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Earliest land plants created modern levels of atmospheric oxygen
Open accessAbstract
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.
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TL;DR
The paper argues that the earliest land plants, colonizing from ∼470 Ma, drove a mid-Paleozoic oxygenation event by increasing organic carbon burial, raising atmospheric O₂ to present levels by 420–400 Ma. It uses a revised COPSE biogeochemical model plus a trait-based cryptogamic vegetation model and bryophyte C/P data to reproduce a sustained +2‰ δ¹³C shift and O₂ rise consistent with charcoal and redox proxies.
Key findings
- Earliest nonvascular plants colonized land from ∼470–445 Ma; the model reproduces a sustained +2‰ δ¹³C increase by ∼445 Ma and an O₂ rise to present levels by 420–400 Ma, consistent with fossil charcoal indicating O₂ >15–17% by 420–400 Ma and Mo isotope evidence for deep-ocean oxygenation at 435–392 Ma.
- A trait-based ecophysiological model predicts cryptogamic vegetation cover at 445 Ma (CO₂ = 8 PAL, O₂ = 0.6 PAL) could achieve global NPP = 18.7 GtC·y⁻¹, ∼30% of today; COPSE with stronger Late Ordovician colonization predicts 30–40% of today’s NPP.
- Bryophyte C/P literature ranges from 800 to 4,300 (mean ∼1,900); assuming C/P = 2,000 vs 1,000 increases modeled organic carbon burial by ∼50% vs ∼25%, δ¹³C by +1.1‰ vs +0.5‰, and O₂ by +0.27 vs +0.11 PAL.
- Reproducing the observed +2‰ δ¹³C shift requires selective P weathering: a 50% increase in P weathering with C/P = 1,000 gives burial ∼95%, δ¹³C +2.2‰, and O₂ +0.74 PAL (to 0.97 PAL at 417 Ma); a 25% increase with C/P = 2,000 gives burial ∼90%, δ¹³C +2.1‰, and O₂ +0.67 PAL; P weathering spikes with C/P = 2,000 peak at O₂ +0.72 PAL at 407 Ma.
Methods
- Revised COPSE biogeochemical model for coupled C, O, N, P, and S cycles and δ¹³C, with biological forcing factors: evolution/colonization (E), weathering enhancement (W), selective phosphorus weathering (F), and terrestrial C/P burial ratio (CP); sensitivity analyses varied geologic forcing and initial conditions.
- Trait-based spatial model of cryptogamic vegetation (bryophyte and lichen) driven by Late Ordovician (445 Ma) climate simulations at different atmospheric CO₂ and O₂; assumed CO₂ = 8 PAL, O₂ = 0.6 PAL, and no ice sheet mask to predict global NPP.
- Data compilation included the early charcoal record, molybdenum isotope record, carbonate δ¹³C spline fit, and bryophyte C/P ratios; experimental P weathering calculation used granite with the moss *Physcomitrella patens*.
Limitations
- Stated: direct evidence linking a phase of land colonization to enhanced weathering and a positive δ¹³C excursion is only established for the Silurian–Devonian boundary; alternative hypotheses for short-lived positive δ¹³C excursions should also be considered.
- Stated: the model does not capture the earlier Late Ordovician–Early Silurian δ³⁴S rise from ∼25 to ∼30‰; other processes not yet included warrant future consideration, such as the effect of increasing atmospheric mass on climate and effects on δ⁷Li and ⁸⁷Sr/⁸⁶Sr.
- Evident: P weathering amplification estimates are dominated by the unmeasured P content of biomass, though the factor must be >5.3; the hypothesis requires early plants to have been much more productive and globally extensive than previously assumed.
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Claims the paper makes
33 Not checked yet · 20 Own interpretation · 10 Own result · 7 No basis given · 2 Own method · 1 Supported so far
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In the paper
“The first appearance of fossil charcoal in the Late Silurian (11) and its ongoing occurrence through the Devonian (12) (Table S1), albeit rare and at low concentrations, indicates O2 > 15 -17% (vol) of the atmosphere (13) (or O2 > ∼ 0.7 PAL assuming a constant N2 reservoir) already by ∼ 420 -400 Ma.”
What the sources say
- Not checked yetcites [11] source
- Supportedcites [12] source
The Prídolí–Emsian section says that charcoal occurrences around 420–400 Ma indicate oxygen above present atmospheric levels, exceeding the cited 15–17% threshold. It also reports modeled values of 23%, 25%, and 22% at 420, 410, and 400 Ma, respectively. The paper does not extend that high-oxygen inference to the entire Devonian. (LLM, conf=0.90)
- Not checked yetcites [13] Belcher CM, McElwain JC (2008) Limits for combustion in low O2 redefine paleoatmospheric predictions for the Mesozoic. Science 321(5893):1197 -1200. source
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In the paper
“Data from modern bryophytes suggests this plentiful early plant material had a much higher molar C:P ratio ( ∼ 2,000) than marine biomass ( ∼ 100)”
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In the paper
“The original baseline model (10) predicts early Paleozoic O2 ∼ 0.23 PAL at a reference time of 445 Ma, supported by an organic carbon burial flux of ∼ 4 × 10 12 mol · y -1 (about half the present-day value) with δ 13 C = 0.03 ‰ .”
What the sources say
- Not checked yetcites [10] source
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In the paper
“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”
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- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“and predict a corresponding rise in O2 to present levels by 420 -400 Ma, consistent with geochemical data.”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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.”
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In the paper
“At atmospheric CO2 = 8 PAL, consistent with Late Ordovician glaciations (20), predicted global NPP is ∼ 19 GtC · y -1 (GtC, billion metric tons of carbon) (Fig. 2), ∼ 30% of today.”
What the sources say
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
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In the paper
“Alternatively, a series of P weathering spikes designed to reproduce the observed sequence of positive δ 13 C excursions (Fig. 1), combined with C/P = 2,000, produces a series of spikes in organic carbon burial and a peak increase of O2 of 0.72 PAL at 407 Ma (Fig. 3, black).”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
In the paper
“this gives a biotic P weathering amplification factor ∼ 24 (range 15 -43), whereas previously we suggested up to 60 (20).”
What the sources say
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“However, the P weathering amplification factor has to be > 5.3 (the ratio of dissolved phosphate entering solution in microcosms with moss to those without)”
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- It may state two separate claims as one. 33
- Probably not a substantial claim. 14
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In the paper
“If instead we assume that buried early plant matter had C/P = 2,000, then given their productivity, even with no effect on weathering fluxes, this increases global organic carbon burial by ∼ 50%, δ 13 C by 1.1 ‰ , and atmospheric O2 by 0.27 PAL (Fig. 3, cyan).”
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
None of the sources cited for this claim have been checked yet.
In the paper
“In the original COPSE model (10), predicted NPP only reaches ∼ 5% of today ' s value in the Late Ordovician and Silurian, but when we assume a stronger Late Ordovician phase of land colonization by nonvascular plants (following ref. 20; SI Materials and Methods ), then COPSE predicts global NPP 30 -40% of today (Fig. 3 A ), consistent with the detailed spatial model.”
