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The current strategy for detecting evidence of ancient life on Mars-a primary goal of NASA's ongoing Mars 2020 mission-is based largely on knowledge of Precambrian life and of its preservation in Earth's early rock record. The fossil record of primitive microorganisms consists mainly of stromatolites and other microbially influenced sedimentary structures, which occasionally preserve microfossils or other geochemical traces of life. Raman spectroscopy is an invaluable tool for identifying such signs of life and is routinely performed on Precambrian microfossils to help establish their organic composition, degree of thermal maturity, and biogenicity. The Mars 2020 rover, Perseverance, is equipped with a deep-ultraviolet (UV) Raman spectrometer as part of the SHERLOC (Scanning Habitable Environments with Raman and Luminescence for Organics and Chemicals) instrument, which will be used in part to characterize the preservation of organic matter in the ancient sedimentary rocks of Jezero crater and therein search for possible biosignatures. To determine the deep-UV Raman spectra characteristic of ancient microbial fossils, this study analyzes individual microfossils from 14 Precambrian cherts using deep-UV (244 nm) Raman spectroscopy. Spectra obtained were measured and calibrated relative to a graphitic standard and categorized according to the morphology and depositional environment of the fossil analyzed and its Raman-indicated thermal maturity. All acquired spectra of the fossil kerogens include a considerably Raman-enhanced and prominent first-order Raman G-band (∼1600 cmsup-1/sup), whereas its commonly associated D-band (∼1350 cmsup-1/sup) is restricted to specimens of lower thermal maturity (below greenschist facies) that thus have the less altered biosignature indicative of relatively well-preserved organic matter. If comparably preserved, similar characteristics would be expected to be exhibited by microfossils or ancient organic matter in rock samples collected and cached on Mars in preparation for future sample return to Earth.

The surface ecosystem of the Precambrian Earth was dominated by marine, planktonic and benthic phototrophic microorganisms, most prominently stromatolite-forming cyanobacteria, anoxygenic photosynthetic bacteria, and associated microbes. Although coupling the early microfossil record to physiologically definitive geochemical and isotopic signatures remains challenging, insights can be derived from isotopic studies of individual fossil microorganisms permineralized in Precambrian cherts. Here, we use correlated optical microscopy, Raman spectroscopy, and secondary ion mass spectrometry (SIMS) to link morphology with the molecular and carbon isotopic composition (δ13Corg) of individual filamentous microfossils (Eomycetopsis sp.) composed of thermally immature kerogen and preserved in two spatially and temporally distinct shallow-marine cherts of the Proterozoic Gaoyuzhuang Formation (∼1,560 Ma, northern China) and Kwagunt Formation (∼850 Ma, Arizona, USA). In both geologic units studied, the Eomycetopsis fossils yielded virtually indistinguishable δ13Corg values (–29.0 ± 2.0‰) that are comparable to values resulting from carbon fixation via the RuBisCO enzyme in the Calvin cycle of oscillatoriacean cyanobacteria, the main oxygenic phototrophs of the Proterozoic.

Analyses by secondary ion mass spectroscopy (SIMS) of 11 specimens of five taxa of prokaryotic filamentous kerogenous cellular microfossils permineralized in a petrographic thin section of the ∼3,465 Ma Apex chert of northwestern Western Australia, prepared from the same rock sample from which this earliest known assemblage of cellular fossils was described more than two decades ago, show their δ13C compositions to vary systematically taxon to taxon from -31‰ to -39‰. These morphospecies-correlated carbon isotope compositions confirm the biogenicity of the Apex fossils and validate their morphology-based taxonomic assignments. Perhaps most significantly, the δ13C values of each of the five taxa are lower than those of bulk samples of Apex kerogen (-27‰), those of SIMS-measured fossil-associated dispersed particulate kerogen (-27.6‰), and those typical of modern prokaryotic phototrophs (-25 ± 10‰). The SIMS data for the two highest δ13C Apex taxa are consistent with those of extant phototrophic bacteria; those for a somewhat lower δ13C taxon, with nonbacterial methane-producing Archaea; and those for the two lowest δ13C taxa, with methane-metabolizing γ-proteobacteria. Although the existence of both methanogens and methanotrophs has been inferred from bulk analyses of the carbon isotopic compositions of pre-2,500 Ma kerogens, these in situ SIMS analyses of individual microfossils present data interpretable as evidencing the cellular preservation of such microorganisms and are consistent with the near-basal position of the Archaea in rRNA phylogenies.

