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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.

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)

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.

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)

Wrinkle structures in ancient sedimentary environments are enigmatic. Wave-tank experiments suggest that wrinkle structures are shaped by microbial mat fragments that are moved by waves over sandy-bed surfaces, and thus are morphological biosignatures.

Elongated stromatolites are often used as indicators of current direction and shoreline orientation, especially in paleoenvironmental reconstructions. However, mechanisms that create shore-parallel, m-scale elongated stromatolite mounds in carbonate sand are not well understood. We propose that this geometry is initiated by microbial growth on the parts of sand bars that experience low wave-induced bed shear stresses. We test this idea by growing microbial mats on carbonate sand bars in a laboratory wave tank. Cyanobacterial mats grow on the bar runnels, where sediment motion is negligible, but are absent from the bar ridges, where the waves generate migrating ripples. When microbially-promoted lithification reinforces and preserves this initial pattern, elongated stromatolites should initiate in the runnels of sand bars, with long wavelengths (5–100 m) and small width-to-wavelength ratios (∼0.3). These dimensions are consistent with modern shore-parallel stromatolites in Hamelin Pool, Western Australia, and with patterns of microbial colonization in other sandy sediments. This model of elongated stromatolite mounds can inform paleoenvironmental reconstructions by clarifying and quantifying feedbacks among waves, sediment transport and microbial growth.

Stromatolites document microbial interactions with sediments and flowing water throughout recorded Earth history and have the potential to illuminate the long-term history of life and environments. Modern stromatolites, however, provide analogs to only a small subset of the structures preserved in Archean and Proterozoic carbonates. Thus, interpretations of secular trends in the shapes and textures of ancient columnar stromatolites require nonuniformitarian, scale-dependent models of microbial responses to nutrient availability, seawater chemistry, influx of sediment grains, shear, and burial. Models that integrate stromatolite scales, macroscopic organization, and shapes could also help test the biogenicity of the oldest stromatolites and other structures whose petrographic fabrics do not preserve direct evidence of microbial activity. An improved understanding of stromatolite morphogenesis in the presence of oxygenic and anoxygenic microbial mats may illuminate the diversity of microbial metabolisms that contributed to stromatolite growth in early oceans.

Laminated carbonates of the Rasthof Formation, deposited in the aftermath of the early Cryogenian low-latitude glaciation (Sturtian, 717–662 Ma), preserve abundant round tests of agglutinated microscopic eukaryotes. Previously, fossil tests were reported in two localities (Ongongo and Okaaru) from microbially laminated carbonates in the Rasthof Formation, which revealed a previous unexplored Cryogenian taphonomic window. In order to better understand the lateral variability in these microfossil assemblages, this work systematically examines fossil tests from two additional localities, South Ombepera and Ombepera, and compares their preservation in thinly and thickly laminated microbial laminites. Cap carbonates in South Ombepera and Ombepera contain abundant, hollow, spheroidal agglutinated tests (50 to 225 µm in diameter). Some of these tests exhibit slitlike or triangular apertures. In contrast, much larger, oval tests with a tapering end dominate the assemblages at Okaaru, whereas oval, laterally compressed and round structures with slits, visors, or central apertures are found at Ongongo. The thinly laminated microbial laminites from Ombepera, South Ombepera, and Okaaru also preserve rare agglutinated tubes attributed to fossils of early Foraminifera. At all four localities, the thinly laminated microbial laminites preserve more microfossils than thickly laminated microbial laminites although these two facies commonly interfinger and are interbedded. This difference shows that conditions present during formation of the thinly laminated microbial laminites favored the preservation of round agglutinators, perhaps during early burial, lithification, and fossilization of the test walls.

