Microbial palaeontology is largely reliant on the interpretation of geologically stable biomarkers or molecular fossils. Biomolecules that are both specific to particular groups of organisms and stable on a geological scale are invaluable for tracing the emergence and diversification of lifeforms, particularly in cases where mineral fossils are lacking. 2-Methylhopanoids and their diagenic product, 2-methylhopanes, are highly abundant bacterial membrane lipids, recoverable from samples in excess of a billion years old. In this work we used degenerate PCR, targeting 2-methylhopanoid biosynthesis genes, and sequencing to show that the ability to produce these molecules in arid biological soil crusts from deserts in diverse geographical locations (Utah, USA, and the Pilbara, Australia) is largely confined to cyanobacteria. These data suggest that 2-methylhopanes can be used as a proxy for cyanobacterial presence within these environments, contributing to our understanding of the emergence of terrestrial life on Earth.

Identifying microbial fossils in the rock record is a difficult task because they are often simple in morphology and can be mimicked by non-biological structures. Biosignatures are essential for identifying putative fossils as being definitively biological in origin, but are often lacking due to geologic effects which can obscure or erase such signs. As such, there is a need for robust biosignature identification techniques. Here we show new evidence for the application of trace elements as biosignatures in microfossils. We found elevated concentrations of magnesium, aluminum, manganese, iron, and strontium colocalized with carbon and sulfur in microfossils from Drummond Basin, a mid-Paleozoic hot spring deposit in Australia. Our results also suggest that trace element sequestrations from modern hot spring deposits persist through substantial host rock alteration. Because some of the oldest fossils on Earth are found in hot spring deposits and ancient hot spring deposits are also thought to occur on Mars, this biosignature technique may be utilized as a valuable tool to aid in the search for extraterrestrial life.

The ancient rocks of the Pilbara region of Western Australia have been an important analog site for the study of possible inhabited environments in the search for life on early Mars for over four decades. Here, we review the evidence for Paleo- to Neoarchean life and the habitats that it occupied in the Pilbara Craton and unconformably overlying Fortescue Group of the Mount Bruce Supergroup. Nine major inhabited environments are described, which range from land to sea, and into the subsurface, showing that life had diversified into, and flourished within, a range of different environments early in Earth history. An important additional component in the search for life on Mars involves the manner in which evidence for early life is preserved. From the examples studied here, early mineralization of organic matter is key to the preservation of reliable biosignatures, in either silica, carbonate, or pyrite, but burial by volcanic ash can also provide excellent preservation.

Earth and Space Science Open Archive PosterOpen AccessYou are viewing the latest version by default [v1]Elemental Relics: Biosignatures for Microbial Life in Terrestrial Hot Springs on Ancient Earth and MarsAuthorsAndrewGangidineMalcolmWalterJeffHavigAndrewCzajaDanielSturmeriDJeffreyHannonSee all authors Andrew GangidineCorresponding Author• Submitting AuthorCranbrook Institute of Scienceview email addressThe email was not providedcopy email addressMalcolm WalterUniversity of New South Walesview email addressThe email was not providedcopy email addressJeff HavigUniversity of Minnesotaview email addressThe email was not providedcopy email addressAndrew CzajaUniversity of Cincinnati Main Campusview email addressThe email was not providedcopy email addressDaniel SturmeriDUniversity of Cincinnati Main CampusiDhttps://orcid.org/0000-0002-9997-6702view email addressThe email was not providedcopy email addressJeffrey HannonUniversity of Wisconsin Madisonview email addressThe email was not providedcopy email address

