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COPSE reloaded: An improved model of biogeochemical cycling over Phanerozoic time
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“The first-order relationship between global marine new production (newp′), global extent of anoxia (AANOX) and atmospheric pO2 (o′) has been obtained from Earth system ocean–atmosphere models (57):”
From Atmosphere-ocean oxygen and productivity dynamics during early animal radiations (2019) · cites this paper as [57]
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“The constants kanox = 12, ku = 0.5 are determined so that 0.25% of the seafloor is overlain by anoxic water masses similar to modern productivity and oxygen levels (newp′ = o′ = 1) and the formula gives a fairly sharp transition of AANOX from 23 to 77% as atmospheric pO2 declines from 13 to 8 atm%, consistent with biogeochemical models for the spatial distribution of oxygen in the oceans (57, 58).”
From Atmosphere-ocean oxygen and productivity dynamics during early animal radiations (2019) · cites this paper as [57]
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Stepwise Earth oxygenation is an inherent property of global biogeochemical cycling
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Self-citationEvolution of Earth’s tectonic carbon conveyor belt
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Evolution of Atmospheric O2 Through the Phanerozoic, Revisited
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“S ee c op yr ig ht f or a pp ro ve d us e. biogeochemistry and climate (e.g., COPSE: Bergman et al. 2004, Lenton et al. 2018; MAGic: Arvidson et al. 2013; SCION: Mills et al. 2021).”
BackgroundMethodSelf-citationAtmosphere oxygen cycling through the Proterozoic and Phanerozoic
Large, R. R., Mukherjee, I., Gregory, D., Steadman, J., Corkrey, R., & Danyushevsky, L. V. (2019) · Mineralium Deposita · doi:10.1007/s00126-019-00873-9 · Cited by 122
Reconciling proxy records and models of Earth's oxygenation during the Neoproterozoic and Palaeozoic
Tostevin, R., & Mills, B. J. W. (2020) · Interface Focus · doi:10.1098/rsfs.2019.0137 · Cited by 128
“Recent reviews are available that describe the latest version of the model [65,66].”
BackgroundSelf-citationUranium isotopes in marine carbonates as a global ocean paleoredox proxy: A critical review
Zhang, F., Lenton, T. M., del Rey, Á., Romaniello, S. J., Chen, X., Planavsky, N. J., Clarkson, M. O., Dahl, T. W., Lau, K., Wang, W., et al. (2020) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2020.05.011 · Cited by 119
Self-citationAtmosphere-ocean oxygen and productivity dynamics during early animal radiations
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Selection for Gaia across Multiple Scales
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Self-citationUncovering the Ediacaran phosphorus cycle
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Self-citationWarm afterglow from the Toarcian Oceanic Anoxic Event drives the success of deep-adapted brachiopods
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Self-citationThe origin and early evolution of plants
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Drivers of the global phosphorus cycle over geological time
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“Mafic rocks generally contain more 83 , plant effect on weathering (black) 19 and global average temperature (GAT) (grey) 82,84 (shading represents the 95% confidence interval in GAT) (panel d ); and phosphorus content of average weatherable crust 17 (shading represents the standard error) (panel e…”
InfluentialSelf-citationThe History of Ocean Oxygenation
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“In panel b, the small light blue bars show empirical constraints from iron/phosphorus ratios in iron-rich chemical sediments, and the larger bars (<1.8 Gya) show model-derived values from Ozaki et al. (2019) (dark blue) and Lenton et al. (2018) (green).”
BackgroundInfluentialSpatial continuous integration of Phanerozoic global biogeochemistry and climate
Mills, B. J. W., Yannick Donnadieu, A., & Goddéris, Y. (2021) · Gondwana Research · doi:10.1016/j.gr.2021.02.011 · Cited by 93
“As discussed in Lenton et al. (2018) this change makes little difference to the model biogeochemistry predictions, but the addition of sulfur degassing fl uxes were not compatible with the COPSE calcium cycle.”