A microbial consortium preserved in shallow water carbonaceous chert of the Paleoarchean (∼3400 Ma) Strelley Pool Formation (Pilbara Craton, northwestern Western Australia) is composed of copiously abundant globular “cobweb-like” swirls of randomly oriented filaments intermingled with laterally extensive flat-laminated masses of subparallel filaments. The shallow-water (possibly mudflat-like) habitat of the biocoenose is established by its geological setting and intimately intermixed association with millimetric quartz radiating laths and rosettes interpreted to be pseudomorphs after evaporitic gypsum/anhydrite. The indigenousness and syngenicity of the permineralized fossils to the encompassing chert; their uncompacted minimally altered “life-like” spatial distribution; and their biogenicity and bacterial affinity are shown by optical microscopic, Raman spectroscopic and confocal laser scanning microscopic analyses in situ that document the mineralogy of their embedding matrix and their cylindrical filamentous morphology, cellularity, and kerogenous composition. By comparison with mid-Precambrian and modern sulfur-cycling ecosystems, and consistent with sulfur isotopic data and their intimate co-existence with copious microgranular pyrite and pseudomorphs evidently after sulfate, the swirled aggregates are interpreted to be anaerobic sulfuretums. In contrast, the intermixed flat-laminated components, like microbes of fossil and modern stromatolitic mat communities, are interpreted to be co-occurring light-requiring phototrophs. Whether such phototrophs were anoxygenic photosynthesizers, like extant photosynthetic bacteria (e.g., chloroflexaleans) or facultatively anaerobic cyanobacteria (cf. Oscillatoria limnetica), their anoxygenic metabolism seems plausibly established. Given its great age, shallow water habitat, and anaerobic/microaerophilic physiology, this evidently first reported occurrence from the fossil record of anoxygenic phototrophs and of a microbial sulfuretum-phototroph consortium is consistent with the supposition that Earth’s Paleoarchean environment was essentially anoxic and suggests that O2-producing photosynthesis may have been a later evolutionary innovation.

Decreasing pH levels in the world’s oceans are widely recognized as a threat to marine life. Bryozoans are among several phyla that produce calcium carbonate skeletons potentially affected by ocean acidification (OA). Depending on species, bryozoan skeletons can consist of calcite, aragonite or have a bimineralic combination of these two minerals. Aragonite is generally more soluble in seawater than calcite, making aragonitic species more vulnerable to OA. Here, for the first time we use Raman spectroscopy to determine the mineral composition of a tropical bryozoan biota. Compared with bryozoan biotas from higher latitudes in which calcite predominates, aragonite was found to occur in a much higher proportion of the 22 cheilostome bryozoan species collected from the shorelines of Penang and Langkawi in Malaysia, where 46% of species are calcitic, 41% aragonitic and 13% bimineralic. All but one of the aragonitic or bimineralic species belong to the ascophorans, whereas calcitic skeletons characteri...

Fluorescence signatures of samarium3+ substituted into apatite for calcium2+ have been shown experimentally to correlate with oxygen availability, such substituted apatite being a principal diagenetic mineral emplaced in shallow-water environments during the Neoproterozoic-Cambrian global phosphogenic event coeval with the “Cambrian Explosion of Life.” The occurrence of Sm3+-substituted apatite, encrusting, infilling and permineralizing components of shallow water microbial communities suggested that quantification of these fluorescence signatures might provide a widely applicable mineral-based paleobarometer by which to measure oxygen concentrations in the environment in which such apatite precipitated. To test this hypothesis, fluorescence spectroscopic analyses have been carried out on the Sm3+-substituted apatite of photic-zone microscopic fossils and fossil-associated ooids of near-shore facies of the Ediacaran Doushantuo (South China) and Early Cambrian Chulaktau (South Kazakhstan) Formations, and of the open ocean apatite-biomineralized phytoplanktonic “scale fossils” (Characodictyon) of the mid-Neoproterozoic (Tonian) Fifteenmile Group (Yukon Canada). Results obtained – quantified by the newly devised apatite oxygen paleobarometer metric – indicate a low (

