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Continental configuration controls ocean oxygenation during the Phanerozoic


The early evolutionary and much of the extinction history of marine animals is thought to be driven by changes in dissolved oxygen concentrations ([O2]) in the ocean1,2,3. In turn, [O2] is widely assumed to be dominated by the geological history of atmospheric oxygen (pO2)4,5. Here, by contrast, we show by means of a series of Earth system model experiments how continental rearrangement during the Phanerozoic Eon drives profound variations in ocean oxygenation and induces a fundamental decoupling in time between upper-ocean and benthic [O2]. We further identify the presence of state transitions in the global ocean circulation, which lead to extensive deep-ocean anoxia developing in the early Phanerozoic even under modern pO2. Our finding that ocean oxygenation oscillates over stable thousand-year (kyr) periods also provides a causal mechanism that might explain elevated rates of metazoan radiation and extinction during the early Palaeozoic Era6. The absence, in our modelling, of any simple correlation between global climate and ocean ventilation, and the occurrence of profound variations in ocean oxygenation independent of atmospheric pO2, presents a challenge to the interpretation of marine redox proxies, but also points to a hitherto unrecognized role for continental configuration in the evolution of the biosphere.

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Fig. 1: Earth system model results for simulations at 2,240 ppm CO2 (series #1).
Fig. 2: Benthic oxygen concentrations for simulations at 2,240 ppm CO2 (series #1).
Fig. 3: Earth system model results for simulations in which we varied pCO2 to approximately ‘correct’ for the palaeogeographical impacts on climate (series #2).

Code availability

The version of the cGENIE code used in this paper is tagged as release v0.9.31 and is available at Necessary boundary condition files are included as part of the code release. Configuration files for the specific experiments presented in the paper can be found in the installation subdirectory: genie-userconfigs/PUBS/published/Pohl_et_al.2022. Details of the experiments, plus the command line needed to run each one, are given in the readme.txt file in that directory. A manual describing code installation, basic model configuration and an extensive series of tutorials is provided ( The FOAM output is hosted on Zenodo (


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We thank J. Mossinger for editorial handling. This project has received financing from the European Union’s Horizon 2020 research and innovation programme under Marie Skłodowska-Curie grant agreement no. 838373. Calculations were partly performed using HPC resources from DNUM CCUB (Centre de Calcul de l’Université de Bourgogne). A.R. acknowledges support from NSF grants 1736771 and EAR-2121165, as well as from the Heising-Simons Foundation. This is a contribution to UNESCO project IGCP 735 ‘Rocks and the Rise of Ordovician Life (Rocks n’ ROL)’.

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Authors and Affiliations



A.P. and A.R. designed the study and wrote the manuscript, with input from all co-authors. A.P. and A.R. conducted the FOAM and cGENIE experiments. A.P., A.R. and A.K. led the analysis of the model results. C.R.S. produced the continental reconstructions.

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Correspondence to Alexandre Pohl.

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Nature thanks Tais Dahl, Shuhai Xiao, Sabin Zahirovic and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

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Extended data figures and tables

Extended Data Fig. 1 Comparison of modern observations of seafloor [O2] with the cGENIE model.

Results are shown for different (modern) continental grids, boundary conditions and biogeochemical cycling parameterization assumptions. Projection is equal-area rectangular and the colour scale is chosen to approximately match that in Fig. 2. a, Present-day distribution of [O2] globally at the seafloor, for which we re-grid the World Ocean Atlas 2018 (ref. 56) data to the modern continental grid of ref. 47 and show the oxygen concentration in the deepest model grid point. Apparent is both the production and southward propagation (via North Atlantic Deep Water (NADW)) of highly oxygenated waters from the North Atlantic and oxygenated deep-water production and mixing around Antarctica. b, Benthic [O2] distribution for the ‘standard’ modern ocean circulation of ref. 47 run at 278 ppm CO2, plus simplified biological export scheme. The large-scale patterns of benthic [O2] are reasonably reproduced, with the exception of too weak mixing of oxygen around the Southern Ocean, itself caused by a weak simulated Antarctic Circumpolar Current (ACC) and the difficulty in adequately representing the Drake Passage at this resolution. The slightly too low compared with observations values in the North Pacific is a further consequence of this. c, Test of substituting the simplified biological export scheme of ref. 47 with the explicit ecosystem model used here in the Phanerozoic series of simulations (but still at 278 ppm CO2). The slightly greater export simulated by the ecological model reduces benthic [O2] by about 10–20 µmol kg−1 while leaving the large-scale patterns largely unaltered. d, Deep-sea oxygenation in the 0 Ma Phanerozoic simulation (as per Fig. 2, 0 Ma) run at 2,240 ppm CO2. Without flux adjustment applied in the simplified cGENIE 2D EMBM atmosphere (see ref. 57), there is virtually no NADW, explaining the relatively poor and south-to-north oxygenation of the Atlantic. The Indian Ocean is also too poorly oxygenated, although the general pattern in the Pacific is reproduced. There are several main reasons for this model–data mismatch. First, directly re-gridding from a relatively coarse resolution GCM (FOAM) creates a highly restricted and shallow Drake Passage (Extended Data Fig. 7a, 0 Ma), precluding a strong ACC forming. More pertinently, the Phanerozoic simulation series (Fig. 2) are all run at 2,240 ppm. The resulting much-warmer-than-modern ocean is associated with the absence of any sea-ice formation and lower seawater oxygen solubility, probably explaining at least some of the spatial pattern and much of the lower global mean [O2] inventory (Extended Data Fig. 8a, 0 Ma). In terms of ventilation and water mass idealized mean age (not shown), the Atlantic, lacking an Atlantic MOC is far too old, whereas the Indian and Pacific oceans are similar to the inverse modelling of ref. 58, despite the aforementioned issues with the Drake Passage and that the 0 Ma simulation is run at 2,240 ppm CO2.