BackgroundMethodResultInfluentialSelf-citationA nutrient control on expanded anoxia and global cooling during the Late Ordovician mass extinction
Qiu, Z., Zou, C., Mills, B. J. W., Xiong, Y., Tao, H., Lu, B., Liu, H., Xiao, W., & Poulton, S. W. (2022) · Communications Earth & Environment · doi:10.1038/s43247-022-00412-x · Cited by 90
Self-citationEvolution of the structure and impact of Earth’s biosphere
Planavsky, N. J., Crowe, S. A., Fakhraee, M., Beaty, B., Reinhard, C. T., Mills, B. J. W., Holstege, C., & Konhauser, K. O. (2021) · Nature Reviews Earth & Environment · doi:10.1038/s43017-020-00116-w · Cited by 102
Self-citationA long-term record of early to mid-Paleozoic marine redox change
Sperling, E. A., Melchin, M. J., Fraser, T., Stockey, R. G., Farrell, U. C., Bhajan, L., Brunoir, T. N., Cole, D. B., Gill, B. C., Lenz, A., et al. (2021) · Science Advances · doi:10.1126/sciadv.abf4382 · Cited by 85
Early Palaeozoic ocean anoxia and global warming driven by the evolution of shallow burrowing
van de Velde, S., Mills, B. J. W., Meysman, F. J. R., Lenton, T. M., & Poulton, S. W. (2018) · Nature Communications · doi:10.1038/s41467-018-04973-4 · Cited by 87
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“…per unit of photosynthate carbon (C) received at CO2 concentrations simulating the atmosphere of the early to mid-Palaeozoic under which land plants first evolved (1500 ppm atmospheric CO2 (a[CO2]); Berner, 2006; Lenton et al., 2018) than under modern-day a[CO2] of 440 ppm (Field et al., 2016).”
BackgroundMethodRevisiting the Great Ordovician Diversification of land plants: Recent data and perspectives
Servais, T., Cascales-Miñana, B., Cleal, C. J., Gerrienne, P., Harper, D. A. T., & Neumann, M. (2019) · Palaeogeography, Palaeoclimatology, Palaeoecology · doi:10.1016/j.palaeo.2019.109280 · Cited by 77
What controls the duration and intensity of ocean anoxic events in the Paleozoic and the Mesozoic?
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Land plant evolution decreased, rather than increased, weathering rates
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Marine anoxia linked to abrupt global warming during Earth’s penultimate icehouse
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The impact of marine nutrient abundance on early eukaryotic ecosystems
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Extreme variability in atmospheric oxygen levels in the late Precambrian
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Self-citationNew paleogeographic and degassing parameters for long-term carbon cycle models
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A tectonically driven Ediacaran oxygenation event
Williams, J., Mills, B. J. W., & Lenton, T. M. (2019) · Nature Communications · doi:10.1038/s41467-019-10286-x · Cited by 70
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The Evolution Pathway of Ammonia-Oxidizing Archaea Shaped by Major Geological Events
Yang, Y., Zhang, C., Lenton, T. M., Yan, X., Zhu, M., Zhou, M., Tao, J., Phelps, T. J., & Cao, Z. (2021) · Molecular Biology and Evolution · doi:10.1093/molbev/msab129 · Cited by 66
“2018) and the COPSE reloaded model (dark blue) (Lenton et al. 2018).”
BackgroundInfluentialSelf-citationVertical decoupling in Late Ordovician anoxia due to reorganization of ocean circulation
Pohl, A., Lu, Z., Lu, W., Stockey, R., Elrick, M., Li, M., Desrochers, A., Shen, Y., He, R., Finnegan, S., et al. (2021) · Nature Geoscience · doi:10.1038/s41561-021-00843-9 · Cited by 65
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“T. M. Lenton, S. J. Daines, B. J. W. Mills, COPSE reloaded: An improved model of biogeochemical cycling over Phanerozoic time.”
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“This model rapidly became the centre of theoretical discussion about Gaia (Betts and Lenton, 2007; Lenton et al., 2018a; Lenton and Lovelock, 2001; McDonald-Gibson et al., 2008; Volk, 2002; Watson and Lovelock, 1983).”
