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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
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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
“Data from modern bryophytes suggests this plentiful early plant material had a much higher molar C:P ratio ( ∼ 2,000) than marine biomass ( ∼ 100)”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn 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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesThe 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
“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”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- 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.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- 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.”
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn 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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA 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
“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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- 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
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesA 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
“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)”
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA 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 presence of these rock-weathering capabilities in two early diverging lineages (mosses and liverworts) suggests it is an ancestral trait.”
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- It may state two separate claims as one. 33
- Probably not a substantial claim. 14
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
“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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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 mean moss biomass in the biotic microcosms was 14.390 mg”
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
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
“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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably 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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
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
“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).”
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
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
“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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably 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”
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
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
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
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
“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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably 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”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim. It may state two separate claims as one.
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.
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
“The evolution of land plants is the obvious candidate”
Tests✓ Asserted: passes✗ Checkable: fails✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim. It may be too vague to check against evidence.
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
“Here we show that neither scenario is correct.”
Load-bearingStrength:boosted· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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.
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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).”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
“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.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
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
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably 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
Strength:hedged· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt 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.”
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably 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
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably 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.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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).”
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably 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)”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt 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
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably 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
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably 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
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“Proc Natl Acad Sci USA 111(2):734–739.”
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Berner, R. A. (2006) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2005.11.032 · Cited by 1,120
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A New Model for Atmospheric Oxygen Over Phanerozoic Time
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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
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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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2 of the 92 references couldn’t be matched to a known paper.
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Cited by
Sorted by citations
Items 101–200 of 353
Palaeozoic And Mesozoic Palaeo-Wildfires: An Overview On Advances In The 21st Century
Jasper, A., Pozzebon–Silva, Â., Carniere, J. S., & Uhl, D. (2021) · The Palaeobotanist · doi:10.54991/jop.2021.13 · Cited by 29
Spaak, G., Edwards, D. S., Foster, C. B., Pagès, A., Summons, R. E., Sherwood, N., & Grice, K. (2017) · Global and Planetary Change · doi:10.1016/j.gloplacha.2017.10.010 · Cited by 29
“Recent work by Lenton et al. (2012, 2016) suggests that the appearance of early land plants could have had a significant impact on global climate, potentially contributing to the Late Ordovician cooling and the rise in atmospheric oxygen.”
BackgroundZhang, J., Li, C., Zhong, Y., Wu, X., Fang, X. M., Liu, M., Chen, D., Gill, B. C., Algeo, T. J., Lyons, T. W., et al. (2024) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2024.118635 · Cited by 28
Climate windows of opportunity for plant expansion during the Phanerozoic
Gurung, K., Field, K. J., Batterman, S. A., Goddéris, Y., Yannick Donnadieu, A., Porada, P., Taylor, L. L., & Mills, B. J. W. (2022) · Nature Communications · doi:10.1038/s41467-022-32077-7 · Cited by 28
“…composition and biogeochemical cycles, it has been hypothesised that plants had a key role in driving both the Hirnantian (~445Ma) and Late Paleozoic (~300Ma) ice ages 10,11 as well as mid-Paleozoic oxygenation of the atmosphere 12 and the more recent Cenozoic cooling 13,14 (66Ma – present).”
BackgroundSelf-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
Nutrient acquisition by symbiotic fungi governs Palaeozoic climate transition
Mills, B. J. W., Batterman, S. A., & Field, K. J. (2017) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2016.0503 · Cited by 28
“We examined the implications of contrasting plant–fungal relationships (figure 3) in response to changes in CO2 by incorporating phosphorus effects on global plant biomass and the ability of fungi to affect plant phosphorus uptake into the COPSE biogeochemical model (Carbon Oxygen Phosphorus Sulphur Evolution) [2,3].”
