zone, to biofilm-coated ooids at about 3 m water depth. Further offshore, grapestones (cemented aggregates of ooids) dominated. Benthic diatoms and Proteobacteria dominated biofilms. Taxa that may promote carbonate precipitation were abundant, particularly those associated with sulfur cycling. Compared to the lipids associated with surface biofilms, relict lipids bound within carbonate exhibited remarkably similar profiles in all grain types. The enhanced abundance of methyl-branched fatty acids and β-hydroxy fatty acids, 1-O-monoalkyl glycerol ethers and hopanoids bound within ooid and grapestone carbonate confirms a clear association of benthic sedimentary bacteria with these grains. Lipids bound within ooid cortices also contain molecular indicators of microbial heterotrophic degradation of organic matter, possibly in locally reducing conditions. These included the loss of labile unsaturated fatty acids, enhanced long-chain fatty acids/short-chain fatty acids, enriched stable carbon isotopes ratios of fatty acids, and very high stanol/stenol ratios. To what extent some of these molecular signals are derived from later heterotrophic endolithic activity remains to be fully resolved. We speculate that some ooid carbonate forms in microbial biofilms and that early diagenetic degradation of biofilms may also play a role in early stage carbonate precipitation around ooids.

Ediacaran sedimentary successions are noted for the preservation of microbes and microbial textures on the surfaces of sandstones and siltstones. Although microorganisms have been preserved in coarse-grained siliciclastic sand throughout geologic history, the exceptional preservation of microbes in Ediacaran sediments suggests the potential for a unique taphonomic window. Here, we identify conditions conducive to the fossilization of filamentous cyanobacteria growing in the presence of siliciclastic sand and demonstrate that the sheaths of filamentous cyanobacteria can become coated by clay minerals within days under oxic conditions. Smooth, extensive mineral coatings develop in the presence of 5.6 to 55.6 mg/L of suspended clay and 0.1 mM or greater concentrations of dissolved silica. Thus, elevated concentrations of seawater silica and the delivery of suspended clays promote microbial preservation on sandy and silty surfaces. These factors likely facilitated microbial fossilization in coarse-grained siliciclastic sand throughout the Ediacaran Period and may have also contributed to microbial fossilization in siliciclastic deposits at other times throughout Earth’s history.

Rationale The meromictic Fayetteville Green Lake (FGL) is of significant geobiological interest because of microbial cycling of sulfur within and below the permanent chemocline and in the euxinic deep waters. Studies of glycerol dibiphytanyl glycerol tetraethers (GDGTs) may help shed light on understanding the activity of archaeal communities in these habitats. Methods Normal-phase and reversed-phase liquid chromatography/mass spectrometry (LC/MS) analysis on total lipid extracts of environmental samples revealed series of GDGTs with different biphytane structures. Comparison of the mass spectrum of biphytane obtained from separated novel GDGTs with that of a synthetic C40 biphytane confirms our structural assignments. Results A unique cyclohexyl ring configured in the middle of a C40 biphytane chain was identified in these novel GDGTs. We suggest the trivial name S-GDGTs for these compounds, where ‘S′ stands for ‘sulfidic’ and ‘six-membered ring’. S-GDGT derivatives composed of biphytanes modified with double bonds and cyclopentane rings were also detected in the samples we analyzed. Intact polar lipid precursors of S-GDGT include compounds with mono- and diglycosyl head groups. Conclusions The carbon isotopic composition of S-GDGTs and their occurrence in FGL, Messel Shale as well as Salt Pond and salt marshes on Cape Cod suggest that S-GDGTs may be produced by chemoautotrophic archaea that prefer sulfidic conditions. Copyright © 2016 John Wiley & Sons, Ltd.