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The role of calcium in regulating marine phosphorus burial and atmospheric oxygenation
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Oxygenation of the Earth aided by mineral–organic carbon preservation
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Self-citationRecent Advances in Geochemical Paleo-Oxybarometers
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Self-citationJurassic greenhouse ice-sheet fluctuations sensitive to atmospheric CO2 dynamics
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The future lifespan of Earth’s oxygenated atmosphere
Ozaki, K., & Reinhard, C. T. (2021) · Nature Geoscience · doi:10.1038/s41561-021-00693-5 · Cited by 56
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LeRoy, M. A., Gill, B. C., Sperling, E. A., McKenzie, N. R., & Park, T. S. (2021) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2020.110209 · Cited by 53
Wei, G., Zhao, M., Sperling, E. A., Gaines, R. R., Kalderon-Asael, B., Shen, J., Li, C., Zhang, F., Li, G., Zhou, C., et al. (2024) · Science Advances · doi:10.1126/sciadv.adk2152 · Cited by 56
“Initial atmospheric O 2 levels were set at 5 to 40% PAL, based on previous estimates generated by well-established numerical models ( 82 , 85 ).”
MethodUpper limits on the extent of seafloor anoxia during the PETM from uranium isotopes
Clarkson, M. O., Lenton, T. M., Andersen, M. B., Bagard, M.-L., Dickson, A. J., & Vance, D. (2021) · Nature Communications · doi:10.1038/s41467-020-20486-5 · Cited by 55
Self-citationSproson, A. D., von Strandmann, P. A. E. P., Selby, D. S., Jarochowska, E., Frýda, J., Hladil, J., Loydell, D. K., Slavík, L., Calner, M., Maier, G., et al. (2021) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2021.117260 · Cited by 49
Self-citationCauses and consequences of the Messinian salinity crisis
Krijgsman, W., Rohling, E. J., Palcu, D. V., Raad, F., Amarathunga, U., Flecker, R., Florindo, F., Roberts, A. P., Sierro, F. J., & Aloisi, G. (2024) · Nature Reviews Earth & Environment · doi:10.1038/s43017-024-00533-1 · Cited by 53
“Most biogeochemical models have ignored evaporite-driven perturbations to seawater chemistry 148,149 , assuming that over time-scales >100 kyr, evaporite precipitation and weathering are balanced.”
Mays, C., & McLoughlin, S. (2022) · Palaios · doi:10.2110/palo.2021.051 · Cited by 50
“…estimate only a modest reduction in pO2 from pre-EPE levels, and retention at consistently higher values than modern pO2 for the Early Triassic (Krause et al. 2018; Lenton et al. 2018; Mills et al. 2021), suggesting that oxygen was not the primary limiting factor to fire ignition and spread.”
BackgroundAtmospheric oxygen of the Paleozoic
Brand, U., Davis, A., Shaver, K. K., Blamey, N., Heizler, M., & Lécuyer, C. (2021) · Earth-Science Reviews · doi:10.1016/j.earscirev.2021.103560 · Cited by 34
Rates of seafloor and continental weathering govern Phanerozoic marine phosphate levels
Sharoni, S., & Halevy, I. (2022) · Nature Geoscience · doi:10.1038/s41561-022-01075-1 · Cited by 44
Carbon isotope evidence for the global physiology of Proterozoic cyanobacteria
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“The earliest land plants evolved in a world where atmospheric CO 2 levels were seven times higher and oxygen levels were 1.4 times lower than today (Lenton et al. 2018).”