Microbial fossils and textures are commonly preserved in Ediacaran and early Cambrian coarse-grained siliciclastic sediments that were deposited in tidal and intertidal marine settings. In contrast, the fossilization of micro-organisms in similar marine environments of post-Cambrian age is less frequently reported. Thus, temporal discrepancies in microbial preservation may have resulted from the opening and closing of a unique taphonomic window during the terminal Proterozoic and early Phanerozoic, respectively. Here, we expand upon previous work to identify environmental factors which may have facilitated the preservation of cyanobacteria growing on siliciclastic sand, by experimentally determining the ability of microbial mats to trap small, suspended mineral grains, and precipitate minerals from ions in solution. We show that (i) fine grains coat the sheaths of filamentous cyanobacteria (e.g., Nodosilinea sp.) residing within the mat, after less than 1 week of cell growth under aerobic conditions, (ii) clay minerals do not coat sterile cellulose fibers and rarely coat unsheathed cyanobacterial cells (e.g., Nostoc sp.), (iii) stronger disturbances (where culture jars were agitated at 170 rpm; 3 mm orbital diameter) produce the smoothest and most extensive mineral veneers around cells, compared with those agitated at slower rotational speeds (150 and 0 rpm), and (iv) mineral veneers coating cyanobacterial cells are ~1 μm in width. These new findings suggest that sheathed filamentous cyanobacteria may be preferentially preserved under conditions of high fluid energy. We integrate these results into a mechanistic model that explains the preservation of microbial fossils and textures in Ediacaran sandstones and siltstones, and in fine-grained siliciclastic deposits that contain exceptionally preserved microbial mats.

Major lineages of modern eukaryotes, represented primarily by microscopic taxa, are thought to have originated during the Neoproterozoic, but microfossils older than 635 Ma rarely have unambiguous relationships to modern microscopic eukaryotes. Here we report exceptionally preserved 715–635 m.y. old eukaryotic tests in limestone strata of Mongolia. The ∼100-μm-long organic-rich three-dimensional tests have flask-like shapes, constricted necks, distinct and often thickened collars, and flexible walls composed of densely packed alveolar structures. The combined morphological and ultrastructural characters of these Cryogenian tests are remarkably similar to the tests of tintinnids, modern planktonic ciliates. Eukaryotes forming recalcitrant organic or mineral-rich tests before 635 Ma may have increased export and burial fraction of organic carbon, driving an increase in atmospheric oxygen and the subsequent radiation of metazoans.

Paleomagnetic data suggest that the early Cryogenian (Sturtian) glaciation extended to sea level at low latitude. The impact of this dramatic environmental change on biota, and the composition of ecosystems in the immediate aftermath of the Sturtian glaciation remain virtually unknown. Here we report the discovery of abundant agglutinated tests in organic-rich carbonates directly overlying Sturtian glacial deposits from two different paleocontinents: the Rasthof Formation of the Congo craton in northern Namibia and the Tsagaan Oloom Formation of the Dzabkhan terrane in Mongolia. The most abundant tests preserve morphological and compositional characters consistent with those found in at least two different families of modern lobose testate amoebae (Amoebozoa), a group of heterotrophic microbial eukaryotes. The presence of spatially and compositionally variable clay minerals, quartz and microcline on the test walls is a signature of widespread biological agglutination. The post-glacial fossil assemblages differ from the most common pre-Sturtian vase-shaped fossil testate amoebae, perhaps as a result of different preservational mechanisms or of the appearance of new forms after the glaciation. The apparent local abundance of eukaryotic body fossils in the post-Sturtian carbonates suggests that the Cryogenian limestones and dolostones may host a currently unexplored fossil record of modern eukaryotes.