Abstract Terrestrial thermal environments can serve as analogs for subsurface environments in the search for life because they regularly host microbial communities, which may leave behind biosignatures. This study focused on an acid-sulfate hydrothermal site as an analog for a potentially habitable environment on Mars. A weathered boulder in the thermal area was dissected, revealing an interior marked with disconnected horizons of differently colored materials, very low pH, and increasing temperature. The mineralogy comprised weathering products from andesite (kaolinite, quartz, clinoptilolite) along with sulfate salts (alunite, jarosite, tschermigite, and copiapite) formed by oxidation of sulfide and ferrous iron. Characterization of organic matter in this boulder and several soil samples yielded interesting but surprising results. Both mass spectrometry and Raman spectroscopy identified organic compounds in portions of the soils and the boulder. Jarosite-associated samples showed more numerous and diverse organic signatures than did Al-bearing silicate samples, despite the lower total organic carbon content of the jarosite-associated soils (0.69 ± 0.07 wt% Corg) compared to the Al-bearing samples (1.28 ± 0.13 wt% Corg). Results from our geochemical, mineralogical, and spectroscopic study of hydrothermal alteration products and salts inform the heterogeneous distribution of inorganic and organic materials that could delineate habitats and demonstrate the limits on organic matter detectability using different analytical techniques. Furthermore, we relate our measurements and results directly to current and upcoming martian missions, and we provide recommendations for detection and characterization of minerals and organics as biosignatures on Mars using instruments on future missions.

Recent discoveries of geyserite and siliceous sinter with textural biosignatures in the ∼3.5 Ga Dresser Formation of the Pilbara Craton, Western Australia, extended the record of inhabited subaerial hot springs on Earth by ∼3 billion years, back to the time when siliceous sinter deposits are known to have formed on Mars (e.g., at Columbia Hills, Gusev Crater). Here, we present more detailed lithostratigraphic, petrographic and geochemical data collected from 100 measured sections across a ∼14 km strike length in the Dresser Formation. The data indicate deposition of a wide range of hot spring and associated deposits in a restricted interval that directly overlies a hydrothermally influenced volcanic caldera lake facies, with shoreline stromatolites. Hot spring deposits show abrupt lateral facies changes and include associated channelized clastic deposits that support fluvial, subaerial hot spring deposition. All Dresser hot spring and associated lithofacies have direct analogs with proximal, middle, and distal apron hot spring facies that are characteristic of those from New Zealand, Yellowstone National Park, USA, and Argentina. Rare earth element and yttrium geochemistry shows that the Dresser geyserite shares identical patterns with Phanerozoic hot spring sinters. This geochemical data further supports textural and contextual evidence that indicate the Dresser geyserite formed as a subaerial hot spring sinter. Further, the Dresser hot spring deposits are temporally associated with a diverse suite of textural biosignatures that indicate a thriving microbial community existed within in a Paleoarchean hot spring field. The results presented here underscore the importance of continued study of the early geological record for astrobiological research. In particular these findings reinforce the long-standing hypothesis that hydrothermal systems are optimal places to search for past life on Mars.

The earliest evidence of life captured in lithified microbial mats (microbialites) predates the onset of oxygen production and yet, modern oxygenic mats are often studied as analogs based on their morphological similarity and their sedimentological and biogeochemical context. Despite their structural similarity to fossil microbialites, the presence of oxygen in most modern microbial mats disqualifies them as appropriate models for understanding early Earth conditions. Here we describe the geochemistry, element cycling and lithification potential of microbial mats that thrive under permanently anoxic conditions in arsenic laden, sulfidic waters feeding Laguna La Brava, a hypersaline lake in the Salar de Atacama of northern Chile. We propose that these anoxygenic, arsenosulfidic, phototrophic mats are a link to the Archean because of their distinctive metabolic adaptations to a reducing environment with extreme conditions of high UV, vast temperature fluctuations, and alkaline water inputs from combined meteoric and volcanic origin, reminiscent of early Earth.

