Abstract. Drilling for the International Continental Scientific Drilling Program (ICDP) Early Jurassic Earth System and Timescale project (JET) was undertaken between October 2020 and January 2021. The drill site is situated in a small-scale synformal basin of the latest Triassic to Early Jurassic age that formed above the major Permian–Triassic half-graben system of the Cheshire Basin. The borehole is located to recover an expanded and complete succession to complement the legacy core from the Llanbedr (Mochras Farm) borehole drilled through 1967–1969 on the edge of the Cardigan Bay Basin, North Wales. The overall aim of the project is to construct an astronomically calibrated integrated timescale for the Early Jurassic and to provide insights into the operation of the Early Jurassic Earth system. Core of Quaternary age cover and Early Jurassic mudstone was obtained from two shallow partially cored geotechnical holes (Prees 2A to 32.2 m below surface (m b.s.) and Prees 2B to 37.0 m b.s.) together with Early Jurassic and Late Triassic mudstone from the principal hole, Prees 2C, which was cored from 32.92 to 651.32 m (corrected core depth scale). Core recovery was 99.7 % for Prees 2C. The ages of the recovered stratigraphy range from the Late Triassic (probably Rhaetian) to the Early Jurassic, Early Pliensbachian (Ibex Ammonoid Chronozone). All ammonoid chronozones have been identified for the drilled Early Jurassic strata. The full lithological succession comprises the Branscombe Mudstone and Blue Anchor formations of the Mercia Mudstone Group, the Westbury and Lilstock formations of the Penarth Group, and the Redcar Mudstone Formation of the Lias Group. A distinct interval of siltstone is recognized within the Late Sinemurian of the Redcar Mudstone Formation, and the name “Prees Siltstone Member” is proposed. Depositional environments range from playa lake in the Late Triassic to distal offshore marine in the Early Jurassic. Initial datasets compiled from the core include radiography, natural gamma ray, density, magnetic susceptibility, and X-ray fluorescence (XRF). A full suite of downhole logs was also run. Intervals of organic carbon enrichment occur in the Rhaetian (Late Triassic) Westbury Formation and in the earliest Hettangian and earliest Pliensbachian strata of the Redcar Mudstone Formation, where up to 4 % total organic carbon (TOC) is recorded. Other parts of the succession are generally organic-lean, containing less than 1 % TOC. Carbon-isotope values from bulk organic matter have also been determined, initially at a resolution of ∼ 1 m, and these provide the basis for detailed correlation between the Prees 2 succession and adjacent boreholes and Global Stratotype Section and Point (GSSP) outcrops. Multiple complementary studies are currently underway and preliminary results promise an astronomically calibrated biostratigraphy, magnetostratigraphy, and chemostratigraphy for the combined Prees and Mochras successions as well as insights into the dynamics of background processes and major palaeo-environmental changes.
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The endosymbiotic origin of mitochondria during eukaryogenesis has long been viewed as an adaptive response to the oxygenation of Earth's surface environment, presuming a fundamentally aerobic lifestyle for the free-living bacterial ancestors of mitochondria. This oxygen-centric view has been robustly challenged by recent advances in the Earth and life sciences. While the permanent oxygenation of the atmosphere above trace concentrations is now thought to have occurred 2.2 billion years ago, large parts of the deep ocean remained anoxic until less than 0.5 billion years ago. Neither fossils nor molecular clocks correlate the origin of mitochondria, or eukaryogenesis more broadly, to either of these planetary redox transitions. Instead, mitochondria-bearing eukaryotes are consistently dated to between these two oxygenation events, during an interval of pervasive deep-sea anoxia and variable surface-water oxygenation. The discovery and cultivation of the Asgard archaea has reinforced metabolic evidence that eukaryogenesis was initially mediated by syntrophic H2 exchange between an archaeal host and an α-proteobacterial symbiont living under anoxia. Together, these results temporally, spatially and metabolically decouple the earliest stages of eukaryogenesis from the oxygen content of the surface ocean and atmosphere. Rather than reflecting the ancestral metabolic state, obligate aerobiosis in eukaryotes is most probably derived, having only become globally widespread over the past 1 billion years as atmospheric oxygen approached modern levels.

