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Concept: Proterozoic


Chemical disequilibrium in planetary atmospheres has been proposed as a generalized method for detecting life on exoplanets through remote spectroscopy. Among solar system planets with substantial atmospheres, the modern Earth has the largest thermodynamic chemical disequilibrium due to the presence of life. However, how this disequilibrium changed over time and, in particular, the biogenic disequilibria maintained in the anoxic Archean or less oxic Proterozoic eons are unknown. We calculate the atmosphere-ocean disequilibrium in the Precambrian using conservative proxy- and model-based estimates of early atmospheric and oceanic compositions. We omit crustal solids because subsurface composition is not detectable on exoplanets, unlike above-surface volatiles. We find that (i) disequilibrium increased through time in step with the rise of oxygen; (ii) both the Proterozoic and Phanerozoic may have had remotely detectable biogenic disequilibria due to the coexistence of O2, N2, and liquid water; and (iii) the Archean had a biogenic disequilibrium caused by the coexistence of N2, CH4, CO2, and liquid water, which, for an exoplanet twin, may be remotely detectable. On the basis of this disequilibrium, we argue that the simultaneous detection of abundant CH4 and CO2 in a habitable exoplanet’s atmosphere is a potential biosignature. Specifically, we show that methane mixing ratios greater than 10-3 are potentially biogenic, whereas those exceeding 10-2 are likely biogenic due to the difficulty in maintaining large abiotic methane fluxes to support high methane levels in anoxic atmospheres. Biogenicity would be strengthened by the absence of abundant CO, which should not coexist in a biological scenario.

Concepts: Water, Solar System, Planet, Mars, Atmosphere, Neptune, Uranus, Proterozoic


It is widely accepted that Earth’s early atmosphere contained less than 0.001 per cent of the present-day atmospheric oxygen (O2) level, until the Great Oxidation Event resulted in a major rise in O2 concentration about 2.4 billion years ago. There are multiple lines of evidence for low O2 concentrations on early Earth, but all previous observations relate to the composition of the lower atmosphere in the Archaean era; to date no method has been developed to sample the Archaean upper atmosphere. We have extracted fossil micrometeorites from limestone sedimentary rock that had accumulated slowly 2.7 billion years ago before being preserved in Australia’s Pilbara region. We propose that these micrometeorites formed when sand-sized particles entered Earth’s atmosphere and melted at altitudes of about 75 to 90 kilometres (given an atmospheric density similar to that of today). Here we show that the FeNi metal in the resulting cosmic spherules was oxidized while molten, and quench-crystallized to form spheres of interlocking dendritic crystals primarily of magnetite (Fe3O4), with wüstite (FeO)+metal preserved in a few particles. Our model of atmospheric micrometeorite oxidation suggests that Archaean upper-atmosphere oxygen concentrations may have been close to those of the present-day Earth, and that the ratio of oxygen to carbon monoxide was sufficiently high to prevent noticeable inhibition of oxidation by carbon monoxide. The anomalous sulfur isotope (Δ(33)S) signature of pyrite (FeS2) in seafloor sediments from this period, which requires an anoxic surface environment, implies that there may have been minimal mixing between the upper and lower atmosphere during the Archaean.

Concepts: Cyanobacteria, Photosynthesis, Oxygen, Iron, Sedimentary rock, Carbon, Atmosphere, Proterozoic


Ediacaran (635-542 million years ago) fossils have been regarded as early animal ancestors of the Cambrian evolutionary explosion of marine invertebrate phyla, as giant marine protists and as lichenized fungi. Recent documentation of palaeosols in the Ediacara Member of the Rawnsley Quartzite of South Australia confirms past interpretations of lagoonal-aeolian deposition based on synsedimentary ferruginization and loessic texture. Further evidence for palaeosols comes from non-marine facies, dilation cracks, soil nodules, sand crystals, stable isotopic data and mass balance geochemistry. Here I show that the uppermost surfaces of the palaeosols have a variety of fossils in growth position, including Charniodiscus, Dickinsonia, Hallidaya, Parvancorina, Phyllozoon, Praecambridium, Rugoconites, Tribrachidium and ‘old-elephant skin’ (ichnogenus Rivularites). These fossils were preserved as ferruginous impressions, like plant fossils, and biological soil crusts of Phanerozoic eon sandy palaeosols. Sand crystals after gypsum and nodules of carbonate are shallow within the palaeosols, even after correcting for burial compaction. Periglacial involutions and modest geochemical differentiation of the palaeosols are evidence of a dry, cold temperate Ediacaran palaeoclimate in South Australia. This new interpretation of some Ediacaran fossils as large sessile organisms of cool, dry soils, is compatible with observations that Ediacaran fossils were similar in appearance and preservation to lichens and other microbial colonies of biological soil crusts, rather than marine animals, or protists.