Subtidal cherts from phosphorite-bearing basal strata of the Early Cambrian Shabakta (Dzhylandy) Formation, Tamda Group of the Maly Karatau Range of South Kazakhstan, contain a diverse assemblage of acanthomorph acritarchs and other well-preserved organic-walled microfossils. Acanthomorphs of the Shabakta lowermost layers are taxonomically essentially identical to those of many similarly aged Early Cambrian organic-walled compression-preserved and permineralized microbiotas preserved in rocks deposited in relatively open marine settings and are biostratigraphically correlative with acanthomorph-containing microfloral assemblages of the Vergale and Rausve Horizons (Regional Stages) of the East European Platform as well as with other assemblages known worldwide from the upper Atdabanian (Series 2 of the International Union of Geological Sciences [IUGS]). Although lower Atdabanian deposits of the Talsy (Lukati) horizon are missing from the Maly Karatau succession studied here, Nemakit-Daldynian through Tommotian (the IUGS Terranovian) microbiotas both of the Shabakta-underlying Berkuta Member of the Kyrshabakta Formation and phosphoritic Chulaktau Formation are comparable to those of the pre-trilobite Rovno and Lontova Horizons (Regional Stages) of the East European Platform. Thus, the phosphatized and chert-permineralized Shabakta assemblages date from of the Early Cambrian part of the Precambrian-Cambrian phosphogenic event, an interpretation supported also by associated small shelly fossils. The studies reported here of the Shabakta microfossils – based on the use of optical microscopy, confocal laser scanning microscopy (CLSM) and Raman spectroscopy – provide information in three dimensions at high spatial resolution about their organismal morphology, cellular anatomy, kerogenous composition, taphonomy, and mode and fidelity of preservation. The combined use of CLSM and Raman, techniques introduced to Precambrian paleobiology only rather recently, is shown to reveal morphological characters of taxonomic, biostratigraphic and palaeoenvironmental significance.

Peritidal cherts and silicified phosphorites of the Early Cambrian Kyrshabakta (Berkuta Member) and Chulaktau formations of South Kazakhstan contain diverse assemblages of cellularly permineralized microorganisms that like other microbiotas of similar age and setting are dominated by cyanobacteria, both filamentous (oscillatoriacean trichomes and empty sheaths) and coccoidal (chroococcaceans). Although these near-shore assemblages contain sphaeromorph acritarchs, they differ from Neoproterozoic and Early Cambrian microbiotas of more open-marine environments by lacking morphologically complex planktonic eukaryotes such as the acanthomorphic acritarchs that are abundant in the Chulaktau-overlying Shabakta Formation. In general composition, the Berkuta and Chulaktau assemblages are similar both to microfossil assemblages of the pre-trilobite Rovno and Lontova Regional Stages of the East European Platform and to Early Cambrian microbiotas known from chert-phosphorite deposits worldwide.The studies reported here are based on use of optical microscopy combined with techniques recently introduced to paleobiology: confocal laser scanning microscopy, and Raman and fluorescence spectroscopy and imagery. Taken together, these provide information in situ, in three dimensions and at high spatial resolution, about the organismal morphology; cellular anatomy; chemical composition; and mode, fidelity, and environment of preservation of the permineralized microfossils. Data are presented suggesting that the substitution of samarium for calcium in fossil-preserving apatite may provide evidence of its oxic or anoxic paleoenvironment of formation.We interpret the Berkuta and Chulaktau assemblages to be composed of 27 taxa assigned to 17 genera of microscopic prokaryotes and eukaryotes that include one new genus and species, Berkutaphycus elongatus.