The Ediacaran Shuram negative carbon isotope excursion (SE) records major paleoceanographic changes during the late Neoproterozoic, possibly linked to a global oceanic oxygenation event, yet its cause(s) remain uncertain. Earlier studies of the upper Ediacaran Doushantuo Formation in South China based on local redox proxies have documented strong spatial redox heterogeneity along shelf-to-basin transects, but variations of δ238U (a global redox proxy) have not yet been examined in deep-water SE carbonates. In this study, we examined δ238U variations through the SE in the upper slope Siduping section. Similar to platform SE sections, Siduping exhibits a shift toward higher δ238U values correlative with the peak of the SE (i.e. maximum negative δ13Ccarb), confirming inferences of global ocean oxygenation during the SE. This raises an apparent paradox, because a global negative carbon isotope excursion implies net oxidant consumption, requiring an ocean-based oxygenation mechanism. We hypothesize that an increase in the efficiency of phosphorus burial due to a plankton-driven shift from dominantly dissolved organic matter (DOM) cycling to greater particulate organic matter (POM) export depleted the ocean of nutrient phosphorus. By producing a steep redox gradient close to the sediment-water interface, we suggest that ocean oxygenation also triggered a globally simultaneous diagenetic event in which isotopically light δ13Ccarb was precipitated in authigenic carbonate minerals. This scenario can account for δ238U differences between shallow-water and deep-water carbonates, which reflect precipitation of relatively larger amounts of authigenic carbonate minerals in shallow-water settings, generating both a larger negative δ13Ccarb shift and a larger early diagenetic δ238U offset.

The Neoproterozoic era witnessed a succession of biological innovations that culminated in diverse animal body plans and behaviours during the Ediacaran–Cambrian radiations. Intriguingly, this interval is also marked by perturbations to the global carbon cycle, as evidenced by extreme fluctuations in climate and carbon isotopes. The Neoproterozoic isotope record has defied parsimonious explanation because sustained 12C-enrichment (low δ13C) in seawater seems to imply that substantially more oxygen was consumed by organic carbon oxidation than could possibly have been available. We propose a solution to this problem, in which carbon and oxygen cycles can maintain dynamic equilibrium during negative δ13C excursions when surplus oxidant is generated through bacterial reduction of sulfate that originates from evaporite weathering. Coupling of evaporite dissolution with pyrite burial drives a positive feedback loop whereby net oxidation of marine organic carbon can sustain greenhouse forcing of chemical weathering, nutrient input and ocean margin euxinia. Our proposed framework is particularly applicable to the late Ediacaran ‘Shuram’ isotope excursion that directly preceded the emergence of energetic metazoan metabolisms during the Ediacaran–Cambrian transition. Here we show that non-steady-state sulfate dynamics contributed to climate change, episodic ocean oxygenation and opportunistic radiations of aerobic life during the Neoproterozoic era.

We present the first version of OMEN-SED (Organic Matter ENabled SEDiment model), a new, onedimensional analytical early diagenetic model resolving organic matter cycling and the associated biogeochemical dynamics in marine sediments designed to be coupled to Earth system models. OMEN-SED explicitly describes organic matter (OM) cycling and the associated dynamics of the most important terminal electron acceptors (i.e. O2, NO3, SO4) and methane (CH4), related reduced substances (NH4, H2S), macronutrients (PO4) and associated pore water quantities (ALK, DIC). Its reaction network accounts for the most important primary and secondary redox reactions, equilibrium reactions, mineral dissolution and precipitation, as well as adsorption and desorption processes associated with OM dynamics that affect the dissolved and solid species explicitly resolved in the model. To represent a redox-dependent sedimentary P cycle we also include a representation of the formation and burial of Fe-bound P and authigenic Ca-P minerals. Thus, OMEN-SED is able to capture the main features of diagenetic dynamics in marine sediments and therefore offers similar predictive abilities as a complex, numerical diagenetic model. Yet, its computational efficiency allows for its coupling to global Earth system models and therefore the investigation of coupled global biogeochemical dynamics over a wide range of climate-relevant timescales. This paper provides a detailed description of the new sediment model, an extensive sensitivity analysis and an evaluation of OMEN-SED's performance through comprehensive comparisons with observations and results from a more complex numerical model. We find that solid-phase and dissolved pore water profiles for different ocean depths are reproduced with good accuracy and simulated terminal electron acceptor fluxes fall well within the range of globally observed fluxes. Finally, we illustrate its application in an Earth system model framework by coupling OMEN-SED to the Earth system model cGENIE and tune the OM degradation rate constants to optimise the fit of simulated benthic OM contents to global observations. We find that the simulated sediment characteristics of the coupled model framework, such as OM degradation rates, oxygen penetration depths and sediment-water interface fluxes, are generally in good agreement with observations and in line with what one would expect on a global scale. Coupled to an Earth system model, OMENSED is thus a powerful tool that will not only help elucidate the role of benthic-pelagic exchange processes in the evolution and the termination of a wide range of climate events, but will also allow for a direct comparison of model output with the sedimentary record - the most important climate archive on Earth.

Recently postulated mechanisms and models can help explain the enduring ‘Gaia’ puzzle of environmental regulation mediated by life. Natural selection can produce nutrient recycling at local scales and regulation of heterogeneous environmental variables at ecosystem scales. However, global-scale environmental regulation involves a temporal and spatial decoupling of effects from actors that makes conventional evolutionary explanations problematic. Instead, global regulation can emerge by a process of ‘sequential selection’ in which systems that destabilize their environment are short-lived and result in extinctions and reorganizations until a stable attractor is found. Such persistence-enhancing properties can in turn increase the likelihood of acquiring further persistence-enhancing properties through ‘selection by survival alone’. Thus, Earth system feedbacks provide a filter for persistent combinations of macroevolutionary innovations.