Three-dimensionally preserved Ediacaran fossils occur globally within sandstone beds. Sandy siliciclastic deposits of the Ediacaran Wood Canyon Formation (WCF) in the Montgomery Mountains, Nevada, contain two fossil morphologies with similar shapes and sizes: one exhibits mm-scale ridges and a distinct lower boundary and the other is devoid of these diagnostic features. We interpret these as taphomorphs of erniettomorphs, soft-bodied organisms with uncertain taxonomic affinities. We explore the cast-and-mould preservation of both taphomorphs by petrography, Raman spectroscopy, X-ray fluorescence microprobe and X-ray diffraction. All fossils and the surrounding sedimentary matrix contain quartz grains, iron-rich chlorite and muscovite. The ridged fossils contain about 70% larger quartz grains compared to the ridgeless taphomorph, indicating a lower abundance of clay minerals in the ridged fossil. Chlorite and muscovite likely originated from smectite and kaolinite precursors that underwent lower greenschist facies metamorphism. Kaolinite and smectite are inferred to have been abundant in sediments around the ridged fossil, which enabled the preservation of a continuous, distinct, clay- and kerogen-rich bottom boundary. The prevalence of quartz in the ridged fossils of the WCF and in erniettomorphs from other localities also suggests a role for this mineral in three-dimensional preservation of erniettomorphs in sandstone and siltstone deposits.

Oxygenic photosynthesis supplies organic carbon to the modern biosphere, but it is uncertain when this metabolism originated. It has previously been proposed1,2 that photosynthetic reaction centres capable of splitting water arose by about 3billion years ago on the basis of the inferred presence of manganese oxides in Archaean sedimentary rocks. However, this assumes that manganese oxides can be produced only in the presence of molecular oxygen3, reactive oxygen species4,5 or by high-potential photosynthetic reaction centres6,7. Here we show that communities of anoxygenic photosynthetic microorganisms biomineralize manganese oxides in the absence of molecular oxygen and high-potential photosynthetic reaction centres. Microbial oxidation of Mn(II) under strictly anaerobic conditions during the Archaean eon would have produced geochemical signals identical to those used to date the evolution of oxygenic photosynthesis before the Great Oxidation Event1,2. This light-dependent process may also produce manganese oxides in the photic zones of modern anoxic water bodies and sediments.

The Martian surface is cold, dry, exposed to biologically harmful radiation and apparently barren today. Nevertheless, there is clear geological evidence for warmer, wetter intervals in the past that could have supported life at or near the surface. This evidence has motivated National Aeronautics and Space Administration and European Space Agency to prioritize the search for any remains or traces of organisms from early Mars in forthcoming missions. Informed by (1) stratigraphic, mineralogical and geochemical data collected by previous and current missions, (2) Earth's fossil record, and (3) experimental studies of organic decay and preservation, we here consider whether, how, and where fossils and isotopic biosignatures could have been preserved in the depositional environments and mineralizing media thought to have been present in habitable settings on early Mars. We conclude that Noachian‐Hesperian Fe‐bearing clay‐rich fluvio‐lacustrine siliciclastic deposits, especially where enriched in silica, currently represent the most promising and best understood astropaleontological targets. Siliceous sinters would also be an excellent target, but their presence on Mars awaits confirmation. More work is needed to improve our understanding of fossil preservation in the context of other environments specific to Mars, particularly within evaporative salts and pore/fracture‐filling subsurface minerals., Key Points Noachian‐Hesperian Fe‐bearing clay‐rich fluvio‐lacustrine siliciclastic sediments are favored in the search for ancient Martian lifeThere is insufficient confidence in the nature of reported silica sinters on Mars or the possibility of preservation in the deep biosphereExperimental taphonomy approaches from paleontology should now be adapted to understand limits on preservation under Martian conditions