What the sources say
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In the paper
“If we include in COPSE an effect of early plants on silicate weathering following ref. 20, assuming C/P = 1,000, this increases organic carbon burial by ∼ 35%, δ 13 C by 0.7 ‰ , and O2 by 0.18 PAL (Fig. 3, magenta).”
What the sources say
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In COPSE, this advent of early land plants alone, with no assumed effect on weathering fluxes, and assumed C/P = 1,000, increases total organic carbon burial by ∼ 25%, δ 13 C by 0.5 ‰ , and atmospheric O2 by 0.11 PAL (Fig. 3, blue).”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
In the paper
“However, we estimate much higher global NPP (Fig. 2) and weathering potential (32).”
What the sources say
- Not checked yetcites [32] Porada P, et al. (2016) High potential for weathering and climate effects of nonvascular vegetation in the Late Ordovician. Nat Commun , 10.1038/ncomms12113. source
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In the paper
“Predicted NPP is sensitive to variations in CO2 and climate (Fig. S1), ice sheet cover (Fig. S2), and O 2 (Table S2), but is consistently higher than the 4.3 GtC · y -1 (7% of today) estimated elsewhere (33).”
What the sources say
- Not checked yetcites [33] Edwards D, Cherns L, Raven JA (2015) Could land-based early photosynthesizing ecosystems have bioengineered the planet in mid-Palaeozoic times? Palaeontology 58(5): 803 -837. source
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In the paper
“The mean moss biomass in the biotic microcosms was 14.390 mg”
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In the paper
“Assuming that early plants caused a sustained 50% increase in phosphorus weathering relative to bulk rock dissolution, with C/P = 1,000, increases total organic carbon burial by ∼ 95%, δ 13 C by 2.2 ‰ , and O2 by 0.74 PAL (to 0.97 PAL at 417 Ma; Fig. 3, yellow).”
hedgeddirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Varying the uplift and degassing forcing of the model within plausible bounds only causes ± 0.08 PAL variation in O2 about the initial state (Fig. S4), although it does cause the effect of the same early plant forcing scenario to range over + 0.4 to 1.0 PAL O2 (Table S4).”
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In the paper
“The relatively high global NPP results obtained are consistent with present-day cryptogamic covers providing ∼ 7% of global NPP, despite making up only 1% of terrestrial vegetation by mass (90) and being restricted to relatively resource-poor habitats, while also operating in an atmosphere with a low CO2/O2 ratio.”
What the sources say
- Not checked yetcites [90] Elbert W, et al. (2012) Contribution of cryptogamic covers to the global cycles of carbon and nitrogen. Nat Geosci 5(7):459 -462. source
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In the paper
“Our model makes additional predictions that can be tested against geochemical data -notably, it predicts a decline in pyrite sulfur burial and associated drop in δ 34 S and increases in seawater SO4 concentration and C/S burial ratio with the rise of the earliest plants (Fig. S5).”
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In the paper
“Assuming that buried early plant matter had a higher C/P = 2,000 causes larger increases in total organic carbon burial ∼ 60%, δ 13 C + 1.2 ‰ , and atmospheric O2 + 0.35 PAL (Fig. 3, green).”
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Probably not a substantial claim. It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“the amplification factors for Ca = 1.4 and Mg = 1.5 from granite”
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In the paper
“Including an additional negative feedback on O2, from increased marine organic C/P burial ratios under anoxic waters (37), increases its initial early Paleozoic level to 0.54 PAL and reduces the effect of the same biological forcing scenarios on O2 by ∼ 10 -30%, giving a maximum increase of + 0.63 PAL (Table S5).”
What the sources say
- Not checked yetcites [37] Van Cappellen P, Ingall ED (1996) Redox stabilization of the atmosphere and oceans by phosphorus-limited marine productivity. Science 271:493 -496. source
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In the paper
“Assuming a sustained 25% increase in phosphorus weathering relative to bulk rock and C/P = 2,000 increases organic carbon burial by ∼ 90%, δ 13 C by 2.1 ‰ , and O2 by 0.67 PAL (Fig. 3, red).”
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. It may state two separate claims as one.
- Rests on an assumption: The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“which assuming 0.43 gC/g biomass and C/P = 2,000 (Table S3) suggests 0.26 μ mol P in biomass, or for C/P = 1,000 -4,000, 0.13 -0.52 μ mol P in biomass”
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In the paper
“Subsequent experiments (21) with the modern liverwort Marchantia paleacea found a 2.5- to 7-fold amplification of Ca weathering and a 9- to 13-fold amplification of P weathering from basalt.”
What the sources say
- Not checked yetcites [21] Quirk J, et al. (2015) Constraining the role of early land plants in Palaeozoic weathering and global cooling. Proc Biol Sci , 10.1098/rspb.2015.1115.
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In the paper
“Furthermore, Early Devonian coaly shales indicate extensive peatlands 410 -400 Ma and have C/N of 44 -119 (28), comparable to that in modern peatlands where N/P and C/P ratios tend to increase with depth to C/P > 3,000 (34).”
What the sources say
- Not checked yetcites [28] Kennedy KL, et al. (2013) Lower Devonian coaly shales of northern New Brunswick, Canada: Plant accumulations in the early stages of Terrestrial colonization. J Sediment Res 83(12):1202 -1215. source
- Not checked yetcites [34] Wang M, Moore TR, Talbot J, Richard PJH (2014) The cascade of C:N:P stoichiometry in an ombrotrophic peatland: from plants to peat. Environ Res Lett 9(2):024003. source
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In the paper
“The evolution of land plants is the obvious candidate”
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In the paper
“Here we show that neither scenario is correct.”
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In the paper
“However, plants and their associated mycorrhizal fungi can increase phosphorus weathering (20 -22), and this could be sustained on longer timescales if they preferentially weather phosphorus relative to alkalinity.”
What the sources say
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
- Not checked yetcites [21] Quirk J, et al. (2015) Constraining the role of early land plants in Palaeozoic weathering and global cooling. Proc Biol Sci , 10.1098/rspb.2015.1115.
- Not checked yetcites [22] Lenton TM, Watson AJ (2000b) Redfield revisited: 2. What regulates the oxygen content of the atmosphere? Global Biogeochem Cycles 14(1):249 -268. source
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It may state two separate claims as one.
In the paper
“The COPSE model we adapt here (10) predicts Early Paleozoic pO2 ∼ 0.2 -0.5 PAL consistent with redox proxy data but, like the other models (7, 8), it does not predict a rise in oxygen until the advent of forests starting ∼ 385 Ma, and continuing until ∼ 300 Ma, which is too late to explain marked changes in geochemical data that occur before ∼ 390 Ma (Fig. 1).”
What the sources say
- Not checked yetcites [10] source
plain- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“Given that earlier plants developed peatlands (28), and had rock-weathering capabilities (20, 21), they could also have affected the global carbon cycle (18, 20).”