Extended Data Fig. 2 Selected redox proxy data versus the corresponding model oxygenation realization.

Benthic oxygen concentrations for simulations at 2,240 ppm CO2 (series #1). Eckert IV projection. Emerged continental masses are shaded white. Results are averaged over the last 5,000 years. Black (white) dots represent anoxic (oxic) conditions and grey points represent possible or intermittent anoxia, for 100 Ma after ref. 28, for 120 Ma after ref. 27, for 180 Ma after ref. 30, for 260 Ma after ref. 26, for 380 and 500 Ma after ref. 59 and for 440 Ma after ref. 60. These time slices have been chosen to represent regularly spaced periods during the Phanerozoic, typified by OAEs. The last 100 Myr are deliberately omitted because (1) the ocean is, overall, well oxygenated during this time interval and (2) our constant boundary conditions (deliberately chosen to isolate the role of tectonics from climate, see main text) do not (and, indeed, do not intend to) reproduce the pronounced cooling trend through the Cenozoic (especially from Early Eocene Climatic Optimum (ca. 50 Ma) onwards), precluding direct model–data comparison.

Extended Data Fig. 3 Deep-ocean circulation for simulations at 2,240 ppm CO2 (series #1).

a, Meridional overturning stream function, in Sv (sverdrup, 1 Sv = 106 m3 s−1). A negative (blue) stream function corresponds to an anticlockwise circulation. b, Annual distribution of convective adjustments across the water column. Emerged continental masses are shaded white. Eckert IV projection. Results are averaged over the last 5,000 years.

Extended Data Fig. 4 Sensitivity of benthic [O2] to the continental reconstruction.

Benthic oxygen concentrations for simulations at 2,240 ppm CO2 (such as series #1) at 440 Ma (a) and 460 Ma (b), using the continental reconstructions of BugPlates31, with topography/bathymetry after ref. 61. Eckert IV projection. Emerged continental masses are shaded white. Results are averaged over the last 5,000 years. Panels a and b are identical to the 440 Ma and 460 Ma panels of Fig. 2, except that simulations have been conducted using another continental reconstruction. Note that, although we simulate sulphate reduction in cGENIE, with SO42− being used as the electron acceptor for the remineralization of organic matter in the ocean interior once dissolved O2 has become depleted (see ref. 48), small negative O2 concentrations can arise when several geochemical reactions compete simultaneously for the same depleted oxygen pool. However, because the product of sulphate reduction—hydrogen sulphide (H2S)—has fast oxidation kinetics in the presence of free oxygen, the transport of H2S from anoxic to oxic areas closely mirrors the transport and fate of ‘negative oxygen’ (see ref. 62) and the overall redox landscape is largely independent of this small modelled oxygen overconsumption.

Extended Data Fig. 5 Sensitivity to the remineralization scheme.

Earth system model results for simulations at 2,240 ppm CO2 (such as #1) but with no dependence of remineralization on temperature. a, Same as Fig. 1. b, Same as Fig. 2.

Extended Data Fig. 6 Sensitivity of benthic [O2] to deep-ocean bathymetry.

Benthic oxygen concentrations for simulations at 2,240 ppm CO2 (such as #1) but with no mid-ocean ridges (see Extended Data Fig. 7). Results are averaged over the last 5,000 years. Emerged continental masses are shaded white. Eckert IV projection.

Extended Data Fig. 7 Bathymetric reconstructions.

Bathymetry with mid-ocean ridges used in series #1 and #2 (a) and flat-bottomed bathymetric reconstructions used in simulations with no mid-ocean ridges (b) (see Extended Data Fig. 6). Only reconstructions for 0 to 140 Ma (both included) differ (see Methods). Emerged continental masses are shaded white. Eckert IV projection.

Extended Data Fig. 8 Time evolution of benthic [O2].

a, Simulations at 2,240 ppm CO2 with temperature-dependent remineralization (series #1). b, Simulations in which we varied pCO2 to approximately ‘correct’ for the palaeogeographical impacts on climate (series #2).

Extended Data Fig. 9 Sensitivity of benthic [O2] to atmospheric forcing (pCO2).

Benthic oxygen concentrations for simulations at 1,120 ppm CO2, with temperature-dependent remineralization and mid-ocean ridges. Emerged continental masses are shaded white. Results are averaged over the last 5,000 years. Eckert IV projection.

Extended Data Fig. 10 Ocean circulation regimes in the cGENIE model.

a, Envelope of benthic [O2] values simulated at various atmospheric CO2 levels using Drake world (in blue, see maps in panel b) and ridge world (in black) (Methods). Regimes of stable equilibria and regimes of stable oscillations for Drake world simulations are numbered and labelled using a blue and a red background, respectively. b, Meridional overturning stream function in Sv (sverdrup, 1 Sv = 106 m3 s−1) and map of benthic ventilation age for each stable equilibrium and the two extreme states of each stable oscillatory regime identified in panel a, using the same numbering and colour coding. Lambert cylindrical equal-area projection. A negative (blue) stream function corresponds to an anticlockwise circulation. c, Tentative comparison with the latest Ordovician simulations of Pohl et al.21. Results are shown at 6,720 ppm and 2,380 ppm CO2, corresponding to the warm and cold states of scenario #1 in ref. 21.

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Pohl, A., Ridgwell, A., Stockey, R.G. et al. Continental configuration controls ocean oxygenation during the Phanerozoic. Nature 608, 523–527 (2022).

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