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 pO 2 â 1/40.1 PAL (present atmospheric level), but that stability is lost at pO 2

The ocean has undergone several profound biogeochemical transformations in its 4-billion-year history, and these were an integral part of the coevolution of life and the planet. This review focuses on changes in ocean redox state as controlled by changes in biological activity, nutrient concentrations, and atmospheric O2. Motivated by disparate interpretations of available geochemical data, we aim to show how quantitative modeling-spanning microbial mats, shelf seas, and the open ocean-can help constrain past ocean biogeochemical redox states and show what caused transformations between them. We outline key controls on ocean redox structure and review pertinent proxies and their interpretation. We then apply this quantitative framework to three key questions: How did the origin of oxygenic photosynthesis transform ocean biogeochemistry? How did the Great Oxidation transform ocean biogeochemistry? and how was ocean biogeochemistry transformed in the Neoproterozoic-Paleozoic?

Contents 531 I. 531 II. 532 III. 534 IV. 535 V. 535 VI. 535 Acknowledgements 536 References 536 SUMMARY: There is growing evidence that life has been on land for billions of years. Microbial mats fuelled by oxygenic photosynthesis were probably present in terrestrial habitats from c. 3.0 billion yr ago (Ga) onwards, creating localized 'oxygen oases' under a reducing atmosphere, which left a characteristic oxidative weathering signal. After the Great Oxidation c. 2.4 Ga, the now oxidizing atmosphere masked that redox signal, but ancient soils record the mobilization of phosphorus and other elements by organic acids in weathering profiles. Evidence for Neoproterozoic 'greening of the land' and intensification of weathering c. 0.85-0.54 Ga is currently equivocal. However, the mid-Palaeozoic c. 0.45-0.4 Ga shows global atmospheric changes consistent with increased terrestrial productivity and intensified weathering by the first land plants.

The advent of genomic-, transcriptomic- and proteomic-based approaches has revolutionized our ability to describe marine microbial communities, including biogeography, metabolic potential and diversity, mechanisms of adaptation, and phylogeny and evolutionary history. New interdisciplinary approaches are needed to move from this descriptive level to improved quantitative, process-level understanding of the roles of marine microbes in biogeochemical cycles and of the impact of environmental change on the marine microbial ecosystem. Linking studies at levels from the genome to the organism, to ecological strategies and organism and ecosystem response, requires new modelling approaches. Key to this will be a fundamental shift in modelling scale that represents micro-organisms from the level of their macromolecular components. This will enable contact with omics data sets and allow acclimation and adaptive response at the phenotype level (i.e. traits) to be simulated as a combination of fitness maximization and evolutionary constraints. This way forward will build on ecological approaches that identify key organism traits and systems biology approaches that integrate traditional physiological measurements with new insights from omics. It will rely on developing an improved understanding of ecophysiology to understand quantitatively environmental controls on microbial growth strategies. It will also incorporate results from experimental evolution studies in the representation of adaptation. The resulting ecosystem-level models can then evaluate our level of understanding of controls on ecosystem structure and function, highlight major gaps in understanding and help prioritize areas for future research programs. Ultimately, this grand synthesis should improve predictive capability of the ecosystem response to multiple environmental drivers.

The progressive oxygenation of the Earth's atmosphere was pivotal to the evolution of life, but the puzzle of when and how atmospheric oxygen (O2) first approached modern levels (∼21%) remains unresolved. Redox proxy data indicate the deep oceans were oxygenated during 435-392 Ma, and the appearance of fossil charcoal indicates O2 >15-17% by 420-400 Ma. However, existing models have failed to predict oxygenation at this time. Here we show that the earliest plants, which colonized the land surface from ∼470 Ma onward, were responsible for this mid-Paleozoic oxygenation event, through greatly increasing global organic carbon burial-the net long-term source of O2 We use a trait-based ecophysiological model to predict that cryptogamic vegetation cover could have achieved ∼30% of today's global terrestrial net primary productivity by ∼445 Ma. Data from modern bryophytes suggests this plentiful early plant material had a much higher molar C:P ratio (∼2,000) than marine biomass (∼100), such that a given weathering flux of phosphorus could support more organic carbon burial. Furthermore, recent experiments suggest that early plants selectively increased the flux of phosphorus (relative to alkalinity) weathered from rocks. Combining these effects in a model of long-term biogeochemical cycling, we reproduce a sustained +2‰ increase in the carbonate carbon isotope (δ(13)C) record by ∼445 Ma, and predict a corresponding rise in O2 to present levels by 420-400 Ma, consistent with geochemical data. This oxygen rise represents a permanent shift in regulatory regime to one where fire-mediated negative feedbacks stabilize high O2 levels.