What the sources say
- Not checked yetcites [28] Kennedy KL, et al. (2013) Lower Devonian coaly shales of northern New Brunswick, Canada: Plant accumulations in the early stages of Terrestrial colonization. J Sediment Res 83(12):1202 -1215. source
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
- Not checked yetcites [21] Quirk J, et al. (2015) Constraining the role of early land plants in Palaeozoic weathering and global cooling. Proc Biol Sci , 10.1098/rspb.2015.1115.
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“Though there are many subsequent δ 13 C fluctuations, including drops back to 0 ‰ , for example, at ∼ 400 Ma, the long-term mean δ 13 C remains ∼ 2 ‰ throughout the rest of the Paleozoic, the Mesozoic, and the Early Cenozoic (17), indicating a sustained increase in global organic carbon burial.”
What the sources say
- Not checked yetcites [17] Saltzman MR, Thomas E (2012) Carbon isotope stratigraphy. The Geologic Time Scale , eds Gradstein FM, Schmitz JGOD, Ogg GM (Elsevier, Boston), pp 207 -232. source
hedgeddirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Rests on an assumption: Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The persistent oxygenation of the ocean and appearance of charcoal can be explained by a rise in atmospheric oxygen occurring by ∼ 400 Ma”
plaindirect- Asserted: passes — The authors put it forward as true.
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The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Land plants typically have a much higher molar C/P ratio ( ∼ 1,000) than marine organic matter ( ∼ 100) due to carbon-rich but phosphorus-poor structural compounds such as sporopollenin, lignin, and, in their fungal mycorrhizal symbionts, chitin; therefore, they can support an increased organic carbon burial flux for the same P weathering flux.”
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Rests on an assumption: On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
None of the sources cited for this claim have been checked yet.
In the paper
“The P weathering flux is partly tied to bulk silicate weathering, for example, due to the dissolution of apatite inclusions in silicate rocks, and the silicate weathering flux of alkalinity is in turn set by negative feedback in the longterm carbon cycle, so is ultimately controlled by the degassing input of CO2 on timescales ≥ 1 My (7, 10).”
plaindirect- Asserted: passes — The authors put it forward as true.
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A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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.”
plaindirect- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“This ocean oxygenation is also supported by a Silurian increase in the C/S ratio of shales (15), and a shift in iron speciation data sometime during 435 -387 Ma (6).”
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“The early charcoal record (Table S1) was compiled from the literature (11, 12, 28, 44 -72) using existing compilations (12, 44 -47) and checking them where possible against the original sources.”
What the sources say
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“This finding is broadly consistent with the sulfur isotope ( δ 34 S) record (38 -40), which shows a marked decline through the Silurian -Early Devonian from ∼ 30 to ∼ 18 ‰ , although available data also suggest an earlier Late Ordovician -Early Silurian rise from ∼ 25 to ∼ 30 ‰ , which the present model does not capture.”
What the sources say
- Not checked yetcites [38] Prokoph A, Shields GA, Veizer J (2008) Compilation and time-series analysis of a marine carbonate δ 18O, δ 13C, 87Sr/86Sr and δ 34S database through Earth history. Earth Sci Rev 87(3 -4):113 -133. source
- Not checked yetcites [39] Gill BC, Lyons TW, Saltzman MR (2007) Parallel, high-resolution carbon and sulfur isotope records of the evolving Paleozoic marine sulfur reservoir. Palaeogeogr Palaeoclimatol Palaeoecol 256(3 -4):156 -173. source
- Not checked yetcites [40] Jones DS, Fike DA (2013) Dynamic sulfur and carbon cycling through the end-Ordovician extinction revealed by paired sulfate -pyrite δ 34S. Earth Planet Sci Lett 363: 144 -155. source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Sensitivity analyses ( SI Materials and Methods ) indicate that our results are robust.”
hedged- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Uncertainties shown in Fig. 1 represent 2 SD of the mean (analytical precision) plus the propagated uncertainty from matching in-house reference materials to the universal standard NIST SRM 3136 where seawater display δ 98/95 Mo = 2.3 ‰ (75, 76).”
What the sources say
- Not checked yetcites [75] Goldberg T, et al. (2013) Resolution of inter-laboratory discrepancies in Mo isotope data: An intercalibration. J Anal At Spectrom 28(5):724 -735. source
- Not checked yetcites [76] Nägler TF, et al. (2014) Proposal for an international molybdenum isotope measurement standard and data representation. Geostand Geoanal Res 38(2):149 -151. source
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The δ 13 C record suggests this increase in organic carbon burial was essentially permanent, producing a new dynamically stable state for atmospheric O2.”
hedgeddirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“new fire-mediated negative feedbacks on O2 were instigated that have played a key role in stabilizing atmospheric O2 concentration up to the present day (22, 43).”
What the sources say
- Not checked yetcites [22] Lenton TM, Watson AJ (2000b) Redfield revisited: 2. What regulates the oxygen content of the atmosphere? Global Biogeochem Cycles 14(1):249 -268. source
- Not checked yetcites [43] Kump LR (1988) Terrestrial feedback in atmospheric oxygen regulation by fire and phosphorus. Nature 335:152 -154. source
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The model is consistent with proxy reconstructions of seawater SO4 concentration, which suggest an Ordovician -Silurian rise from ∼ 6 to ∼ 10 mM (41), and with a Silurian increase in the molar C/S ratio of shales from ∼ 5 to ∼ 16 (15).”
What the sources say
- Not checked yetcites [41] Algeo TJ, Luo GM, Song HY, Lyons TW, Canfield DE (2015) Reconstruction of secular variation in seawater sulfate concentrations. Biogeosciences 12(7):2131 -2151. source
- Not checked yetcites [15] Berner RA, Raiswell R (1983) Burial of organic carbon and pyrite sulfur in sediments over phanerozoic time: A new theory. Geochim Cosmochim Acta 47(5):855 -862. source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“We explored the sensitivity of predicted global NPP to variations in atmospheric CO2 and corresponding climate state (Fig. S1), to constraining vegetation cover with extensive Late Ordovician ice sheet cover (Fig. S2), and to varying O2 in combination with CO2 (Table S2).”
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In this new steady state, oxidative weathering was increased (becoming less sensitive to variations in O2)”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
In the paper
“In our previously reported (20) weathering experiments with granite, the mean amounts of phosphate weathered into aqueous solution were as follows: control microcosms = 0.0137 μ mol P, biotic microcosms = 0.0726 μ mol P.”
What the sources say
- Not checked yetcites [20] Lenton TM, Crouch M, Johnson M, Pires N, Dolan L (2012) First plants cooled the Ordovician. Nat Geosci 5(2):86 -89. source
plain- Asserted: passes — The authors put it forward as true.
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- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“Then, in a sensitivity analysis, we considered uncertainty in geologic forcing (91), and alternative initial conditions (altering the feedback structure of the model).”
What the sources say
- Not checked yetcites [91] Royer DL, Donnadieu Y, Park J, Kowalczyk J, Goddéris Y (2014) Error analysis of CO2 and O2 estimates from the long-term geochemical model GEOCARBSULF. Am J Sci 314(9):1259 -1283. source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
The paper relies on this without support.