BackgroundInfluentialFire-prone Rhamnaceae with South African affinities in Cretaceous Myanmar amber
Shi, C., Wang, S., Cai, H., Zhang, H., Long, X., Tihelka, E., Song, W., Feng, Q., JIANG, R.-X., Cai, C., et al. (2022) · Nature Plants · doi:10.1038/s41477-021-01091-w · Cited by 43
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Decoupled oxygenation of the Ediacaran ocean and atmosphere during the rise of early animals
Shi, W., Mills, B. J. W., Li, C., Poulton, S. W., Krause, A. J., He, T., Zhou, Y., Cheng, M., & Shields, G. (2022) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2022.117619 · Cited by 43
Self-citationSilurian wildfire proxies and atmospheric oxygen
Glasspool, I. J., & Gastaldo, R. A. (2022) · Geology · doi:10.1130/g50193.1 · Cited by 41
Babakhani, P., Dale, A. W., Woulds, C., Moore, O., Xiao, K., Curti, L., & Peacock, C. L. (2025) · Nature Geoscience · doi:10.1038/s41561-024-01606-y · Cited by 43
Earth History of Oxygen and the iprOxy
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Krissansen‐Totton, J., Kipp, M. A., & Catling, D. C. (2021) · Geobiology · doi:10.1111/gbi.12440 · Cited by 38
“…et al., 2009); however, the total carbonate burial flux may be as high as 30– 40 Tmol C/year (Milliman, 1993), which would imply a carbonate weathering flux around 20– 30 Tmol C/year, assuming a ~10 Tmol C/year contribution from silicate weathering (Gaillardet et al., 1999; Lenton et al., 2018).”
BackgroundMethodResultInfluentialZhao, Z., Pang, X., Zou, C., Dickson, A. J., Basu, A., Guo, Z., Pan, S., Nielsen, A. T., Schovsbo, N. H., Jing, Z., et al. (2023) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2023.118013 · Cited by 39
Limited Carbon Cycle Response to Increased Sulfide Weathering Due to Oxygen Feedback
Maffre, P., Swanson‐Hysell, N. L., & Goddéris, Y. (2021) · Geophysical Research Letters · doi:10.1029/2021gl094589 · Cited by 40
“Hence, geochemical cycle models, such as COPSE (Lenton et al., 2018), generally assume those fluxes to be proportional to sulfate concentration.”
MethodZhao, C., Zhang, K., Xiao, L., Uhl, D., Shi, Z., Zhao, W., Zhao, Q., Sun, Y., & Liu, B. (2023) · Chemical Geology · doi:10.1016/j.chemgeo.2023.121677 · Cited by 36
Dzombak, R., & Sheldon, N. D. (2020) · Soil Systems · doi:10.3390/soilsystems4040073 · Cited by 36
“Prior to the evolution and expansion of land plants, changes in the flux of P from continents to oceans are thought to have been a primary control on marine productivity, and consequently, on the concentration of oxygen in the atmosphere [10,11].”
BackgroundBelcher, C. M., Mills, B. J. W., Vitali, R., Baker, S. J., Lenton, T. M., & Watson, A. J. (2021) · Nature Communications · doi:10.1038/s41467-020-20772-2 · Cited by 39
Self-citationBackground Earth system state amplified Carnian (Late Triassic) environmental changes
Corso, J. D., Mills, B. J. W., Chu, D., Newton, R. J., & Song, H. (2021) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2021.117321 · Cited by 37
Self-citationLithium isotope behaviour during basalt weathering experiments amended with organic acids
von Strandmann, P. A. E. P., Liu, X., Liu, C., Wilson, D. J., Hammond, S. J., Tarbuck, G. M., Aristilde, L., Krause, A. J., & Fraser, W. T. (2022) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2022.04.032 · Cited by 36
“…caused significant global increases in chemical weathering, leading to an increased drawdown of atmospheric CO 2 , and potentially triggering cold periods and mass extinctions in the Devonian-Carboniferous (Berner, 1998, 2006; Dahl and Arens, 2020; Lenton et al., 2018; Lenton et al., 2012).”