BackgroundMethodInfluentialSelf-citationTranscriptomic and Metabolomic Response to High Light in the Charophyte Alga Klebsormidium nitens
Serrano-Pérez, E., Romero–Losada, A. B., Morales-Pineda, M., García‐Gómez, M. E., Couso, I., García‐González, M., & Romero–Campero, F. J. (2022) · Frontiers in Plant Science · doi:10.3389/fpls.2022.855243 · Cited by 23
Zhang, J., Li, C., Fang, X. M., Li, W., Deng, Y., Tu, C., Algeo, T. J., Lyons, T. W., & Zhang, Y. (2022) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2022.117858 · Cited by 27
Pohl, A., Harper, D. A. T., Yannick Donnadieu, A., Hir, G. L., Nardin, É., & Servais, T. (2017) · Lethaia · doi:10.1111/let.12247 · Cited by 27
Graham, L., Graham, J. M., Knack, J. J., Trest, M. T., Piotrowski, M., & Arancibia-Ávila, P. (2017) · INTERNATIONAL JOURNAL PLANT SCIENCES · doi:10.1086/693019 · Cited by 18
“Models suggest that early moss-like plants might have strongly influenced early Paleozoic atmospheric chemistry and climate (Graham et al. 2004; Lenton et al. 2012, 2016; Porada et al. 2016).”
BackgroundKoehler, M. C., Stüeken, E. E., Hillier, S., & Prave, A. R. (2019) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2019.109321 · Cited by 26
White, J. D., Montañez, I. P., Wilson, J. P., Poulsen, C. J., McElwain, J. C., DiMichele, W. A., Hren, M. T., Macarewich, S. I., Richey, J. D., & Matthaeus, W. J. (2020) · American Journal of Science · doi:10.2475/09.2020.01 · Cited by 25
Evolution and Classification of Ferns and Lycophytes
Sessa, E. B. (2018) · Springer eBooks · doi:10.1007/978-3-319-75103-0_9 · Cited by 24
Harnessing hypoxia as an evolutionary driver of complex multicellularity
Hammarlund, E. U. (2020) · Interface Focus · doi:10.1098/rsfs.2019.0101 · Cited by 25
Ward, J. F., Verdel, C., Campbell, M. J., Leonard, N. D., & Nguyen, A. D. (2019) · Precambrian Research · doi:10.1016/j.precamres.2019.105471 · Cited by 24
Yin, J., Li, H., & Xiao, K. (2023) · Minerals · doi:10.3390/min13040547 · Cited by 26
Gould, S. B., Garg, S. G., Handrich, M. R., Sathi, S. N., Gruenheit, N., Tielens, A. G. M., & Martin, W. F. (2019) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2019.1491 · Cited by 22
“The rise of oxygen concentration to modern levels coincides with the conquering of land by streptophyte algae some 500 Ma and the beginning of massive carbon burial on land [6,9,10].”
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Biodiversity and Global Change
Lenton, T. M. (2019) · Cambridge University Press eBooks · doi:10.1017/9781108668675.005 · Cited by 23
Self-citationClimate change shapes the future evolution of plant metabolism
Xu, S. Y., & Weng, J. (2020) · Advanced Genetics · doi:10.1002/ggn2.10022 · Cited by 20
Legitimacy and limitations of valuing the oxygen production of ecosystems
Chen, H., Costanza, R., & Ida Kubiszewski, A. (2022) · Ecosystem Services · doi:10.1016/j.ecoser.2022.101485 · Cited by 21
The expansion of land plants during the Late Devonian contributed to the marine mass extinction
Smart, M. S., Filippelli, G. M., Gilhooly, W., Ozaki, K., Reinhard, C. T., Marshall, J. E. A., & Whiteside, J. H. (2023) · Communications Earth & Environment · doi:10.1038/s43247-023-01087-8 · Cited by 24
Self-citationSpiridonov, A. (2017) · The Journal of Geology · doi:10.1086/691184 · Cited by 24
“The Silurian period was an environmentally and macroevolutionary volatile time interval (Cooper et al. 2014; Crampton et al. 2016), punctuated by many oceanic biogeochemical events (Jeppsson 1987, 1998; Saltzman 2005; Lenton et al. 2016).”
