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 (δ¹³C) 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. Significance The rise of atmospheric oxygen over Earth’s history has received much recent interdisciplinary attention. However, the puzzle of when and how atmospheric oxygen reached modern levels remains unresolved. Many recent studies have argued for a major oxygenation event—of uncertain cause—in the Neoproterozoic Era >541 Ma, enabling the rise of animals. Previous modelling work has predicted a late Paleozoic oxygen rise (<380 Ma) due to the rise of forests. Here we show that neither scenario is correct. Instead, the earliest plants, which colonized the land from 470 Ma onward, first increased atmospheric oxygen to present levels by 400 Ma, and this instigated fire-mediated feedbacks that have stabilized high oxygen levels ever since, shaping subsequent evolution.

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.

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 Tgyr⁻¹ and for phosphorus it ranges from 0.46 to 4.6 Tgyr⁻¹. Estimates of chemical weathering are between 0.058 and 1.1 km³ 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.

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 CO2 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.

Research into biotic interactions has been a core theme of ecology for over a century. However, despite the obvious role that biota play in the global carbon cycle, the effects of biotic interactions on carbon pools and fluxes are poorly understood. Here we develop a conceptual framework that illustrates the importance of biotic interactions in regulating carbon cycling based on a literature review and a quantitative synthesis by means of meta‐analysis. Our study focuses on blue carbon ecosystems—vegetated coastal ecosystems that function as the most effective long‐term CO₂sinks of the biosphere. We demonstrate that a multitude of mutualistic, competitive and consumer–resource interactions between plants, animals and microbiota exert strong effects on carbon cycling across various spatial scales ranging from the rhizosphere to the landscape scale. Climate change‐sensitive abiotic factors modulate the strength of biotic‐interaction effects on carbon fluxes, suggesting that the importance of biota‐mediated carbon cycling will change under future climatic conditions. Strong effects of biotic interactions on carbon cycling imply that biosphere‐climate feedbacks may not be sufficiently represented in current Earth system models. Inclusion of new functional groups in these models, and new approaches to simplify species interactions, may thus improve the predictions of biotic effects on the global climate.

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 process-based 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.