BackgroundColwyn, D. A., Sheldon, N. D., Maynard, J. B., Gaines, R. R., Hofmann, A., Wang, X., Guéguen, B., Asael, D., Reinhard, C. T., & Planavsky, N. J. (2019) · Geobiology · doi:10.1111/gbi.12360 · Cited by 36
Ordovician cyanobacterial calcification: A marine fossil proxy for atmospheric CO2
Liu, L., Liang, L., Wu, Y., Zhou, X., Jia, L., & Riding, R. (2019) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2019.115950 · Cited by 33
Iron and sulfur cycling in the cGENIE.muffin Earth system model (v0.9.21)
van de Velde, S., Hülse, D., Reinhard, C. T., & Ridgwell, A. J. (2021) · Geoscientific model development · doi:10.5194/gmd-14-2713-2021 · Cited by 45
Merdith, A., Atkins, S., & Tetley, M. G. (2019) · Frontiers in Earth Science · doi:10.3389/feart.2019.00332 · Cited by 35
Functional traits of fossil plants
McElwain, J. C., Matthaeus, W. J., Barbosa, C., Chondrogiannis, C., Dea, K. O., Jackson, B., Knetge, A. B., Kwaśniewska, K., Nair, R. K. F., White, J. D., et al. (2024) · New Phytologist · doi:10.1111/nph.19622 · Cited by 37
On the geological and scientific legacy of petrogenic organic carbon
Blattmann, T. M., Letsch, D., & Eglinton, T. I. (2018) · American Journal of Science · doi:10.2475/08.2018.02 · Cited by 33
Evaporite weathering and deposition as a long-term climate forcing mechanism
Shields, G., & Mills, B. J. W. (2020) · Geology · doi:10.1130/g48146.1 · Cited by 35
“…(halides, sulfates) into rivers unaccompanied by carbonate alkalinity, for example: In established long-term biogeochemical models (Berner, 1991; Lenton et al., 2018; Mills et al., 2018), evaporite fluxes are negated by assuming a close balance between the weathering and deposition of Ca-and…”
Self-citationA persistent non-uniformitarian paleomagnetic field in the Devonian?
van der Boon, A., Biggin, A. J., Thallner, D., Hounslow, M. W., Bono, R. K., Nawrocki, J., Wójcik, K., Paszkowski, M., Königshof, P., Backer, T. D., et al. (2022) · Earth-Science Reviews · doi:10.1016/j.earscirev.2022.104073 · Cited by 33
Respiratory evolution in archosaurs
Brocklehurst, R. J., Schachner, E. R., Codd, J. R., & Sellers, W. I. (2020) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2019.0140 · Cited by 33
Zhou, N., Wang, Y., Li, Y., Porter, A. S., Kürschner, W. M., Li, L., Lu, N., & McElwain, J. C. (2019) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2019.109547 · Cited by 33
Zheng, X., Beard, B. L., Neuman, M., Fahnestock, M. F., Bryce, J. G., & Johnson, C. M. (2022) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2022.117653 · Cited by 33
Ice, Fire, or Fizzle: The Climate Footprint of Earth's Supercontinental Cycles
Jellinek, M. A., Lenardic, A., & Pierrehumbert, R. T. (2020) · Geochemistry Geophysics Geosystems · doi:10.1029/2019gc008464 · Cited by 31
“…Hayes & Waldbauer, 2006; Higgins & Schrag, 2003; Jagoutz et al., 2016; Johnston et al., 2012; Kump, 2018; Kump et al., 2000; Le Hir et al., 2009; Lenton et al., 2018; Mills et al., 2014, 2014; Roberson et al., 2011; Ridgwell & Zeebe, 2005; Rothman, 2015; Rothman et al., 2003; Rothman & Forney,…”
BackgroundMethodResultInfluentialA generic hierarchical model of organic matter degradation and preservation in aquatic systems
Shang, H. (2023) · Communications Earth & Environment · doi:10.1038/s43247-022-00667-4 · Cited by 31
Porter, A. S., Evans-Fitz.Gerald, C., Yiotis, C., Montañez, I. P., & McElwain, J. C. (2019) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2019.05.037 · Cited by 32
Schwieterman, E. W., Olson, S. L., Pidhorodetska, D., Reinhard, C. T., Ganti, A., Fauchez, T. J., Bastelberger, S., Crouse, J., Ridgwell, A. J., & Lyons, T. W. (2022) · The Astrophysical Journal · doi:10.3847/1538-4357/ac8cfb · Cited by 36
“Steady-state P levels are unlikely to have dramatically exceeded ∼2 times POL at any point in Earth’s history (Reinhard et al. 2017b; Lenton et al. 2018; Reinhard & Planavsky 2022).”