Microbial structures in Neoproterozoic cap carbonates record the environmental processes present in the aftermath of global glaciation. The Rasthof Formation of northern Namibia is a unique carbonate depositional sequence that formed during post-glacial transgression and highstand following the Chuos glaciation. Carbon isotope profiles from four examined localities reveal that onlap was diachronous over post-glacial, syn-rift topography. The lower Rasthof Formation consists primarily of dark gray thinly ( The close spatial association of deep-water microbialaminite facies in the Rasthof cap carbonate with carbonate clastic dikes suggests that the emplacement of dikes produced both dm-sized broad folds and cm-scale laterally discontinuous roll-up structures. The emplacement of the dikes, most likely due to the release of fluids into incompletely lithified mats, deformed cement-rich thick laminites into broad folds, while thinly laminated and more slowly lithifying mats were rolled into roll-up structures. Microbialaminite facies in the Rasthof cap carbonate thus not only reflect the depositional and environmental processes that operated in the aftermath of the Sturtian glaciation, but may also provide clues for the formation of roll-up structures found in even older Precambrian carbonates.

Stromatolites may be Earth’s oldest macroscopic fossils; however, it remains controversial what, if any, biological processes are recorded in their morphology. Although the biological interpretation of many stromatolite morphologies is confounded by the influence of sedimentation, conical stromatolites form in the absence of sedimentation and are, therefore, considered to be the most robust records of biophysical processes. A qualitative similarity between conical stromatolites and some modern microbial mats suggests a photosynthetic origin for ancient stromatolites. To better understand and interpret ancient fossils, we seek a quantitative relationship between the geometry of conical stromatolites and the biophysical processes that control their growth. We note that all modern conical stromatolites and many that formed in the last 2.8 billion years display a characteristic centimeter-scale spacing between neighboring structures. To understand this prominent—but hitherto uninterpreted—organization, we consider the role of diffusion in mediating competition between stromatolites. Having confirmed this model through laboratory experiments and field observation, we find that organization of a field of stromatolites is set by a diffusive time scale over which individual structures compete for nutrients, thus linking form to physiology. The centimeter-scale spacing between modern and ancient stromatolites corresponds to a rhythmically fluctuating metabolism with a period of approximately 20 hr. The correspondence between the observed spacing and the day length provides quantitative support for the photosynthetic origin of conical stromatolites throughout geologic time.

Thin, filamentous, non-heterocystous, benthic cyanobacteria (Subsection III) from some marine, lacustrine and thermal environments aggregate into macroscopic cones and conical stromatolites. We investigate the uptake and storage of inorganic carbon by cone-forming cyanobacteria from Yellowstone National Park using high-resolution stable isotope mapping of labeled carbon (H13CO3−) and immunoassays. Observations and incubation experiments in actively photosynthesizing enrichment cultures and field samples reveal the presence of abundant cyanophycin granules in the active growth layer of cones. These ultrastructurally heterogeneous granules rapidly accumulate newly fixed carbon and store 18% of the total particulate labeled carbon after 120 mins of incubation. The intracellular distribution of labeled carbon during the incubation experiment demonstrates an unexpectedly large contribution of PEP carboxylase to carbon fixation, and a large flow of carbon and nitrogen toward cyanophycin in thin filamentous, non-heterocystous cyanobacteria. This pattern does not occur in obvious response to a changing N or C status. Instead, it may suggest an unusual interplay between the regulation of carbon concentration mechanisms and accumulation of photorespiratory products that facilitates uptake of inorganic C and reduces photorespiration in the dense, surface-attached communities of cyanobacteria from Subsection III., Academia Sinica (Beijing, China) (postdoctoral scholar award), National Science Council (China) (NSC 102-2116-M-006-018-MY2), National Science Council (China) (NSC 101-2116-M-001-007), Massachusetts Institute of Technology (Solomon Buchsbaum Fund)

Conical stromatolites are thought to be robust indicators of the presence of photosynthetic and phototactic microbes in aquatic environments as early as 3.5 billion years ago. However, phototaxis alone cannot explain the ubiquity of disrupted, curled, and contorted laminae in the crests of many Mesoproterozoic, Paleoproterozoic, and some Archean conical stromatolites. Here, we demonstrate that cyanobacterial production of oxygen in the tips of modern conical aggregates creates contorted laminae and submillimeter-to-millimeter-scale enmeshed bubbles. Similarly sized fossil bubbles and contorted laminae may be present only in the crestal zones of some conical stromatolites 2.7 billion years old or younger. This implies not only that cyanobacteria built Proterozoic conical stromatolites but also that fossil bubbles may constrain the timing of the evolution of oxygenic photosynthesis.