The balance of evidence suggests that oxygenic photosynthesis had evolved by 3.0-2.7 Ga, several hundred million years prior to the Great Oxidation ≈2.4 Ga. Previous work has shown that if oxygenic photosynthesis spread globally prior to the Great Oxidation, this could have supported widespread aerobic ecosystems in the surface ocean, without oxidising the atmosphere. Here we use a suite of models to explore the implications for carbon cycling and the Great Oxidation. We find that recycling of oxygen and carbon within early aerobic marine ecosystems would have restricted the balanced fluxes of methane and oxygen escaping from the ocean, lowering the atmospheric concentration of methane in the Great Oxidation transition and its aftermath. This in turn would have minimised any bi-stability of atmospheric oxygen, by weakening a stabilising feedback on oxygen from hydrogen escape to space. The result would have been a more reversible and probably episodic rise of oxygen at the Great Oxidation transition, consistent with existing geochemical evidence. The resulting drop in methane levels to ≈10 ppm is consistent with climate cooling at the time but adds to the puzzle of what kept the rest of the Proterozoic warm. A key test of the scenario of abundant methanotrophy in oxygen oases before the Great Oxidation is its predicted effects on the organic carbon isotope (δ13Corg) record. Our open ocean general circulation model predicts δCorg13≈-30 to -45‰ consistent with most data from 2.65 to 2.45 Ga. However, values of δCorg13≈-50‰ require an extreme scenario such as concentrated methanotroph production where shelf-slope upwelling of methane-rich water met oxic shelf water.

Ocean acidification triggered by Siberian Trap volcanism was a possible kill mechanism for the Permo-Triassic Boundary mass extinction, but direct evidence for an acidification event is lacking. We present a high-resolution seawater pH record across this interval, using boron isotope data combined with a quantitative modeling approach. In the latest Permian, increased ocean alkalinity primed the Earth system with a low level of atmospheric CO2 and a high ocean buffering capacity. The first phase of extinction was coincident with a slow injection of carbon into the atmosphere, and ocean pH remained stable. During the second extinction pulse, however, a rapid and large injection of carbon caused an abrupt acidification event that drove the preferential loss of heavily calcified marine biota.

Tectonic drivers of degassing and weathering processes are key long-term controls on atmospheric CO2. However, there is considerable debate over the changing relative importance of different carbon sources and sinks. Existing geochemical models have tended to rely on indirect methods to derive tectonic drivers, such as inversion of the seawater 87Sr/86Sr curve to estimate uplift or continental basalt area. Here we use improving geologic data to update the representation of tectonic drivers in the COPSE biogeochemical model. The resulting model distinguishes CO2 sinks from terrestrial granite weathering, total basalt weathering, and seafloor alteration. It also distinguishes CO2 sources from subduction zone metamorphism and from igneous intrusions. We reconstruct terrestrial basaltic area from data on the extent of large igneous provinces and use their volume to estimate their contribution to degassing. We adopt a recently published reconstruction of subduction-related degassing, and relate seafloor weathering to ocean crust creation rate. Revised degassing alone tends to produce unrealistically high CO2, but this is counteracted by the inclusion of seafloor alteration and global basalt weathering, producing a good overall fit to Mesozoic-Cenozoic proxy CO2 estimates and a good fit to 87Sr/86Sr data. The model predicts that seafloor alteration and terrestrial weathering made similar contributions to CO2 removal through the Triassic and Jurassic, after which terrestrial weathering increased and seafloor weathering declined. We predict that basalts made a greater contribution to silicate weathering than granites through the Mesozoic, before the contribution of basalt weathering declined over the Cenozoic due to decreasing global basaltic area.

The controls on the 'Redfield' N : P stoichiometry of marine phytoplankton and hence the N : P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource-allocation trade-offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per-ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N : P ratios and N : P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.

Here we describe a new trait-based model for cellular resource allocation that we use to investigate the relative importance of different drivers for small cell size in phytoplankton. Using the model, we show that increased investment in nonscalable structural components with decreasing cell size leads to a trade-off between cell size, nutrient and light affinity, and growth rate. Within the most extreme nutrient-limited, stratified environments, resource competition theory then predicts a trend toward larger minimum cell size with increasing depth. We demonstrate that this explains observed trends using a marine ecosystem model that represents selection and adaptation of a diverse community defined by traits for cell size and subcellular resource allocation. This framework for linking cellular physiology to environmental selection can be used to investigate the adaptive response of the marine microbial community to environmental conditions and the adaptive value of variations in cellular physiology. © 2013, by the Association for the Sciences of Limnology and Oceanography, Inc.