BackgroundEnabling the water-to-land transition
Ralf Reski, A. (2018) · Nature Plants · doi:10.1038/s41477-018-0101-5 · Cited by 20
Evolutionary Adaptations of Parasitic Flatworms to Different Oxygen Tensions
Martínez-González, J. D. J., Guevara‐Flores, A., & Mena, I. P. D. A. (2022) · Antioxidants · doi:10.3390/antiox11061102 · Cited by 17
“Currently, it is accepted that this change began with the origin of cyanobacteria and the development of oxygenic photosynthesis [1,2] and continued later with the emergence and diversification of photosynthetic pigments in different types of algae and plants [3,4].”
BackgroundA timeline of bacterial and archaeal diversification in the ocean
Martínez-Gutiérrez, C. A., Uyeda, J. C., & Aylward, F. O. (2023) · eLife · doi:10.7554/elife.88268 · Cited by 22
Paleo-redox context of the Mid-Devonian Appalachian Basin and its relevance to biocrises
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Beer, C., Porada, P., Ekici, A., & Brakebusch, M. (2018) · The Cryosphere · doi:10.5194/tc-12-741-2018 · Cited by 20
“Furthermore, it is applicable when estimating various impacts of lichens and bryophytes on biogeochemical cycles (Lenton et al., 2016; Porada et al., 2016b, 2017).”
BackgroundSelf-citationHammarlund, E. U., Smith, M. P., Rasmussen, J. A., Nielsen, A. T., Canfield, D. E., & Harper, D. A. T. (2018) · Geobiology · doi:10.1111/gbi.12315 · Cited by 21
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Three enzymes governed the rise of O2 on Earth
Mrnjavac, N., Degli Esposti, M., Mizrahi, I., Martin, W. F., & Allen, J. F. (2024) · Biochimica et Biophysica Acta (BBA) - Bioenergetics · doi:10.1016/j.bbabio.2024.149495 · Cited by 18
“This conclusion is in line with other recent studies [16] that implicate land plants as the cause of late O 2 rise in Earth history (for a recent review see [7]).”
BackgroundResultInfluentialA Systems Approach to Understanding How Plants Transformed Earth's Environment in Deep Time
Matthaeus, W. J., Macarewich, S. I., Richey, J. D., Montañez, I. P., McElwain, J. C., White, J. D., Wilson, J. P., & Poulsen, C. J. (2023) · Annual Review of Earth and Planetary Sciences · doi:10.1146/annurev-earth-080222-082017 · Cited by 18
Crustal carbonate build-up as a driver for Earth’s oxygenation
Alcott, L., Walton, C. R., Planavsky, N. J., Shorttle, O., & Mills, B. J. W. (2024) · Nature Geoscience · doi:10.1038/s41561-024-01417-1 · Cited by 22
Self-citationBowles, A. M. C., Williams, T. A., Donoghue, P. C. J., Campbell, D. A., & Williamson, C. J. (2024) · New Phytologist · doi:10.1111/nph.19860 · Cited by 24
“The evolution of terrestrial flora (Embryophyta) (de Vries & Archibald, 2018) transformed Earth’s continents, atmosphere and climate (Lenton et al ., 2016; Bengtson et al ., 2017; Leebens-Mack et al ., 2019; Delaux & Schornack, 2021; Bowles et al ., 2023), and promoted the diversification of multiple…”
BackgroundEvolving circuitries in plant signaling cascades
Zegers, J. M. S., Irisarri, I., de Vries, S., & de Vries, J. (2024) · Journal of Cell Science · doi:10.1242/jcs.261712 · Cited by 19
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Tognacca, R. S., Kubaczka, M. G., Servi, L., Rodríguez, F. S., Herz, M. A. G., & Petrillo, E. (2020) · Transcription · doi:10.1080/21541264.2020.1796473 · Cited by 17
“[11] Lenton TM, Dahl TW, Daines SJ, et al.”
BackgroundMarine oxygenation, deoxygenation, and life during the Early Paleozoic: An overview
Zhang, J., Edwards, C. T., Diamond, C., Lyons, T. W., & Zhang, Y. (2021) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2021.110715 · Cited by 18
Shepherd, M., Giordano, D., Verde, C., & Poole, R. K. (2022) · Antioxidants · doi:10.3390/antiox11071222 · Cited by 20
“Atmospheric O2 concentrations reached modern levels in the Devonian period as a result of photosynthesis [21], and with the consequent change in the oxidation state of the atmosphere, H2S was eliminated as a primary electron donor, and NO and CO became less advantageous as electron acceptors [22].”