Carbonate muds deposited on continental shelves are abundant and well-preserved throughout the geologic record because shelf strata are difficult to subduct and peritidal carbonate units often form thick, rheologically strong units that resist penetrative deformation. Much of what we know about pre-Mesozoic ocean chemistry, carbon cycling, and global change is derived from isotope and trace element geochemistry of platform carbonates. Paleomagnetic data from the same sediments would be invaluable, placing records of paleolatitude, paleogeography, and perturbations to the geomagnetic field in the context and relative chronology of chemostratigraphy. To investigate the depositional and early diagenetic processes that contribute to magneitzation in carbonates, we surveyed over 500 core and surface samples of peritidal, often microbially bound carbonate muds spanning the last not, vert, similar 1000 yr and deposited on top of Pleistocene aeolianites in the Triple Goose Creek region of northwest Andros Island, Bahamas. Sedimentological, geochemical, magnetic and ferromagnetic resonance properties divide the sediment columns into three biogeochemical zones. In the upper sediments, the dominant magnetic mineral is magnetite, produced by magnetotactic bacteria and dissimiliatory microbial iron metabolism. At lower depths, above or near mean tide level, microbial iron reduction dissolves most of the magnetic particles in the sediment. In some cores, magnetic iron sulfides precipitate in a bottom zone of sulfate reduction, likely coupled to the oxidation of decaying mangrove roots. The remanent magnetization preserved in all oriented samples appears indistinguishable from the modern local geomagnetic field, which reflects the post-depositional origin of magnetic particles in the lower zone of the parasequence. While we cannot comment on the effects of late-stage diagenesis or metamorphism on remanence in carbonates, we postulate that early-cemented, thin-laminated parasequence tops in ancient peritidal carbonates are mostly likely to preserve syn-depositional paleomagnetic directions and magnetofossil stratigraphies.

The terrestrial cycling of Si is thought to have a large influence on the terrestrial and marine primary production, as well as the coupled biogeochemical cycles of Si and C. Biomineralization of silica is widespread among terrestrial eukaryotes such as plants, soil diatoms, freshwater sponges, silicifying flagellates and testate amoebae. Two major groups of testate (shelled) amoebae, arcellinids and euglyphids, produce their own silica particles to construct shells. The two are unrelated phylogenetically and acquired biomineralizing capabilities independently. Hyalosphenids, a group within arcellinids, are predators of euglyphids. We demonstrate that hyalosphenids can construct shells using silica scales mineralized by the euglyphids. Parsimony analyses of the current hyalosphenid phylogeny indicate that the ability to “steal” euglyphid scales is most likely ancestral in hyalosphenids, implying that euglyphids should be older than hyalosphenids. However, exactly when euglyphids arose is uncertain. Current fossil record contains unambiguous euglyphid fossils that are as old as 50 million years, but older fossils are scarce and difficult to interpret. Poor taxon sampling of euglyphids has also prevented the development of molecular clocks. Here, we present a novel molecular clock reconstruction for arcellinids and consider the uncertainties due to various previously used calibration points. The new molecular clock puts the origin of hyalosphenids in the early Carboniferous (∼370 mya). Notably, this estimate coincides with the widespread colonization of land by Si-accumulating plants, suggesting possible links between the evolution of Arcellinid testate amoebae and the expansion of terrestrial habitats rich in organic matter and bioavailable Si., National Aeronautics and Space Administration (Astrobiology Institute Award NNA13AA90A and NASA Exobiology award 6927184), CNPq-MIT Seed Fund (551952/2011-3)