Concepts: Animal, Geology, Invertebrate, Paleontology, Cambrian explosion, Proterozoic, Ediacaran, Dickinsonia


The transition from dominant bacterial to eukaryotic marine primary productivity was one of the most profound ecological revolutions in the Earth’s history, reorganizing the distribution of carbon and nutrients in the water column and increasing energy flow to higher trophic levels. But the causes and geological timing of this transition, as well as possible links with rising atmospheric oxygen levels and the evolution of animals, remain obscure. Here we present a molecular fossil record of eukaryotic steroids demonstrating that bacteria were the only notable primary producers in the oceans before the Cryogenian period (720-635 million years ago). Increasing steroid diversity and abundance marks the rapid rise of marine planktonic algae (Archaeplastida) in the narrow time interval between the Sturtian and Marinoan ‘snowball Earth’ glaciations, 659-645 million years ago. We propose that the incumbency of cyanobacteria was broken by a surge of nutrients supplied by the Sturtian deglaciation. The ‘Rise of Algae’ created food webs with more efficient nutrient and energy transfers, driving ecosystems towards larger and increasingly complex organisms. This effect is recorded by the concomitant appearance of biomarkers for sponges and predatory rhizarians, and the subsequent radiation of eumetazoans in the Ediacaran period.

Concepts: Photosynthesis, Oxygen, Evolution, Plant, Ecology, Proterozoic, Snowball Earth, Neoproterozoic


The first significant buildup in atmospheric oxygen, the Great Oxidation Event (GOE), began in the early Paleoproterozoic in association with global glaciations and continued until the end of the Lomagundi carbon isotope excursion ca. 2,060 Ma. The exact timing of and relationships among these events are debated because of poor age constraints and contradictory stratigraphic correlations. Here, we show that the first Paleoproterozoic global glaciation and the onset of the GOE occurred between ca. 2,460 and 2,426 Ma, ∼100 My earlier than previously estimated, based on an age of 2,426 ± 3 Ma for Ongeluk Formation magmatism from the Kaapvaal Craton of southern Africa. This age helps define a key paleomagnetic pole that positions the Kaapvaal Craton at equatorial latitudes of 11° ± 6° at this time. Furthermore, the rise of atmospheric oxygen was not monotonic, but was instead characterized by oscillations, which together with climatic instabilities may have continued over the next ∼200 My until ≤2,250-2,240 Ma. Ongeluk Formation volcanism at ca. 2,426 Ma was part of a large igneous province (LIP) and represents a waning stage in the emplacement of several temporally discrete LIPs across a large low-latitude continental landmass. These LIPs played critical, albeit complex, roles in the rise of oxygen and in both initiating and terminating global glaciations. This series of events invites comparison with the Neoproterozoic oxygen increase and Sturtian Snowball Earth glaciation, which accompanied emplacement of LIPs across supercontinent Rodinia, also positioned at low latitude.

Concepts: Cyanobacteria, Oxygen, Earth, Proterozoic, Snowball Earth, Rodinia, Great Oxygenation Event, Geological history of oxygen


Protection against arsenic damage in organisms positioned deep in the tree of life points to early evolutionary sensitization. Here, marine sedimentary records reveal a Proterozoic arsenic concentration patterned to glacial-interglacial ages. The low glacial and high interglacial sedimentary arsenic concentrations, suggest deteriorating habitable marine conditions may have coincided with atmospheric oxygen decline after ~2.1 billion years ago. A similar intensification of near continental margin sedimentary arsenic levels after the Cryogenian glaciations is also associated with amplified continental weathering. However, interpreted atmospheric oxygen increase at this time, suggests that the marine biosphere had widely adapted to the reorganization of global marine elemental cycles by glaciations. Such a glacially induced biogeochemical bridge would have produced physiologically robust communities that enabled increased oxygenation of the ocean-atmosphere system and the radiation of the complex Ediacaran-Cambrian life.