This report reviews how terrestrial hot spring systems can sustain diverse and abundant microbial communities and preserve their fossil records. Hot springs are dependable water sources, even in arid environments. They deliver reduced chemical species and other solutes to more oxidized surface environments, thereby providing redox energy and nutrients. Spring waters have diverse chemical compositions, and their outflows create thermal gradients and chemical precipitates that sustain diverse microbial communities and entomb their remnants. These environments probably were important habitats for ancient benthic microbial ecosystems, and it has even been postulated that life arose in hydrothermal systems. Thermal spring communities are fossilized in deposits of travertine, siliceous sinter, and iron minerals (among others) that are found throughout the geological record back to the oldest known well-preserved rocks at 3.48 Ga. Very few are known before the Cenozoic, but it is likely that there are many more to be found. They preserve fossils ranging from microbes to trees and macroscopic animals. Features on Mars whose morphological and spectroscopic attributes resemble spring deposits on Earth have been detected in regions where geologic context is consistent with the presence of thermal springs. Such features represent targets in the search for evidence of past life on that planet.

Ooids are accretionary grains commonly reported from turbulent, shallow‐water environments. They have long been associated with microbially dominated ecosystems and often occur in close proximity to, or embedded within, stromatolites, yet have historically been thought to form solely through physicochemical processes. Numerous studies have revealed both constructive and destructive roles for microbes colonizing the surfaces of modern calcitic and aragonitic ooids, but there has been little evidence for the operation of these processes during the Archean and Proterozoic, when both ooids and microbially dominated ecosystems were more widespread. Recently described carbonate ooids from the 2.9 Ga Pongola Supergroup, South Africa, include well‐preserved examples composed of diagenetic dolomite interpreted to have formed from a high‐Mg‐calcite precursor. Spatial distributions of organic matter and elements associated with metabolic activity (N, S, and P) were interpreted as evidence for a biologically induced origin. Here, we describe exceptionally well‐preserved ooids composed of calcite, collected from Earth's oldest known carbonate lake system, the ~2.72 Ga Meentheena Member (Tumbiana Formation), Fortescue Group, Western Australia. We used optical microscopy, Raman spectroscopy, XRD , SEM ‐ EDS , LA ‐ ICP ‐ MS , EA ‐ IRMS , and a novel micro‐ XRF instrument to investigate an oolite shoal deposited between stromatolites that preserve abundant evidence for microbial activity. We report an extremely fine, radial‐concentric, calcitic microfabric that is similar to the primary and early diagenetic fabrics of calcitic ooids reported from modern temperate lakes. Early diagenetic silica has trapped isotopically light and thermally mature organic matter. The close association of organic matter with mineral phases and microfabrics related to primary and early diagenetic processes suggest incorporation of organic matter occurred during accretion, likely due to the presence of microbial biofilms. We conclude that the oldest known calcitic ooids were likely formed through processes similar to those that mediate the accretion of ooids in similar environments today, including formation within a microbial biosphere.

Early in Mars’ history, life may have flourished in lacustrine environments developed during a comparatively warmer and wetter climate regime. Sustained subaqueous environments are high priority landing sites for the Mars 2020 mission, which is likely to focus on in situ analysis of geological features present in these environments, and on sample return. However, the community lacks experience investigating ancient, microbially dominated lacustrine environments, in part due to their rarity. Among the rocks that were formed during the Archean and that host biosignatures, only the Fortescue Group (2775–2630 Ma) is known to contain well-preserved lake deposits. Intervals within this group preserve extensive fluvio-lacustrine depositional facies and several categories of biosignatures recording the presence of early, microbially dominated ecosystems. The broader depositional environment of lacustrine units within the Fortescue Group, which formed on plateaus renewed by episodes of ongoing basaltic volcanism, is analogous to the broader depositional setting of paleolakes that may be encountered on Mars. Repeated lake high stands and drying events provide an analogue for climate change related to obliquity variations in early Martian environments, insights into the effect of climate changes on the lacustrine microbial biosphere, and an exploration model for taphonomic windows preserving microbial biosignatures in elements of these systems.