The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks.
“The major long-term source of oxygen to the atmosphere is the burial of organic carbon in sedimentary rocks (which represents the net flux of photosynthesis minus various pathways of respiration and oxidation).”
The paper relies on this without support.
Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks (δ13C).
“Increases in global organic carbon burial are recorded as positive shifts in the isotopic composition of carbonate rocks ( δ 13 C).”
The paper relies on this without support.
On long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the molar ratio of carbon-to-phosphorus in material that is buried.
“In simple terms, on long timescales, the global organic carbon burial flux is determined by the supply flux of the ultimate limiting nutrient phosphorus from weathering and the (molar) ratio of carbon-to-phosphorus in material that is buried”
The paper relies on this without support.
The C/P ratio of early land plants was similar to that of extant bryophytes.
“supported by the high C/P ratio of early land plant material”
“The C/P ratio of extant bryophytes (Table S3) was compiled from data in the literature (34, 80 -88).”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“Sedimentary recycling of organic carbon during the Archaean and Proterozoic can thus help reconcile increases in oxygen, and presumed associated increases in biological productivity, with the lack of a secular trend in δ13Ccarb until the Phanerozoic17 (where there is a shift from 0 to 2‰ associated with the rise of land plants13).”
From Atmospheric oxygen regulation at low Proterozoic levels by incomplete oxidative weathering of sedimentary organic carbon (2017) · cites this paper as [13]
In the citing paper
“The rise of oxygen to present levels simply required an increase in organic carbon burial rate, which may have begun in the Neoproterozoic Era10, but in our model was not completed until the mid-Palaeozoic Era with the colonization of the land by plants and fungi, liberating nutrient phosphorus from rocks and producing high C/P material for burial1113.”
From Atmospheric oxygen regulation at low Proterozoic levels by incomplete oxidative weathering of sedimentary organic carbon (2017) · cites this paper as [13]
Citations
Cites
The rise of oxygen in Earth's early ocean and atmosphere
Lyons, T. W., Reinhard, C. T., & Planavsky, N. J. (2014) · Nature · doi:10.1038/nature13068 · Cited by 2,810 · cited 2×
“Proc Natl Acad Sci USA 111(2):734–739.”
BackgroundGEOCARBSULF: A combined model for Phanerozoic atmospheric O2 and CO2
Berner, R. A. (2006) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2005.11.032 · Cited by 1,120
Burial of organic carbon and pyrite sulfur in sediments over phanerozoic time: a new theory
Berner, R. A., & Raiswell, R. (1983) · Geochimica et Cosmochimica Acta · doi:10.1016/0016-7037(83)90151-5 · Cited by 1,044
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“Ferruginous shales (77) are defined by the Fe speciation proxy when FeHR/FeT > 0.”
BackgroundContribution of cryptogamic covers to the global cycles of carbon and nitrogen
Elbert, W., Weber, B., Burrows, S., Steinkamp, J., Büdel, B., Andreae, M. O., & Pöschl, U. (2012) · Nature Geoscience · doi:10.1038/ngeo1486 · Cited by 957
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“Euxinic shales are defined (78, 79) by theMo enrichment proxy whenMo > 25 ppm (white circles in Fig.”
MethodThe Neoproterozoic oxygenation event: Environmental perturbations and biogeochemical cycling
Och, L. M., & Shields-Zhou, G. A. (2012) · Earth-Science Reviews · doi:10.1016/j.earscirev.2011.09.004 · Cited by 646
Statistical analysis of iron geochemical data suggests limited late Proterozoic oxygenation
Sperling, E. A., Wolock, C. J., Morgan, A. S., Gill, B. C., Kunzmann, M., Halverson, G. P., Macdonald, F. A., Knoll, A. H., & Johnston, D. T. (2015) · Nature · doi:10.1038/nature14589 · Cited by 637 · cited 3×
“This ocean oxygenation is also supported by a Silurian increase in the C/S ratio of shales (15), and a shift in iron speciation data sometime during 435–387 Ma (6).”
BackgroundCOPSE: A new model of biogeochemical cycling over Phanerozoic time
Bergman, N. M. (2004) · American Journal of Science · doi:10.2475/ajs.304.5.397 · Cited by 599
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Sun, R., Grasby, S. E., Shen, J., Xiao, J., & Yin, R. (2022) · Chemical Geology · doi:10.1016/j.chemgeo.2022.120830 · Cited by 9
Earth: Atmospheric Evolution of a Habitable Planet
Olson, S. L., Schwieterman, E. W., Reinhard, C. T., & Lyons, T. W. (2018) · Springer eBooks · doi:10.1007/978-3-319-55333-7_189 · Cited by 24
Morris, J. L., Edwards, D., & Richardson, J. B. (2018) · Elsevier eBooks · doi:10.1016/b978-0-12-813012-4.00004-8 · Cited by 8
黄, 建., 刘, 晓., 何, 永., 沈, 树., 侯, 增., 李, 曙., 李, 昶., 姚, 丽., & 黄, 季. (2021) · SCIENTIA SINICA Terrae · doi:10.1360/n072020-0399 · Cited by 8
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Herbivory and its effect on Phanerozoic oxygen concentrations
Laakso, T. A., Strauss, J. V., & Peterson, K. J. (2020) · Geology · doi:10.1130/g47085.1 · Cited by 5
The preservation of cause and effect in the rock record
D’Antonio, M. P., Ibarra, D. E., & Boyce, C. K. (2022) · Paleobiology · doi:10.1017/pab.2022.33 · Cited by 7
“…in time until reaching a suitable evolutionary origin or by identifying an evolutionary origin and scanning forward in time until reaching a suitable perturbation or trend (e.g., Berner 1992, 2006; Algeo et al. 2001; Heckman et al. 2001; Kennedy et al. 2006; Lenton et al. 2012, 2016; Kump 2014).”