Concepts: Cyanobacteria, Photosynthesis, Life, Water, Earth, Oceanography, Ice age, Proterozoic


Cyanobacteria are among the most diverse prokaryotic phyla, with morphotypes ranging from unicellular to multicellular filamentous forms, including those able to terminally (i.e., irreversibly) differentiate in form and function. It has been suggested that cyanobacteria raised oxygen levels in the atmosphere around 2.45-2.32 billion y ago during the Great Oxidation Event (GOE), hence dramatically changing life on the planet. However, little is known about the temporal evolution of cyanobacterial lineages, and possible interplay between the origin of multicellularity, diversification of cyanobacteria, and the rise of atmospheric oxygen. We estimated divergence times of extant cyanobacterial lineages under Bayesian relaxed clocks for a dataset of 16S rRNA sequences representing the entire known diversity of this phylum. We tested whether the evolution of multicellularity overlaps with the GOE, and whether multicellularity is associated with significant shifts in diversification rates in cyanobacteria. Our results indicate an origin of cyanobacteria before the rise of atmospheric oxygen. The evolution of multicellular forms coincides with the onset of the GOE and an increase in diversification rates. These results suggest that multicellularity could have played a key role in triggering cyanobacterial evolution around the GOE.

Concepts: Cyanobacteria, Photosynthesis, Oxygen, Archaea, Bacteria, Proterozoic, Great Oxygenation Event, Geological history of oxygen


The oxygenation of Earth’s surface fundamentally altered global biogeochemical cycles and ultimately paved the way for the rise of metazoans at the end of the Proterozoic. However, current estimates for atmospheric oxygen (O2) levels during the billion years leading up to this time vary widely. On the basis of chromium (Cr) isotope data from a suite of Proterozoic sediments from China, Australia, and North America, interpreted in the context of data from similar depositional environments from Phanerozoic time, we find evidence for inhibited oxidation of Cr at Earth’s surface in the mid-Proterozoic (1.8 to 0.8 billion years ago). These data suggest that atmospheric O2 levels were at most 0.1% of present atmospheric levels. Direct evidence for such low O2 concentrations in the Proterozoic helps explain the late emergence and diversification of metazoans.

Concepts: Photosynthesis, Oxygen, Carbon dioxide, Iron, Earth, Aluminium, Geology, Proterozoic


Marine planktonic cyanobacteria contributed to the widespread oxygenation of the oceans towards the end of the Pre-Cambrian and their evolutionary origin represents a key transition in the geochemical evolution of the Earth surface. Little is known, however, about the evolutionary events that led to the appearance of marine planktonic cyanobacteria. I present here phylogenomic (135 proteins and two ribosomal RNAs), Bayesian relaxed molecular clock (18 proteins, SSU and LSU) and Bayesian stochastic character mapping analyses from 131 cyanobacteria genomes with the aim to unravel key evolutionary steps involved in the origin of marine planktonic cyanobacteria. While filamentous cell types evolved early on at around 2,600-2,300 Mya and likely dominated microbial mats in benthic environments for most of the Proterozoic (2,500-542 Mya), marine planktonic cyanobacteria evolved towards the end of the Proterozoic and early Phanerozoic. Crown groups of modern terrestrial and/or benthic coastal cyanobacteria appeared during the late Paleoproterozoic to early Mesoproterozoic. Decrease in cell diameter and loss of filamentous forms contributed to the evolution of unicellular planktonic lineages during the middle of the Mesoproterozoic (1,600-1,000 Mya) in freshwater environments. This study shows that marine planktonic cyanobacteria evolved from benthic marine and some diverged from freshwater ancestors during the Neoproterozoic (1,000-542 Mya).

Concepts: Cyanobacteria, Photosynthesis, Gene, Archaea, Bacteria, Evolution, Earth, Proterozoic


It is unclear why atmospheric oxygen remained trapped at low levels for more than 1.5 billion years following the Paleoproterozoic Great Oxidation Event. Here, we use models for erosion, weathering and biogeochemical cycling to show that this can be explained by the tectonic recycling of previously accumulated sedimentary organic carbon, combined with the oxygen sensitivity of oxidative weathering. Our results indicate a strong negative feedback regime when atmospheric oxygen concentration is of order pO2∼0.1 PAL (present atmospheric level), but that stability is lost at pO2<0.01 PAL. Within these limits, the carbonate carbon isotope (δ(13)C) record becomes insensitive to changes in organic carbon burial rate, due to counterbalancing changes in the weathering of isotopically light organic carbon. This can explain the lack of secular trend in the Precambrian δ(13)C record, and reopens the possibility that increased biological productivity and resultant organic carbon burial drove the Great Oxidation Event.

Concepts: Cyanobacteria, Photosynthesis, Oxygen, Carbon dioxide, Carbon, Proterozoic, Great Oxygenation Event, Geological history of oxygen