BackgroundKozik, N. P., Young, S. A., Ahlberg, P., Lindskog, A., & Owens, J. (2023) · Global and Planetary Change · doi:10.1016/j.gloplacha.2023.104183 · Cited by 17
Zhang, X., Liu, J., Wang, Y., Jiayu, R., Zhan, R., Xu, H., & Tang, P. (2018) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2018.02.005 · Cited by 18
Adiatma, Y., Saltzman, M. R., Young, S. A., Griffith, E. M., Kozik, N. P., Edwards, C. T., Leslie, S. A., & Bancroft, A. M. (2019) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2019.109341 · Cited by 18
Self-citationLow atmospheric CO2 levels before the rise of forested ecosystems
Dahl, T. W., Harding, M. A. R., Brugger, J., Feulner, G., Norrman, K., Lomax, B. H., & Junium, C. K. (2022) · Nature Communications · doi:10.1038/s41467-022-35085-9 · Cited by 19
Self-citationEarth: Atmospheric Evolution of a Habitable Planet
Olson, S. L., Schwieterman, E. W., Reinhard, C. T., & Lyons, T. W. (2018) · eScholarship (California Digital Library) · Cited by 18
Seawater signatures of Ordovician climate and environment
Young, S. A., Edwards, C. T., Ainsaar, L., Lindskog, A., & Saltzman, M. R. (2022) · Geological Society London Special Publications · doi:10.1144/sp532-2022-258 · Cited by 17
Self-citationZhao, K., Zhu, G.-Y., Zheng, W., Sun, G.-Y., Li, T., & Zhang, Z.-Y. (2023) · Global and Planetary Change · doi:10.1016/j.gloplacha.2023.104214 · Cited by 16
Hill, R., Wang, Z., Williams, G. D. Z., Polyak, V. J., Singh, A., Kipp, M. A., Asmerom, Y., & Vengosh, A. (2024) · Chemical Geology · doi:10.1016/j.chemgeo.2024.122214 · Cited by 15
Origin and phylogenetic relationships of [4Fe–4S]‐containing O 2 sensors of bacteria
Barth, C., Weiss, M. C., Roettger, M., Martin, W. F., & Unden, G. (2018) · Environmental Microbiology · doi:10.1111/1462-2920.14411 · Cited by 15
“O2 levels of 0.02–0.04 atm resulted in mild oxygenation of ocean surface waters, a situation that persisted for almost 2 billion years (Lenton et al., 2016) because deep ocean oxygenation was not completed until roughly 580–430 million years ago (Stolper and Keller, 2018).”
BackgroundInfluentialThe Evolution of Symbiotic Plant–Microbe Signalling
Clear, M. R., & Hom, E. F. Y. (2019) · Wiley eBooks · doi:10.1002/9781119312994.apr0684 · Cited by 18
Ocean oxygenation and ecological restructuring caused by the late Paleozoic evolution of land plants
Stacey, J., Wallace, M. W., Hood, A. V. S., Shuster, A., Corlett, H., Reed, C. P., & Moynihan, C. (2024) · Geology · doi:10.1130/g52502.1 · Cited by 15
Diversity dynamics of Devonian terrestrial palynofloras from China: Regional and global significance
Shen, Z., Monnet, C., Cascales‐Miñana, B., Gong, Y., Dong, X., Kroeck, D. M., & Servais, T. (2019) · Earth-Science Reviews · doi:10.1016/j.earscirev.2019.102967 · Cited by 15
Rosa, D., Leach, D., Guarnieri, P., & Bekker, A. (2022) · Mineralium Deposita · doi:10.1007/s00126-022-01125-z · Cited by 15
Chen, K., Liao, H., Bellenger, J., & Lutzoni, F. M. (2019) · Environmental Microbiology · doi:10.1111/1462-2920.14605 · Cited by 14
“Mosses (phylumBryophyta) are amajor component of boreal, alpine and arctic vegetations, and contribute significantly to global nutrient cycles (Brown and Bates, 1990; Turetsky, 2003; Lenton et al., 2016).”