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Zhang, J., Li, W., Fang, X. M., Wu, X., Li, C., & Zhang, Y. (2024) · Journal of the Geological Society · doi:10.1144/jgs2023-078 · Cited by 7
Atmospheric oxygenation as a potential trigger for climate cooling
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Learning to See ‘Green’ in an Ecological Crisis
Lawrence, N., & Calvo, P. (2023) · Sustainable development goals series · doi:10.1007/978-3-030-94391-2_9 · Cited by 9
Stratospheric dynamics modulates ozone layer response to molecular oxygen variations
Józefiak, I., Sukhodolov, T., Egorova, T. A., Chiodo, G., Peter, T., Rieder, H. E., Sedláčék, J., Stenke, A., & Rozanov, E. (2023) · Frontiers in Earth Science · doi:10.3389/feart.2023.1239325 · Cited by 6
Steinthorsdottir, M., Montañez, I. P., Royer, D. L., Mills, B. J. W., & Hönisch, B. (2024) · Elsevier eBooks · doi:10.1016/b978-0-323-99762-1.00074-7 · Cited by 12
Self-citationHorne, J. E., & Goldblatt, C. (2024) · Geochemistry Geophysics Geosystems · doi:10.1029/2023gc011252 · Cited by 8
Using deep learning to integrate paleoclimate and global biogeochemistry over the Phanerozoic Eon
Zheng, D., Merdith, A., Goddéris, Y., Yannick Donnadieu, A., Gurung, K., & Mills, B. J. W. (2024) · Geoscientific model development · doi:10.5194/gmd-17-5413-2024 · Cited by 10
Self-citationBharti, M. K., Siddique, R. A., Ranjan, K., Chandra, D., & Singh, N. P. (2021) · Springer eBooks · doi:10.1007/978-981-15-9758-9_3 · Cited by 6
Saintilan, N. J., Archer, C., Maden, C., Samankassou, E., Bernasconi, S. M., Szumigala, D. J., Mahaffey, Z., West, A., & Spangenberg, J. E. (2022) · Mineralium Deposita · doi:10.1007/s00126-022-01123-1 · Cited by 6
“1) in dolomitized carbonaceous limestone in a hot passive margin setting in the global context of ocean oxygenation and the rise of land plants in the Devonian (after Algeo and Scheckler 1998; Lenton et al. 2016; Wallace et al. 2017)”
BackgroundA Subsurface Stepping Stone Hypothesis for the Conquest of Land by Arthropods
Frumkin, A., & Chipman, A. D. (2023) · Diversity · doi:10.3390/d16010006 · Cited by 6
The origin of early-Paleozoic banded iron formations in NW China
Yang, X., Mao, J., Poulton, S. W., Dong, A., Liang, T., Wang, D., & Zhang, X. (2021) · Gondwana Research · doi:10.1016/j.gr.2021.02.008 · Cited by 6
Hydrocarbon resources resulting from plant terrestrialization: A review
Liu, Y., Feng, J., Wang, C., Mao, L., & Shen, Z. (2026) · Earth-Science Reviews · doi:10.1016/j.earscirev.2026.105457 · Cited by 5
Evolution of terrestrial herbivory: nutrient stoichiometry, body size, and dietary diversity
Boyce, C. K. (2023) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2023.1304831 · Cited by 5
What does not kill Gaia makes her stronger: impacts of external perturbations on biosphere evolution
Arthur, R., Nicholson, A., & Mayne, N. (2024) · Monthly notices of the Royal Astronomical Society · doi:10.1093/mnras/stae2003 · Cited by 4
Mid-Devonian ocean oxygenation enabled the expansion of animals into deeper-water habitats
Bubphamanee, K., Kipp, M. A., Meixnerová, J., Stüeken, E. E., Ivany, L. C., Bartholomew, A. J., Algeo, T. J., Brocks, J. J., Dahl, T. W., Kinsley, J., et al. (2025) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2501342122 · Cited by 6
“Together, terrestrial and marine productivity would cause a global increase in organic carbon burial and thus leave surplus O 2 in the atmosphere ( 57 , 63 , 64 ).”
Self-citationUnmiraculous facultative anaerobes (comment on DOI 10.1002/bies.201600174)
Martin, W. F. (2017) · BioEssays · doi:10.1002/bies.201700041 · Cited by 6
“Newer data even have it that the high levels of atmospheric oxygen arose only a few hundred million years ago [21], with very recent findings indicating that modern oxygen levels were attained following the origin of land plants, only some 400 million years ago [22].”
BackgroundHe, R., Elrick, M., Day, J. E., Lu, W., & Lu, Z. (2022) · Frontiers in Marine Science · doi:10.3389/fmars.2022.874759 · Cited by 5
“Both proxy (Mo isotope, U isotope, I/Ca ratio, Ce anomaly and charcoal abundance) and model (COPSE and GEOCARBSULF) results suggest atmospheric pO2 levels increased during the Devonian (Dahl et al., 2010; Glasspool and Scott, 2010; Lenton et al., 2016; Wallace et al., 2017; Krause et al., 2018; Lu et al., 2018; Elrick…”
BackgroundLu, X., Gilleaudeau, G. J., & Kendall, B. (2023) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2023.10.034 · Cited by 5
Mechanistic view of plant adaptation under iron deficiency stress
Saha, S., & Singh, S. (2024) · Elsevier eBooks · doi:10.1016/b978-0-443-18927-2.00022-4 · Cited by 4
The Early Earth as an Analogue for Exoplanetary Biogeochemistry
Stüeken, E. E., Olson, S. L., Moore, E. K., & Foley, B. J. (2024) · Reviews in Mineralogy and Geochemistry · doi:10.2138/rmg.2024.90.14 · Cited by 5
Don’t mind the “charcoal gap”: A reassessment of Devonian wildfire
Glasspool, I. J., & Gastaldo, R. A. (2024) · Geology · doi:10.1130/g52648.1 · Cited by 5
Analysis of gases in fossil amber
Landis, G. P., & Berner, R. A. (2018) · American Journal of Science · doi:10.2475/05.2018.06 · Cited by 5
Hess, A. V., & Trop, J. M. (2019) · Palaios · doi:10.2110/palo.2019.020 · Cited by 5
“…Formation; Mbr.¼Member, FWWB¼ fair-weather wave base; SWB¼ storm wave base; VA¼Virginia; WV¼West Virginia; MD¼Maryland; PA¼ Pennsylvania; NY ¼ New York. surface-water oxygen levels were rising (Lenton et al. 2016; Lu et al. 2018), there is evidence for oxygen limitation in the Appalachian Basin.”
BackgroundEarth: Atmospheric Evolution of a Habitable Planet
Olson, S. L., Schwieterman, E. W., Reinhard, C. T., & Lyons, T. W. (2018) · Springer eBooks · doi:10.1007/978-3-319-30648-3_189-1 · Cited by 5
Khan, K., & Aken, O. V. (2022) · Journal of Experimental Botany · doi:10.1093/jxb/erac351 · Cited by 5
“…levels were greater than in the original aquatic environment, and the explosion of land plant growth further promoted the rise of oxygen, which reached present-day levels by 400 MYA (Scott and Glasspool, 2006; Alboresi et al., 2010; Lyons et al., 2014; Lenton et al., 2016; Dahl and Arens, 2020).”
BackgroundSilurian freshwater arthropod from northwest China
Zong, R., Edgecombe, G. D., Liu, B., Wang, Y., Yin, J., Ma, J., & Xu, H. (2023) · Papers in Palaeontology · doi:10.1002/spp2.1488 · Cited by 5
“The diversification and expansion of vascular plants not only resulted in an increase in oxygen concentration, but also provided an adequate food source and a new ecological niche for arthropods after they left the sea (Labandeira 2013; Lenton et al. 2016).”