Nature Communications 8: Article number: 15263 (2017); Published: 9 May 2017; Updated: 16 August 2017. The original manuscript presented evidence that the deposits studied were fluvial rather than marine to support the conclusion that evidence of life on land had been discovered in the 3.4 Ga Dresser Formation, Pilbara Craton, Australia.

Assessing the habitability of Mars and detecting life, if it ever existed there, depends on knowledge of whether the combined environmental stresses experienced on Mars are compatible with life as we know it and whether a record of that life could ever be detected. So far, only few investigations were performed to understand the combined effect of different environmental stresses on survival and growth of anaerobic and extremophilic organisms. In the space experiment MEXEM (formerly known as MASE-in-SPACE) the hypothesis will be tested that selected terrestrial organisms, enrichment cultures and original samples from extreme Mars-analogue environments on Earth are able to withstand the Martian environ- mental stress factors due to their highly effective cellular and molecular adaptation and repair mechanisms. In addition, artificially fossilized and aged isolates from Mars-analogue environments on Earth will be examined and assessed with respect to their suitability for biosigna- ture identification. MEXEM samples will be (i) oxygen-depleted natural sediment samples, (ii) natural sediments spiked with selected, defined strains representative for the respective analogue site, (iii) individual (facultative) anaerobic / micro-aerophilic species including ciliates and viruses, (iv) defined mixtures of these biological entities, (v) isolated strains from samples collected inside the ISS and (vi) artificially fossilized isolates from the natural environments. Most of these samples and isolates were obtained from Mars-analogue envi- ronments on Earth in the frame of the EC funded project MASE (Grant Agreement 607297) and from the space experiment EXTREMOPHILES (PI C. Moissl-Eichinger). MEXEM will be flown outside on the ISS in the new exobiology facility ESA is building now. It offers the possibility to simulate of the martian environment, in particular the martian UV cl e, which cannot be done in the lab, but also martian atmosphere and pressure in LEO.

The ca. 3.48 Ga Dresser Formation, Pilbara Craton, Western Australia, is well known for hosting some of Earth's earliest convincing evidence of life (stromatolites, fractionated sulfur/carbon isotopes, microfossils) within a dynamic, low-eruptive volcanic caldera affected by voluminous hydrothermal fluid circulation. However, missing from the caldera model were surface manifestations of the volcanic-hydrothermal system (hot springs, geysers) and their unequivocal link with life. Here we present new discoveries of hot spring deposits including geyserite, sinter terracettes and mineralized remnants of hot spring pools/vents, all of which preserve a suite of microbial biosignatures indicative of the earliest life on land. These include stromatolites, newly observed microbial palisade fabric and gas bubbles preserved in inferred mineralized, exopolymeric substance. These findings extend the known geological record of inhabited terrestrial hot springs on Earth by ∼3 billion years and offer an analogue in the search for potential fossil life in ancient Martian hot springs.

ABSTRACT To investigate the function of 2-methylhopanoids in modern cyanobacteria, the hpnP gene coding for the radical S -adenosyl methionine (SAM) methylase protein that acts on the C-2 position of hopanoids was deleted from the filamentous cyanobacterium Nostoc punctiforme ATCC 29133S. The resulting Δ hpnP mutant lacked all 2-methylhopanoids but was found to produce much higher levels of two bacteriohopanepentol isomers than the wild type. Growth rates of the Δ hpnP mutant cultures were not significantly different from those of the wild type under standard growth conditions. Akinete formation was also not impeded by the absence of 2-methylhopanoids. The relative abundances of the different hopanoid structures in akinete-dominated cultures of the wild-type and Δ hpnP mutant strains were similar to those of vegetative cell-dominated cultures. However, the Δ hpnP mutant was found to have decreased growth rates under both pH and osmotic stress, confirming a role for 2-methylhopanoids in stress tolerance. Evidence of elevated photosystem II yield and NAD(P)H-dependent oxidoreductase activity in the Δ hpnP mutant under stress conditions, compared to the wild type, suggested that the absence of 2-methylhopanoids increases cellular metabolic rates under stress conditions. IMPORTANCE As the first group of organisms to develop oxygenic photosynthesis, Cyanobacteria are central to the evolutionary history of life on Earth and the subsequent oxygenation of the atmosphere. To investigate the origin of cyanobacteria and the emergence of oxygenic photosynthesis, geobiologists use biomarkers, the remnants of lipids produced by different organisms that are found in geologic sediments. 2-Methylhopanes have been considered indicative of cyanobacteria in some environmental settings, with the parent lipids 2-methylhopanoids being present in many contemporary cyanobacteria. We have created a Nostoc punctiforme Δ hpnP mutant strain that does not produce 2-methylhopanoids to assess the influence of 2-methylhopanoids on stress tolerance. Increased metabolic activity in the mutant under stress indicates compensatory alterations in metabolism in the absence of 2-methylhopanoids.