Paleotemperatures inferred from the isotopic compositions (δ18O and δ30Si) of marine cherts suggest that Earth's oceans cooled from 70 ± 15 °C in the Archean to the present ∼15 °C. This interpretation, however, has been subject to question due to uncertainties regarding oceanic isotopic compositions, diagenetic or metamorphic resetting of the isotopic record, and depositional environments. Analyses of the thermostability of reconstructed ancestral enzymes provide an independent method by which to assess the temperature history inferred from the isotopic evidence. Although previous studies have demonstrated extreme thermostability in reconstructed archaeal and bacterial proteins compatible with a hot early Earth, taxa investigated may have inhabited local thermal environments that differed significantly from average surface conditions. We here present thermostability measurements of reconstructed ancestral enzymatically active nucleoside diphosphate kinases (NDKs) derived from light-requiring prokaryotic and eukaryotic phototrophs having widely separated fossil-based divergence ages. The ancestral environmental temperatures thereby determined for these photic-zone organisms--shown in modern taxa to correlate strongly with NDK thermostability--are inferred to reflect ancient surface-environment paleotemperatures. Our results suggest that Earth's surface temperature decreased over geological time from ∼65-80 °C in the Archean, a finding consistent both with previous isotope-based and protein reconstruction-based interpretations. Interdisciplinary studies such as those reported here integrating genomic, geologic, and paleontologic data hold promise for providing new insight into the coevolution of life and environment over Earth history.

Vase-shaped microfossils (VSMs) occur in dolomitic extraclasts of indeterminate provenance within the basal diamictite of the Neoproterozoic Urucum Formation (Jacadigo Group) of west-central Brazil, having an age constrained between 889±44 Ma (K-Ar; basement rocks) and 587±7 Ma (40Ar/39Ar age of early metamorphic cryptomelane in overlying manganese ore). Early isopachous carbonate cement entombed these VSMs, preserving rare direct evidence of original wall composition that is carbonaceous (now kerogenous) in practically all specimens. Some tests are siliceous or composed of a quartz-kerogen mixture; secondary replacement explains some features of these tests, but original biomineralization seems more likely for others. This interpretation, coupled with test morphology, suggests affinity to arcellinid testate amoebae. Five VSM taxa are recognized in the deposit:Cycliocyrillium simplexPorter, Meisterfeld, and Knoll, 2003, andC.torquataPorter, Meisterfeld, and Knoll, 2003, originally described in the Chuar Group (USA), and three new monospecific genera—Palaeoamphora urucumensen. gen. n. sp.,Limeta lageniformisn. gen. n. sp., andTaruma ratan. gen. n. sp. Most of the taxonomically important characteristics of these VSMs occur also in extant testate amoebae, but the combinations of some characters, such as organic-walled tests having exceptionally long necks that exhibit terminal apertures (L.lageniformisn. gen. n. sp.), are evidently novel additions to the known diversity of Neoproterozoic VSMs. Evidence of glacially influenced deposition in the conformably overlying Santa Cruz Formation may indicate that the Urucum Formation slightly preceded or was penecontemporaneous with a major Neoproterozoic glaciation, although the VSM-hosting extraclasts must be older, possibly rivaling the age of the testate amoebae of the Chichkan Formation (766±7 Ma) that are currently regarded as the oldest record of protists in the geological record.