BackgroundGrosjean, E., Jarrett, A. J. M., Boreham, C. J., Wang, L., Johnson, L. M., Hope, J. M., Ranasinghe, P., Brocks, J. J., Bailey, A. H. E., Butcher, G. A., et al. (2023) · Organic Geochemistry · doi:10.1016/j.orggeochem.2023.104688 · Cited by 5
Quinton, P. C., Rygel, M. C., & Heins, M. K. (2021) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2021.110440 · Cited by 6
Kriegshauser, L., Knosp, S., Grienenberger, E., et al. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.09.16.300285 · Cited by 5
“Through photosynthetic activity and rock weathering, early land plants contributed to the rise of atmospheric oxygen, carbon sequestration and the development of soils (Lenton et al., 2016; Porada et al., 2016; Retallack, 1997).”
BackgroundA Timeline of Bacterial and Archaeal Diversification in the OceanPreprint
Martínez-Gutiérrez, C. A., Uyeda, J. C., & Aylward, F. O. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.10.27.514092 · Cited by 5
“…ocean was the result of an increment of the burial of organic carbon in sedimentary rocks due to the diversification of the earliest land plants 25,57,59 , which has been also associated with increased phosphorus weathering rates 57,60 , global impacts on the global element cycles 61 , and an…”
BackgroundConvergent evolution and adaptive diversification of root symbioses
Liu, Z., Hu, B., Flemetakis, E., Hänsch, R., Franken, P., & Rennenberg, H. (2025) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/brv.70077 · Cited by 5
“This adaptation was facilitated not only by innovations in plant morphology (e.g. cuticles, stomata, and vascular tissues) (Pires & Dolan, 2012; Lenton et al ., 2016; Kenrick & Crane, 1997; Julca et al ., 2021), but also through The origin and evolution of plant root symbioses symbiotic…”
A Global Conservation Blind Spot: Neglect of Bryophytes Undermines Biodiversity Targets
Yin, X., Liu, Y., Kong, F., Shu, L., PENG, C., & Ang, L. P. (2026) · Global Change Biology · doi:10.1111/gcb.70789 · Cited by 4
“…Ordovician period (Harris et al. 2022), alongside early fungi and bacteria (Beraldi-Campesi 2013; Heckman et al. 2001; Sánchez-Baracaldo et al. 2022) and played a role in raising atmospheric oxygen to near-modern levels through enhanced carbon burial and phosphorus weathering (Lenton et al. 2016).”
Ma, D.-C., Li, W., Chen, Z., Fang, X. M., Cheng, J.-F., & Jia, X.-L. (2023) · Palaeoworld · doi:10.1016/j.palwor.2023.06.006 · Cited by 4
Nakajima, H. (2025) · Functional Plant Biology · doi:10.1071/fp24335 · Cited by 4
Hu, D., Sun, L., Xu, Y., Zhang, X., Zhang, X., & Shen, Y. (2025) · Chemical Geology · doi:10.1016/j.chemgeo.2025.122890 · Cited by 5
Extreme environmental tolerance and space survivability of the moss, Physcomitrium patens
Maeng, C.-H., Hiwatashi, Y., Nakamura, K., Matsuda, O., Mita, H., Tomita‐Yokotani, K., Yokobori, S., Yamagishi, A., Kume, A., & Fujita, T. (2025) · iScience · doi:10.1016/j.isci.2025.113827 · Cited by 6
“4 Since then, bryophytes have survived without becoming extinct despite changes in the Earth’s environment (e.g., mass extinction in Earth’s history).”
The Fractional MacroEvolution Model: a simple quantitative scaling macroevolution model
Lovejoy, S. M., & Spiridonov, A. (2024) · Paleobiology · doi:10.1017/pab.2023.38 · Cited by 4
“In addition, presumably the appearance, global spread, and deep impact on the climate of terrestrial plants in the Silurian is highly significant (Lenton et al. 2016).”
BackgroundA novel carbon cycle turbulence index identifies environmental and ecological perturbations
Li, Z.-H., Guo, Z., Chen, Z., Poulton, S. W., Bao, Y., Zhao, L., & Zhang, F.-F. (2021) · Geochemical Perspectives Letters · doi:10.7185/geochemlet.2137 · Cited by 4
Hidden functional complexity in the flora of an early land ecosystem
D'Ario, M., Lane, B., Junod, M. F., Leslie, A. B., Mosca, G., & Smith, R. S. (2023) · New Phytologist · doi:10.1111/nph.19228 · Cited by 5
“…(Kenrick et al ., 2012) and likely altering global sedimentation patterns, biogeochemical cycling and perhaps oxygenation levels (Algeo & Scheckler, 1998; Gibling & Davies, 2012; Morris et al ., 2015; Lenton et al ., 2016; Wallace et al ., 2017; McMahon & Davies, 2018; Davies et al ., 2021).”
BackgroundHu, J., Wang, M., Wang, H., Li, S., Santosh, M., Zhao, Y., Li, D., Yan, Q.-H., Dong, R., & Ren, H. (2022) · Ore Geology Reviews · doi:10.1016/j.oregeorev.2022.105165 · Cited by 4
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“Notwithstanding the circumstance that the majority of eukaryote evolution occurred in oxygen poor environments (Martin and Mentel, 2008; Müller et al., 2012; Lyons et al., 2014; Lenton et al. 2016) (Fig.”
BackgroundInfluentialUsing Deep Learning to integrate paleoclimate and global biogeochemistry over Phanerozoic timePreprint
Zheng, D., Merdith, A., Goddéris, Y., et al. (2024) · doi:10.5194/gmd-2023-230 · Cited by 4
Self-citationZhang, X.-L., Wang, Y., & Wang, Y. (2024) · Palaeoworld · doi:10.1016/j.palwor.2024.02.004 · Cited by 3
Stolper, D. A., Pu, X., Lloyd, M. K., Christensen, N. I., Bucholz, C. E., & Lange, R. A. (2022) · Geochemistry Geophysics Geosystems · doi:10.1029/2021gc010196 · Cited by 4
“Why such a delay may have occurred is not known, but various ideas on what caused the final increase in atmospheric O2 levels to values that resulted in the oxygenation of the deep ocean include the rise of land plants (Lenton et al., 2016; Wallace et al., 2017), changes in internal feedbacks in global biogeochemical…”
BackgroundLi, J., & Azmy, K. (2023) · Marine and Petroleum Geology · doi:10.1016/j.marpetgeo.2023.106408 · Cited by 3
Probing the limits of habitability: a catalogue of rocky exoplanets in the habitable zone
Bohl, A., Lawrence, L., Lowry, G., & Kaltenegger, L. (2026) · Monthly Notices of the Royal Astronomical Society · doi:10.1093/mnras/stag028 · Cited by 12
“Land plants conquered the land masses at about 0.5Gyr ago (about 4Gyr after Earth’s formation)(Lenton et al. 2016); dinosaurs roamed between about 250 to 66 million years ago; and humans evolved only recently on this large timescale, with the oldest Homo sapiens fossil claim about 300,000 years ago…”
Carbon and sulfur cycling across the Silurian-Devonian boundary in the Qujing Basin, South China
Sun, L., Zhang, X., Zhu, M., Xu, Y., Zhao, W., & Shen, Y. (2024) · Chemical Geology · doi:10.1016/j.chemgeo.2024.121952 · Cited by 2
McMahon, S., Loron, C., Cooper, L. M., et al. (2023) · Geological Magazine · doi:10.1017/s0016756824000049 · Cited by 3
“…the balance of Earth ’ s biomass from prokaryotic to eukaryotic dominance and permanently altered the planet ’ s atmosphere, soil and climate (Lenton et al. 2016; McMahon & Parnell, 2018) Previous studies have identified more than a dozen distinct cyanobacterial morphologies in the Rhynie…”
BackgroundSedimentary 17O‐Nitrate Evidence for Phanerozoic Aridity and Humidity Oscillations in South China
Fan, W., Liu, X., Zhou, M., et al. (2024) · Geophysical Research Letters · doi:10.1029/2024gl111475 · Cited by 3
“Sea‐level changes, for instance, have interrupted terrestrial climatic records (Scotese et al., 2021), and while charcoal records indicate humid climates as early as the late Silurian (Lenton et al., 2016; Scotese et al., 2021), other regions, such as the South China Block, only begin in the middle Devonian (Scotese e…”
InfluentialRinkevičiūtė, S., Daumantas, L., Radzevičius, S., et al. (2025) · Lethaia · doi:10.18261/let.58.1.2 · Cited by 2
Shen, J., Ding, K., Yu, Z., Zhang, Y., Ni, J., & Wu, Y. (2025) · Plants · doi:10.3390/plants14081178 · Cited by 2
Oxygen reductase origin followed the great oxidation event and terminated the Lomagundi excursion
Trost, K., Gennis, R. B., Allen, J. F., Mills, D. B., & Martin, W. F. (2025) · Biochimica et Biophysica Acta (BBA) - Bioenergetics · doi:10.1016/j.bbabio.2025.149575 · Cited by 2
“Following the LJE, δ 13 C values fall to levels indicating roughly 1–10% of present atmospheric O 2 levels (PAL) for almost 2 billion years until the appearance of land plants [49–51].”