BackgroundZygnematophycean algae: Possible models for cellular and evolutionary biology
Ohtaka, K., & Sekimoto, H. (2022) · Seminars in Cell and Developmental Biology · doi:10.1016/j.semcdb.2022.03.042 · Cited by 13
Hg isotope evidence for oceanic oxygenation during the Cambrian Explosion
Fan, H., Fu, X., Yang, R., Wen, H., Hu, C., Ward, J. F., Zhang, H., Zhang, H., & Zhang, X. (2023) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2023.10.020 · Cited by 15
The Bretskyan hierarchy, multiscale allopatry, and geobiomes—on the nature of evolutionary things
Spiridonov, A., & Eldredge, N. (2024) · Paleobiology · doi:10.1017/pab.2023.37 · Cited by 14
“The new characteristics of previously unoccupied environments impose new sets of physical and geo-chemical constraints and feedbacks on these constraints (Kenrick et al. 2012; Lenton et al. 2016).”
BackgroundWu, J., Deng, Q., Qiu, Y., Liu, C., Lin, C., Ru, Y., Sun, Y., Lai, J., Liu, L., Shen, X., et al. (2024) · New Phytologist · doi:10.1111/nph.20040 · Cited by 17
Boyce, C. K., Ibarra, D. E., Nelsen, M. P., & D’Antonio, M. P. (2022) · Geobiology · doi:10.1111/gbi.12519 · Cited by 15
Ma, C., Tang, Y., & Ying, J. (2022) · Earth-Science Reviews · doi:10.1016/j.earscirev.2022.104101 · Cited by 14
Simulating the long-term carbon cycle in the Phanerozoic: Current status and future developments
Zhang, Y., Benjamin, J. W. M., He, T., Yang, T., & Zhu, M. (2022) · Chinese Science Bulletin (Chinese Version) · doi:10.1360/tb-2022-0813 · Cited by 13
Power, competition, and the nature of history
Vermeij, G. J. (2019) · Paleobiology · doi:10.1017/pab.2019.34 · Cited by 13
“Lenton et al. (2016) estimate that productivity before the advent of vascular plants was about one-third the present average.”
BackgroundCole, D., Ozaki, K., & Reinhard, C. (2021) · Global Biogeochemical Cycles · doi:10.1029/2021gb007052 · Cited by 7
“As a result, the transition of the Earth system from a relatively stable mid-Proterozoic world to something closer to the modern state, perhaps by the late Paleozoic (e.g., Dahl et al., 2010; Lenton et al., 2016; Wallace et al., 2017), represents a vast period of time across which we have a relatively poor understandi…”
BackgroundInfluentialSelf-citationDynamic deep marine oxygenation during the Early and Middle Paleozoic
Ostrander, C. M., Clemente, J. N. R., Stockey, R., Strauss, J. V., Fraser, T. A., Nielsen, S. G., & Sperling, E. A. (2025) · Science Advances · doi:10.1126/sciadv.adw5878 · Cited by 16
Mrnjavac, N., Nagies, F. S. P., Wimmer, J. L. E., Kapust, N., Knopp, M. R., Trost, K., Modjewski, L., Bremer, N., Mentel, M., Esposti, M. D., et al. (2024) · FEBS Letters · doi:10.1002/1873-3468.14906 · Cited by 15
“Evidence for the appearance of O 2 in the atmosphere by the GOE 2.4 billion years ago is universally accepted and uncontroversial [1,2,10,11].”
