The progressive oxygenation of the Earth's atmosphere was pivotal to the evolution of life, but the puzzle of when and how atmospheric oxygen (O 2 ) 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 O 2 >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 burialthe net long-term source of O 2 . 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 O 2 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 O 2 levels.oxygen | Paleozoic | phosphorus | plants | weathering

Abstract. Lichens and bryophytes are abundant globally and they may even form the dominant autotrophs in (sub)polar ecosystems, in deserts and at high altitudes. Moreover, they can be found in large amounts as epiphytes in old-growth forests. Here, we present the first process-based model which estimates the net carbon uptake by these organisms at the global scale, thus assessing their significance for biogeochemical cycles. The model uses gridded climate data and key properties of the habitat (e.g. disturbance intervals) to predict processes which control net carbon uptake, namely photosynthesis, respiration, water uptake and evaporation. It relies on equations used in many dynamical vegetation models, which are combined with concepts specific to lichens and bryophytes, such as poikilohydry or the effect of water content on CO 2 diffusivity. To incorporate the great functional variation of lichens and bryophytes at the global scale, the model parameters are characterised by broad ranges of possible values instead of a single, globally uniform value. The predicted terrestrial net uptake of 0.34 to 3.3 Gt yr −1 of carbon and global patterns of productivity are in accordance with empirically-derived estimates. Considering that the assimilated carbon can be invested in processes such as weathering or nitrogen fixation, lichens and bryophytes may play a significant role in biogeochemical cycles.

Abstract. Bryophyte and lichen cover on the forest floor at high latitudes exerts an insulating effect on the ground. In this way, the cover decreases mean annual soil temperature and can protect permafrost soil. Climate change, however, may change bryophyte and lichen cover, with effects on the permafrost state and related carbon balance. It is, therefore, crucial to predict how the bryophyte and lichen cover will react to environmental change at the global scale. To date, current global land surface models contain only empirical representations of the bryophyte and lichen cover, which makes it impractical to predict the future state and function of bryophytes and lichens. For this reason, we integrate a process-based model of bryophyte and lichen growth into the global land surface model JSBACH (Jena Scheme for Biosphere-Atmosphere Coupling in Hamburg). The model simulates bryophyte and lichen cover on upland sites. Wetlands are not included. We take into account the dynamic nature of the thermal properties of the bryophyte and lichen cover and their relation to environmental factors. Subsequently, we compare simulations with and without bryophyte and lichen cover to quantify the insulating effect of the organisms on the soil.We find an average cooling effect of the bryophyte and lichen cover of 2.7 K on temperature in the topsoil for the region north of 50 • N under the current climate. Locally, a cooling of up to 5.7 K may be reached. Moreover, we show that using a simple, empirical representation of the bryophyte and lichen cover without dynamic properties only results in an average cooling of around 0.5 K. This suggests that (a) bryophytes and lichens have a significant impact on soil temperature in high-latitude ecosystems and (b) a processbased description of their thermal properties is necessary for a realistic representation of the cooling effect. The advanced land surface scheme, including a dynamic bryophyte and lichen model, will be the basis for an improved future projection of land-atmosphere heat and carbon exchange.

Lichens and bryophytes may significantly affect global biogeochemical cycles by fixation of nitrogen and biotic enhancement of surface weathering rates. Most of the studies suggesting these effects, however, are either conceptual or rely on upscaling of regional estimates to obtain global numbers. Here we use a different method, based on estimates of net carbon uptake, to quantify the impacts of lichens and bryophytes on biogeochemical cycles at the global scale. We focus on three processes, namely, nitrogen fixation, phosphorus uptake, and chemical weathering. Our estimates have the form of potential rates, which means that we quantify the amount of nitrogen and phosphorus needed by the organisms to build up biomass, also accounting for resorption and leaching of nutrients. Subsequently, we use potential phosphorus uptake on bare ground to estimate chemical weathering by the organisms, assuming that they release weathering agents to obtain phosphorus. The predicted requirement for nitrogen ranges from 3.5 to 34 Tg yr −1 and for phosphorus it ranges from 0.46 to 4.6 Tg yr −1 . Estimates of chemical weathering are between 0.058 and 1.1 km 3 yr −1 of rock. These values seem to have a realistic order of magnitude, and they support the notion that lichens and bryophytes have the potential to play an important role for biogeochemical cycles.