InfluentialMitchell, C. E., Sheets, H. D., Melchin, M. J., & Holmden, C. (2025) · Cambridge University Press eBooks · doi:10.1017/9781009684064 · Cited by 2
Carbon/phosphorus burial ratio reveals a rapid spread of land plants during the Late Ordovician
Cai, J., Tarhan, L. G., Lenton, T. M., Qiu, R., Peacock, C. L., Planavsky, N. J., Ju, P., Zhao, W., Xu, Z., Zhang, H., et al. (2026) · Nature Ecology & Evolution · doi:10.1038/s41559-026-02995-6 · Cited by 2
Self-citationHu, M., Myrow, P. M., Fike, D. A., Pasquo, M. D., Zatoń, M., Fischer, W. W., & Coates, M. I. (2024) · Geological Society of America Bulletin · doi:10.1130/b36728.1 · Cited by 2
Garza, H. K., Catlos, E. J., Brookfield, M. E., Malkowski, M. A., Chamberlain, K. R., Loewy, S., & Stockli, D. F. (2024) · Geological Magazine · doi:10.1017/s0016756824000384 · Cited by 2
“…on the evolution of other organisms and ecosystems, establishing themselves as a critical component of the planet ’ s biodiversity and ecological complexity Garwood & Edgecombe, 2011; Kenrick et al. , 2012; Dunlop et al. , 2013; Lenton et al. , 2016; Wallace et al. , 2017; Dahl &Arens, 2020).”
Future directions for understanding the coevolution of life and oxygen
Alcott, L., Bowyer, F. T., & Agić, H. (2025) · Communications Earth & Environment · doi:10.1038/s43247-025-02689-0 · Cited by 2
“The third stepwise event, the Paleozoic Oxygenation Event (POE; ~450 – 350 Ma 20,21 ), is associated with the rise of the terrestrial biosphere during the mid-Paleozoic 22,23 , and culminated in the oxygenation of the deep ocean and an oxygenated world more reminiscent of the modern day.”
A Timeline of Bacterial and Archaeal Diversification in the OceanPreprint
Martínez-Gutiérrez, C. A., Uyeda, J. C., & Aylward, F. O. (2023) · eLife · doi:10.7554/elife.88268.2 · Cited by 2
Advancing studies on global biocrusts distributionPreprint
Wang, S., Ma, L., Yang, L., et al. (2023) · doi:10.5194/egusphere-2023-2131 · Cited by 2
Environmental effects of land plants evolution in the phanerozoic
Cai, J., Ju, P., Liu, M., Xue, J., & Zhao, M. (2025) · Chinese Science Bulletin (Chinese Version) · doi:10.1360/tb-2025-0173 · Cited by 1
Otoo, B. K. A. (2025) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2025.1606225 · Cited by 2
Astronomical calibration of the Hirnantian Isotopic Carbon Excursion
Ouyang, J., Fang, Q., Wu, H., Chu, R., Shi, M., Gai, C., Zhong, Y., Yang, T., & Li, H. (2025) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2025.113498 · Cited by 1
Liu, C., Qi, J., Cózar, P., Coronado, I., Li, F., Liu, W., Li, M., Wang, P., Li, X., Hu, X., et al. (2025) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2025.113528 · Cited by 1
Holland, F. S. M., Stüeken, E. E., Dan, W., Zhang, W., Zhu, Y., & Mikhail, S. (2026) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2026.119890 · Cited by 3
Cooling-induced intensification of ocean anoxia in the mid-Paleozoic
Wang, Y., Wignall, P. B., Mills, B. J. W., Dickson, A. J., Loydell, D. K., Xiong, Y., Xu, Z., Peakall, J., & Poulton, S. W. (2026) · Science Advances · doi:10.1126/sciadv.aec8573 · Cited by 1
“…OAEs are also distinct from their Mesozoic counterparts in that they occurred against a backdrop of generally low atmospheric and oceanic oxygenation levels, with near-modern levels of oxygenation only developing during the Late Silurian to Devonian (the Paleozoic Oxygenation Event) ( 10 , 11 ).”
Self-citationTrujillo‐Esquivel, E., Lorenzo-Manzanarez, J. L., Espinal‐Centeno, A., Méndez-Álvarez, D., Aguilar‐Cruz, A., Brieba, L. G., & Cruz‐Ramírez, A. (2026) · The Plant Cell · doi:10.1093/plcell/koag089 · Cited by 1
Earth system instability explains redox paradox during late Cambrian SPICE event
Sheng, K., Li, Z., Zhou, J., Lin, Y.-B., Xiong, G., Ren, G., Chen, Z., Lenton, T. M., Shen, S., & Zhang, F. (2026) · Science Advances · doi:10.1126/sciadv.aeb9224 · Cited by 2
“One hypothesis posits delayed oxygenation [ ~ 0.2 present atmospheric level (PAL)] until the middle Paleozoic with the expansion of land plants ultimately driving the rise to modern levels, termed the Paleozoic oxygenation event ( 12 ).”
InfluentialSelf-citationPhylogenomic frameworks from deep archaeplastid evolution to embryophyte diversification.