Raman spectroscopy has been used extensively in thermal maturation studies of kerogen, but has not been used to examine the maturation of organic cements in agglutinated foraminifera. Here, we use Raman spectroscopy to document the existence of carbonaceous matter and silica in recent and fossil agglutinated foraminifera, and to measure thermal alteration effects in fossil foraminifera. The distribution of carbonaceous matter through the test (shell) walls of agglutinated foraminifera suggests that this carbonaceous material is derived from primary organic cement and not from random contamination. Fossil specimens exhibit three broad stages of maturation: (1) Immature specimens are characterized by moderately strong fluorescence, broad, low intensity Raman peaks (relative to fluorescence), and a tendency for the G-band to occur at lower wave numbers. These attributes are consistent with the presence of amorphous carbonaceous matter and minor organic degradation. (2) Mature samples (oil window) exhibit high fluorescence, increased relative D- and G-band intensities, and a decreased width of the D-band. (3) Postmature samples exhibit low levels of fluorescence and high relative D- and G-band intensities, a tendency for the G-band to be located at higher wave numbers, an increase in the D:G band ratio, and an increase of the relative intensity of the silica peak. This stage is consistent with the presence of highly ordered carbonaceous matter and diagenetic quartz. These findings indicate that Raman spectroscopic analysis of fossil agglutinated foraminifera can be used as a quick and easy tool to assess thermal maturity and estimate optimal temperatures for hydrocarbon generation.

Neoproterozoic carbonate successions provide a new taphonomic window into the diversification of eukaryotes. We report recently discovered macroscopic organic warty sheets (MOWS) in macerates of limestone from the ca. 662–635 Ma Taishir Formation (Tsagaan Olom Group, Mongolia). Sheets are applanate. One surface contains raised ridges and conspicuous, ∼ 100-µm-tall warty protuberances with depressed tops that enclose internal cavities containing cellular structures. The Taishir MOWS may be the remains of unusual bacterial, protistan, or fungal biofilms, or a previously undocumented, extinct taxon. However, multiple lines of evidence including the morphology of warty protuberances and the presence of cellular architecture within protuberances support the interpretation of MOWS as marine algae, perhaps a member of the Rhodophyta. Regardless of their specific taxonomic affiliation, MOWS increase the diversity of biota reported from the Cryogenian glacial interlude and indicate the presence of macroscopic and morphologically complex multicellular organisms in the Cryogenian.

The Cryogenian was a time of climatic extremes, with two extended and severe global glaciations bracketing hothouse conditions. The effect of these extreme climate conditions on ocean chemistry and the marine biosphere remain poorly understood. Most of the previous studies of the fossil record from this interval focus on benthic organisms, with few examples of organisms with an inferred planktonic lifestyle and no firm evidence for photosynthetic organisms. Here, we present helically coiled, straight, and curved fossils composed of fine crystalline or framboidal pyrite in limestone samples from the Ikiakpuk formation of Arctic Alaska. These structures are morphologically identical to fossils of Obruchevella, a cyanobacterial form genus reported in pre-Sturtian and post-Marinoan strata, but not in deposits from the Cryogenian non-glacial interlude. We interpret fossils of the Ikiakpuk formation as planktonic cyanobacteria based on their morphology, which is identical to that of some modern planktonic cyanobacteria. Further evidence for a planktonic lifestyle comes from the preservation of these pyritized fossils in deep-water facies that lack evidence of microbial lamination. They provide the first direct evidence for bacterial primary productivity in the pelagic realm during the Cryogenian non-glacial interlude. Keywords: Alaska; metasomatism; paleoecology; United States; Cryogenian; Neoproterozoic; Proterozoic; upper Precambrian; Precambrian, Simons Foundation (Grant 344707)

We investigated the influence of organic substrates and phosphate concentration on the rates of dissimilatory microbial sulfate reduction and the 34S/32S isotopic fractionation produced by several Desulfovibrio species. Our experiments corroborate the previously reported species-specific correlation between sulfur isotope fractionation and cell-specific sulfate reduction rates. We also identify cell size as a key factor that contributes to the species-effect of this correlation. Phosphate limitation results in larger cells and contributes to a small decrease in sulfur isotope fractionation concomitant with an apparent increase in cell-specific sulfate reduction rates. Sulfur isotope fractionation in phosphate-limited cultures asymptotically approaches a lower limit of approximately 5‰ as cell-specific sulfate reduction rates increase to >100 fmol cell−1 day−1. These experimental results test models that link the reversibilities of enzymatic steps in dissimilatory sulfate reduction to sulfur isotope fractionation and show that these models can provide consistent predictions across large variations in physiological states experienced by sulfate reducing bacteria.