It has been hypothesized that predecessors of today's bryophytes significantly increased global chemical weathering in the Late Ordovician, thus reducing atmospheric CO2 concentration and contributing to climate cooling and an interval of glaciations. Studies that try to quantify the enhancement of weathering by non-vascular vegetation, however, are usually limited to small areas and low numbers of species, which hampers extrapolating to the global scale and to past climatic conditions. Here we present a spatially explicit modelling approach to simulate global weathering by non-vascular vegetation in the Late Ordovician. We estimate a potential global weathering flux of 2.8 (km3 rock) yr−1, defined here as volume of primary minerals affected by chemical transformation. This is around three times larger than today's global chemical weathering flux. Moreover, we find that simulated weathering is highly sensitive to atmospheric CO2 concentration. This implies a strong negative feedback between weathering by non-vascular vegetation and Ordovician climate.

Abstract. The cycling of carbon (C) between the Earth surface and the atmosphere is controlled by biological and abiotic processes that regulate C storage in biogeochemical compartments and release to the atmosphere. This partitioning is quantified using various forms of C-use efficiency (CUE) – the ratio of C remaining in a system to C entering that system. Biological CUE is the fraction of C taken up allocated to biosynthesis. In soils and sediments, C storage depends also on abiotic processes, so the term C-storage efficiency (CSE) can be used. Here we first review and reconcile CUE and CSE definitions proposed for autotrophic and heterotrophic organisms and communities, food webs, whole ecosystems and watersheds, and soils and sediments using a common mathematical framework. Second, we identify general CUE patterns; for example, the actual CUE increases with improving growth conditions, and apparent CUE decreases with increasing turnover. We then synthesize > 5000 CUE estimates showing that CUE decreases with increasing biological and ecological organization – from unicellular to multicellular organisms and from individuals to ecosystems. We conclude that CUE is an emergent property of coupled biological–abiotic systems, and it should be regarded as a flexible and scale-dependent index of the capacity of a given system to effectively retain C.

Abstract. It is important that climate models can accurately simulate the terrestrial carbon cycle in the Arctic due to the large and potentially labile carbon stocks found in permafrost-affected environments, which can lead to a positive climate feedback, along with the possibility of future carbon sinks from northward expansion of vegetation under climate warming. Here we evaluate the simulation of tundra carbon stocks and fluxes in three land surface schemes that each form part of major Earth system models (JSBACH, Germany; JULES, UK; ORCHIDEE, France). We use a site-level approach in which comprehensive, high-frequency datasets allow us to disentangle the importance of different processes. The models have improved physical permafrost processes and there is a reasonable correspondence between the simulated and measured physical variables, including soil temperature, soil moisture and snow.We show that if the models simulate the correct leaf area index (LAI), the standard C3 photosynthesis schemes produce the correct order of magnitude of carbon fluxes. Therefore, simulating the correct LAI is one of the first priorities. LAI depends quite strongly on climatic variables alone, as we see by the fact that the dynamic vegetation model can simulate most of the differences in LAI between sites, based almost entirely on climate inputs. However, we also identify an influence from nutrient limitation as the LAI becomes too large at some of the more nutrient-limited sites. We conclude that including moss as well as vascular plants is of primary importance to the carbon budget, as moss contributes a large fraction to the seasonal CO 2 flux in nutrient-limited conditions. Moss photosynthetic activity can be strongly influenced by the moisture content of moss, and the carbon uptake can be significantly different from vascular plants with a similar LAI.The soil carbon stocks depend strongly on the rate of input of carbon from the vegetation to the soil, and our analysis suggests that an improved simulation of photosynthesis would also lead to an improved simulation of soil carbon stocks. However, the stocks are also influenced by soil carbon burial (e.g. through cryoturbation) and the rate of heterotrophic respiration, which depends on the soil physical state. More detailed below-ground measurements are needed to fully evaluate biological and physical soil processes. Furthermore, even if these processes are well modelled, the soil carbon profiles cannot resemble peat layers as peat accumulation processes are not represented in the models.Thus, we identify three priority areas for model development: (1) dynamic vegetation including (a) climate and (b) nutrient limitation effects; (2) adding moss as a plant functional type; and an (3) improved vertical profile of soil carbon including peat processes.

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