Bastürk, M. N., Kunz, C. F., Darienko, T., Pröschold, T., de Vries, J., & Chen, H.-H. (2026) · Current Biology · doi:10.1016/j.cub.2026.03.019 · Cited by 5
The contribution and responses of mosses to the global carbon cycle
Mohanasundaram, B., & Pandey, S. (2026) · Trends in Plant Science · doi:10.1016/j.tplants.2026.06.012 · Cited by 1
Willow communities, optimal absorption of carbon dioxide from the atmosphere
Gorobets, A., & Silva, M. P. (2020) · IOP Conference Series Earth and Environmental Science · doi:10.1088/1755-1315/595/1/012038 · Cited by 1
(2022) · Nature Geoscience · doi:10.1038/s41561-022-01030-0 · Cited by 1
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Ocean redox evolution past and present
Hardisty, D. S., & Lau, K. (2024) · Elsevier eBooks · doi:10.1016/b978-0-323-99762-1.00033-4 · Cited by 3
Загоскина, Н. В. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms252212339 · Cited by 1
“Only under certain conditions can they exist in conditions of its deficiency or even complete absence [37–40].”
CHUPRYNA, A., Aubineau, J., Bankole, O. M., Fru, E. C., Konhauser, K. O., Nesterovskyi, V. A., Martyshyn, A., Soldatenko, Y., Becker-Kerber, B., Sansjofre, P., et al. (2025) · Comptes Rendus Géoscience · doi:10.5802/crgeos.296 · Cited by 2
Nakajima, H., Okazawa, A., Kubuki, S., Shen, Q., & Itoh, K. (2019) · BioMetals · doi:10.1007/s10534-019-00169-5 · Cited by 1
Walking the forest imaginary: A breath between us
Santos, J. P. (2021) · International Journal of Education through Art · doi:10.1386/eta_00049_3 · Cited by 1
Effects of variability of meteorological measures on soil temperature in permafrost regionsPreprint
Beer, C., Porada, P., Ekici, A., & Brakebusch, M. (2016) · doi:10.5194/tc-2016-210 · Cited by 1
“Furthermore, it is applicable to estimate various impacts of lichens and bryophytes on biogeochemical cycles (Porada et al., 2016b; Lenton et al., 2016).”
BackgroundSelf-citationNitrogenase inhibition limited oxygenation of the Proterozoic atmospherePreprint
Allen, J. F., Thake, B., & Martin, W. F. (2018) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/475236 · Cited by 1
Re-evaluating Reality in the Age of VR: Toward an Embodied VR
Aghilidehkordi, B. (2020) · ERA: Education and Research Archive (University of Alberta) · doi:10.7939/r3-fbsq-tg17 · Cited by 1
(2022) · doi:10.5194/gmd-2022-12 · Cited by 1
Self-citationPietluch, F., Mackiewicz, P., Sidorczuk, K., & Gagat, P. (2022) · bioRxiv · doi:10.1101/2022.07.01.497312 · Cited by 1
“Archaeplastidans have participated in carbon dioxide fixation, oxygen production and other global biogeochemical cycles for hundreds of millions of years, but they also donated their plastids to many protist groups thereby further spreading photosynthesis to multiple branches of the tree of life (4, 9).”
BackgroundContemporary atmospheric oxygen levels maximize global protection by ozonePreprint
Józefiak, I., Sukhodolov, T., Egorova, T. A., Chiodo, G., Stenke, A., Rieder, H. E., Peter, T., & Rozanov, E. (2022) · Research Square · doi:10.21203/rs.3.rs-2301615/v1 · Cited by 1
Modifying the Anthropocene Equation with One Health Concept
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A Historical Perspective on Long‐Term Global Biogeochemical Box Models
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“In the early stages of biosphere terrestrialization, land plants changed the Earth System by increasing atmospheric oxygen levels [79] and may have contributed to global cooling in the Late Ordovician by removing CO 2 from the atmosphere [80].”
THE CLIMATIC SIGNIFICANCE OF LATE ORDOVICIAN–EARLY SILURIAN BLACK SHALES
Pohl, A., Yannick Donnadieu, A., Yannick Donnadieu, A., Hir, G. L., Hir, G. L., Ferreira, D., & Ferreira, D. (2019) · Abstracts with programs - Geological Society of America · doi:10.1130/abs/2019am-336415
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The origin and evolution of vascular tissue regulators TMO5 and LHW
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Hoysted, G. A., Kowal, J., Pressel, S., et al. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.09.29.318022
“…evidence that the global environment underwent major changes throughout the Palaeozoic Era (541-250 Ma) (Berner et al., 2006; Bergman et al., 2004; Lenton et al., 2016), consisting of a step-wise increase of the Earth’s atmospheric oxygen ([O2]), and a simultaneous decline in atmospheric [CO2].”
BackgroundPlate motion drives variability in ocean oxygenation through the PhanerozoicPreprint
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Metadata
- Authors
- Lenton, T. M., Dahl, T. W., Daines, S. J., Mills, B. J. W., Ozaki, K., Saltzman, M. R., & Porada, P.
- Affiliations
- University of Exeter (GB)
- University of Copenhagen (DK)
- University of Leeds (GB)
- Georgia Institute of Technology (US)
- The Ohio State University (US)
- Stockholm University (SE)
- ORCID
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2016-08-15
- Volume
- 113
- Issue
- 35
- Pages
- 9704-9709
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC5024600
- OpenAlex
- W2508469640
- Cited by
- 343
- References
- 92
- Citations by year
- 2026: 25 · 2025: 33 · 2024: 45 · 2023: 48 · 2022: 47 · 2021: 34 · 2020: 21 · 2019: 32 · 2018: 31 · 2017: 23 · 2016: 4
- FWCI
- 24.94
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Research at the University of Copenhagen (University of Copenhagen) (publishedVersion)
- White Rose Research Online (University of Leeds, The University of Sheffield, University of York) (acceptedVersion)
- Open Research Exeter (University of Exeter) (acceptedVersion)
- Figshare (submittedVersion)
- Tokyo Tech Research Repository (Tokyo Institute of Technology) (submittedVersion)
- Funders
- Leverhulme Trust (RPG-2013-106)
- Natural Environment Research Council (NE/I005978/2)
- Royal Society (Wolfson Research Merit Award)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Stratigraphy of Fossils
- Physical Sciences › Environmental Science › Global and Planetary Change › Earth Systems and Cosmic Evolution
- Physical Sciences › Earth and Planetary Sciences › Geochemistry and Petrology › Geological and Geochemical Analysis
- Keywords
- atmospheric oxygenation, early land plants, organic carbon burial, bryophytes, terrestrial net primary productivity, C:P ratio, carbonate carbon isotopes, biogeochemical cycles, fire feedbacks, redox proxies
- MeSH terms
- Origin of Life, Atmosphere, Carbon Dioxide, Carbon Isotopes, Biological Evolution, Oceans and Seas, Oxidation-Reduction, Oxygen, Phosphorus, Photosynthesis, Soil, Models, Statistical, Biomass, Earth, Planet, Geologic Sediments, Embryophyta, History, Ancient
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