Although the carbonate compensation depth (CCD) for calcite, generally located in the depth range 4000–5000 m, is often proposed as a physiological barrier to deep-ocean colonization, many organisms with calcareous exoskeletons are found in the deepest oceanic trenches. Serpulid polychaetes inhabiting unprotected calcareous tubes are unlikely deep-sea inhabitants, yet, they are found at all oceanic depths from intertidal to hadal. Here we review and revise the published and unpublished records of Serpulidae from below 5000 m depth. We also describe tube ultrastructure and mineralogical content of available deep-sea serpulid tubes to obtain insights into their biomineralisation. Species belonging to the genera Bathyditrupa, Bathyvermilia, Hyalopomatus, Pileolaria (spirorbin) and Protis were found at depths from 5020 to 9735 m. However, only specimens of Protis sp. were truly hadal (>6000 m) being found at 6200–9700 m. Hadal specimens of Protis have irregularly oriented prismatic tube microstructure similar to that found in more shallow-water representatives of the genus. Initial EDX analysis suggested a mostly calcitic composition (i.e., the most stable CaCO3 polymorph) on the basis of high Mg levels. Surprisingly, however, tubes of Bathyditrupa hovei and a species of Protis analysed using the more reliable method of laser Raman spectroscopy were found to be composed of aragonite. The compensation depth for this less stable CaCO3 polymorph in the oceans is usually 2000–3000 m. We found no obvious structural adaptations to life at extreme depths in the studied serpulid tubes and how serpulids are able to biomineralise and maintain their tubes below the CCD remains to be explained.

Fossil evidence of photosynthesis, documented in the geological record by microbially laminated stromatolites, microscopic fossils, and carbon isotopic data consistent with the presence of Rubisco-mediated CO2-fixation, extends to ~3500 million years ago. Such evidence, however, does not resolve the time of origin of oxygenic photosynthesis from its anoxygenic photosynthetic evolutionary precursor. Though it is evident that cyanobacteria, the earliest-evolved O2-producing photoautotrophs, existed before ~2450 million years ago - the onset of the "Great Oxidation Event" (GOE) that forever altered Earth's environment - O2-producing photosynthesis seems certain to have originated hundreds of millions of years earlier. How did Earth's biota respond to the GOE? Four lines of evidence are here suggested to reflect this major environmental transition: (1) rRNA phylogeny-correlated metabolic and biosynthetic pathways document evolution from an anaerobic (pre-GOE) to a dominantly oxygen-requiring (post-GOE) biosphere; (2) consistent with the rRNA phylogeny of cyanobacteria, their fossil record evidences the immediately post-GOE presence of cyanobacterial nostocaceans characterized by specialized cells that protect their oxygen-labile nitrogenase enzyme system; (3) the earliest known fossil eukaryotes, obligately aerobic phytoplankton and putative algae, closely post-date the GOE; and (4) microbial sulfuretums are earliest known from rocks deposited during and immediately after the GOE, their apparent proliferation evidently spurred by an increase of environmental oxygen and a resulting upsurge of metabolically useable sulfate and nitrate. Though the biotic response to the GOE is a question new to paleobiology that is yet largely unexplored, additional evidence of its impact seems certain to be uncovered.

During the Sino-Japanese conflict of the 1920s, Japanese botanist Ichiro Ohga was presented single-seeded fruit of Nelumbo nucifera var. China Antique collected by a local farmer from a dry lakebed in Northeast China (then, “Manchuria”). Ohga studied the fruit and published his findings. Years later, we tested the germination of Nelumbo fruit from the same locality. The oldest seed sprouted, having a germination time of ~3 days, was radiocarbon dated to be ~1300 years old. These cold- and drought-tolerant seeds exhibited shoot-before-root emergence and a primary green plumule capable of “dim-light” photosynthesis. Such traits and the notable long-term viability of the fruit spurred the interest of Ray Ming, University of Illinois that has now led to the sequencing of the Nelumbo genome. Analyses of this genome may provide insight into the biochemistry of Nelumbo on wax-biosynthesis genes, and application of aging-related thermostable proteins to the extension of seed-life and improvement of food quality of economic crops. Here, we review the history of these long-lived Nelumbo fruit, and their occurrence, discovery, collection, propagation, and methods of seedling care. The robust impermeable wax- and suberin-covered pericarp is a major factor contributing to their remarkable longevity. New findings are presented on the modern and 459- and 464-year-old pericarp anatomy, impermeability to water, and whole fruit and pericarp mechanical properties, and comparison of the mode of fruit weight-gain during imbibition and germination time relative to fruit maturity.