BackgroundMercury isotopes show vascular plants had colonized land extensively by the early Silurian
Yuan, W., Liu, M., Chen, D., Xing, Y., Spicer, R. A., Chen, J., Them, T. R., Wang, X., Li, S., Guo, C., et al. (2023) · Science Advances · doi:10.1126/sciadv.ade9510 · Cited by 30
“Profound impact on the atmosphere and surface of Earth Despite some differences in the estimation of absolute oxygen content, both COPSE ( 40 ) and GEOCARBONSULF ( 23 ) models suggest two episodes of distinct increase of atmospheric p O 2 ; one was from the Ordovician to the end of the Silurian,…”
Implications of different stellar spectra for the climate of tidally locked Earth-like exoplanets
Eager-Nash, J. K., Reichelt, D. J., Mayne, N. J., Lambert, F. H., Sergeev, D. E., Ridgway, R. J., Manners, J. C., Boutle, I., Lenton, T. M., & Kohary, K. (2020) · Astronomy and Astrophysics · doi:10.1051/0004-6361/202038089 · Cited by 34
Self-citationSpatial pattern of marine oxygenation set by tectonic and ecological drivers over the Phanerozoic
Wang, X., Algeo, T. J., Li, C., & Zhu, M. (2023) · Nature Geoscience · doi:10.1038/s41561-023-01296-y · Cited by 28
“2h), which promoted organic burial in terrestrial environments and elevated atmospheric p O ,12,15 , leading to higher surface-ocean dissolved O 2 concentrations, while concurrently enhancing nutrient fluxes to the ocean 16,17 , thus boosting marine primary productivity and O 2 consumption in the…”
BackgroundEarly Triassic super-greenhouse climate driven by vegetation collapse
Xu, Z., Yu, J., Yin, H., Merdith, A., Hilton, J. M., Allen, B. J., Gurung, K., Wignall, P. B., Dunhill, A. M., Shen, J., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-60396-y · Cited by 27
“57 – 59 for a range of estimates of this factor, and Methods 10 and 11 for model runs with different factors).”
Self-citation
Metadata
- Authors
- Lenton, T. M., Daines, S. J., & Mills, B. J. W.
- Affiliations
- University of Exeter (GB)
- University of Leeds (GB)
- Corresponding author
- Lenton, Timothy M.
- Venue
- Earth-Science Reviews
- Publisher
- Elsevier BV
- Published
- 2018-03
- Volume
- 178
- Pages
- 1-28
- ISSN
- 0012-8252 (ISSN-L)
- OpenAlex
- W2772093563
- Cited by
- 350
- References
- 141
- Citations by year
- 2026: 40 · 2025: 46 · 2024: 42 · 2023: 53 · 2022: 57 · 2021: 37 · 2020: 37 · 2019: 27 · 2018: 10
- FWCI
- 14.43
- Type
- journal article
- Language
- en
- Open access
- green
- hdl.handle.net
- Locations
- Earth-Science Reviews (publishedVersion)
- White Rose Research Online (University of Leeds, The University of Sheffield, University of York) (acceptedVersion)
- Open Research Exeter (University of Exeter) (submittedVersion)
- Figshare (submittedVersion)
- Funders
- University of Leeds
- Natural Environment Research Council (NE/P013651/1, NE/N018508/1)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Stratigraphy of Fossils
- Physical Sciences › Earth and Planetary Sciences › Atmospheric Science › Geology and Paleoclimatology Research
- Physical Sciences › Environmental Science › Global and Planetary Change › Earth Systems and Cosmic Evolution
- Keywords
- atmospheric oxygen, atmospheric CO2, sulfate, strontium isotopes, ocean anoxia, seafloor weathering, volcanic rock weathering, land plants, carbon cycle modeling
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