InfluentialLithium isotope stratigraphy and Ordovician weathering
Adiatma, Y., Saltzman, M. R., Liu, X., Wang, X.-K., & Edwards, C. T. (2024) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2024.119030 · Cited by 12
Self-citationMarine sulphate captures a Paleozoic transition to a modern terrestrial weathering environment
Waldeck, A. R., Olson, H. C., Crockford, P. W., Couture, A. M., Cowie, B. R., Hodgin, E. B., Bergmann, K., Dewing, K., Grasby, S. E., Clark, R. J., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-57282-y · Cited by 15
Biotic effects on oxygen consumption during weathering: Implications for the second rise of oxygen
Kanzaki, Y., & Kump, L. R. (2017) · Geology · doi:10.1130/g38869.1 · Cited by 12
Watanabe, Y., & Tajika, E. (2021) · Earth Planets and Space · doi:10.1186/s40623-021-01527-9 · Cited by 12
Regional synthesis of the Ordovician geology and stratigraphy of China
Zhang, Y., Zhan, R., Zhen, Y. Y., Wang, W., Liang, Y., Fang, X. M., Wu, R., Yan, K., Zhang, J., & Li, W. (2022) · Geological Society London Special Publications · doi:10.1144/sp533-2022-128 · Cited by 13
Phanerozoic oceanic and climatic perturbations in the context of Tethyan evolution
Tian, L., Song, H., Liu, Y., Wu, Y., Chu, D., & Song, H. (2023) · Science China Earth Sciences · doi:10.1007/s11430-023-1205-6 · Cited by 13
“Numerical simulations based on COPSE and GEOCARBSULF models further support this long-term trend, confirming the rapid increase in atmospheric O 2 during the Devonian (Berner, 2006; Lenton et al., 2016).”
MethodCyst size trends in the genus Leiosphaeridia across the Mulde (lower Silurian) biogeochemical event
Spiridonov, A., Venckutė-Aleksienė, A., & Radzevičius, S. (2017) · Bulletin of Geosciences · doi:10.3140/bull.geosci.1679 · Cited by 13
Graham, L. E., Graham, J. M., Wilcox, L. W., Cook, M. E., Arancibia‐Ávila, P., & Knack, J. J. (2018) · International Journal of Plant Sciences · doi:10.1086/698709 · Cited by 14
“…early bryophyte-like land plants, which contributed to soil stabilization and organic C content and, by producing degradation-resistant tissues, helped to foster a period of plant-driven CO2 drawdown prior to the rise of vascular plants (Graham et al. 2004a; Field et al. 2015; Lenton et al. 2016).”
BackgroundInfluentialHoysted, G. A., Kowal, J., Pressel, S., Duckett, J. G., Bidartondo, M. I., & Field, K. J. (2021) · Mycorrhiza · doi:10.1007/s00572-021-01033-6 · Cited by 13
“Geochemical models support fossil and stable isotope evidence indicating 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 stepwise increase of the Earth’s atmospheric oxygen ([O2]), and a simult…”
BackgroundOzaki, K., Cole, D. B., Reinhard, C. T., & Tajika, E. (2022) · Geoscientific model development · doi:10.5194/gmd-15-7593-2022 · Cited by 14
Self-citationLife's a beach – the colonization of the terrestrial environment
Plackett, A. R. G., & Coates, J. C. (2016) · New Phytologist · doi:10.1111/nph.14295 · Cited by 12
Time-resolved oxidative signal convergence across the algae–embryophyte dividePreprint
Rieseberg, T. P., Dadras, A., Darienko, T., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.03.11.584470 · Cited by 12
Okafor, M.-M., & Azmy, K. (2023) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2023.111978 · Cited by 11
Model based Paleozoic atmospheric oxygen estimates: a revisit to GEOCARBSULF
Zhang, S., Planavsky, N. J., Krause, A. J., Bolton, E. W., & Mills, B. J. W. (2018) · American Journal of Science · doi:10.2475/05.2018.05 · Cited by 11
Self-citationWang, H., Wang, D., Wei, G., Ling, H., Struck, U., Wei, W., Yao, S., Cheng, C., Li, J., Sun, Y., et al. (2021) · Precambrian Research · doi:10.1016/j.precamres.2021.106501 · Cited by 11
Evolution of ABA signaling pathways
Cuming, A. C. (2019) · Advances in botanical research · doi:10.1016/bs.abr.2019.06.003 · Cited by 13
Porada, P., Tamm, A., Kleidon, A., et al. (2018) · doi:10.5194/bg-2018-324 · Cited by 11
Self-citationA roadmap to developing unified streptophyte algal model systems
Carrillo‐Carrasco, V. P., Hernández‐García, J., Girou, C., Grubor, I., Keller, J., Lim, E., Schmidt, V., Sørensen, I., Vosolsobě, S., Buschmann, H., et al. (2025) · Current Biology · doi:10.1016/j.cub.2025.05.023 · Cited by 11
“Land plants have established today’s oxygen levels 3 and the web of symbiotic interactions in the soil 4 .”