This corrects the article DOI: 10.1038/ncomms15263.

The island of Seram, eastern Indonesia, experienced a complex Neogene history of multiple metamorphic and deformational events driven by Australia–SE Asia collision. Geological mapping, and structural and petrographic analysis has identified two main phases in the island's tectonic, metamorphic, and magmatic evolution: (1) an initial episode of extreme extension that exhumed hot lherzolites from the subcontinental lithospheric mantle and drove ultrahigh-temperature metamorphism and melting of adjacent continental crust; and (2) subsequent episodes of extensional detachment faulting and strike-slip faulting that further exhumed granulites and mantle rocks across Seram and Ambon. Here we present the results of sixteen 40Ar/39Ar furnace step heating experiments on white mica, biotite, and phlogopite for a suite of twelve rocks that were targeted to further unravel Seram's tectonic and metamorphic history. Despite a wide lithological and structural diversity among the samples, there is a remarkable degree of correlation between the 40Ar/39Ar ages recorded by different rock types situated in different structural settings, recording thermal events at 16 Ma, 5.7 Ma, 4.5 Ma, and 3.4 Ma. These frequently measured ages are defined, in most instances, by two or more 40Ar/39Ar ages that are identical within error. At 16 Ma, a major kyanite-grade metamorphic event affected the Tehoru Formation across western and central Seram, coincident with ultrahigh-temperature metamorphism and melting of granulite-facies rocks comprising the Kobipoto Complex, and the intrusion of lamprophyres. Later, at 5.7 Ma, Kobipoto Complex rocks were exhumed beneath extensional detachment faults on the Kaibobo Peninsula of western Seram, heating and shearing adjacent Tehoru Formation schists to form Taunusa Complex gneisses. Then, at 4.5 Ma, 40Ar/39Ar ages record deformation within the Kawa Shear Zone (central Seram) and overprinting of detachment faults in western Seram. Finally, at 3.4 Ma, Kobipoto Complex migmatites were exhumed on Ambon, at the same time as deformation within the Kawa Shear Zone and further overprinting of detachments in western Seram. These ages support there having been multiple synchronised episodes of high-temperature extension and strike-slip faulting, interpreted to be the result of Western Seram having been ripped off from SE Sulawesi, extended, and dragged east by subduction rollback of the Banda Slab.

Significance An ancient deep-sea mud-inhabiting 1,800-million-year-old sulfur-cycling microbial community from Western Australia is essentially identical both to a fossil community 500 million years older and to modern microbial biotas discovered off the coast of South America in 2007. The fossils are interpreted to document the impact of the mid-Precambrian increase of atmospheric oxygen, a world-changing event that altered the history of life. Although the apparent 2-billion-year-long stasis of such sulfur-cycling ecosystems is consistent with the null hypothesis required of Darwinian evolution—if there is no change in the physical-biological environment of a well-adapted ecosystem, its biotic components should similarly remain unchanged—additional evidence will be needed to establish this aspect of evolutionary theory.