Advancing studies on global biocrust distribution
Wang, S., Ma, L., Yang, L., Ma, Y., Zhang, Y., Zhao, C., & Chen, N. (2024) · SOIL · doi:10.5194/soil-10-763-2024 · Cited by 12
Exploring Neoproterozoic climate and biogeochemical evolution in the SCION model
Mills, B. J. W., Hir, G. L., Merdith, A., Gurung, K., Bowyer, F. T., Krause, A. J., Sánchez‐Baracaldo, P., Hunter, S. J., & Zhang, Y. (2025) · Global and Planetary Change · doi:10.1016/j.gloplacha.2025.104791 · Cited by 12
“They have also been used to test the hypothesis that early Paleozoic oxygen levels were lower than present (Lenton et al., 2016; Krause et al., 2018), and that Permo-Carboniferous oxygen levels were higher (Berner, 2006; Bergman et al., 2004) and they have been used to reconstruct marine sulfate…”
Self-citationDiversification to extinction: oceanic and climatic context of the Ordovician
Liu, M., Bao, X., Harper, D., Algeo, T. J., Zhao, M., Saltzman, M., Zhang, W., Chen, D.-Z., Yuan, S., Chen, Y.-H., et al. (2025) · Earth-Science Reviews · doi:10.1016/j.earscirev.2025.105194 · Cited by 10
Self-citationFuchsman, C. A., & Stüeken, E. E. (2020) · Environmental Microbiology · doi:10.1111/1462-2920.15220 · Cited by 9
Yang, X., Yan, D., Zhang, B., & Zhang, L. (2023) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2023.111859 · Cited by 10
Liu, X., Krause, A. J., Wilson, D. J., Fraser, W. T., Joachimski, M. M., Brand, U., Stigall, A. L., Qie, W., Chen, B., Yang, X., et al. (2025) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2025.02.036 · Cited by 10
“…that there could have been decreases in atmospheric CO 2 (from 2000 ppm to less than 1000 ppm) and increases in O 2 concentrations, which would have been beneficial in creating more habitable niches for multicellular biota (e.g., Copper and Scotese, 2003; Foster et al., 2017; Lenton et al., 2016).”
Ordovician marine Charophyceae and insights into land plant derivations
Liu, L., Han, J., Zhang, Z., Tang, Q., Pang, K., Li, R., Wu, Y., Hua, H., Guo, B., Cai, C., et al. (2025) · Nature Plants · doi:10.1038/s41477-025-02003-y · Cited by 10
Metal-rich stars are less suitable for the evolution of life on their planets
Shapiro, A. V., Brühl, C., Klingmüller, K., Steil, B., Shapiro, A. I., Witzke, V., Костогрыз, Н. М., Gizon, L., Solanki, S. K., & Lelieveld, J. (2023) · Nature Communications · doi:10.1038/s41467-023-37195-4 · Cited by 10
“The surface boundary conditions for O 2 and CO 2 on paleo-Earth are established following available geological reconstructions 24,25,27 , see Supplementary Table 1.”
InfluentialThe conquest of spaces: Exploring drivers of morphological shifts through phylogenetic palaeoecology
Lamsdell, J. C. (2021) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2021.110672 · Cited by 11
“…(Carbone and Narbonne, 2014; M ´ angano and Buatouis, 2015; Cribb et al., 2019) and the impact of the radiation of vascular land plants on atmospheric oxygen and continental weathering during the Devonian (Dahl et al., 2010; Lenton et al., 2016; Pawlik et al., 2020; S ø nderholm and Bjerrum, 2021).”
The origin and distribution of the main oxygen sensing mechanism across metazoans
Song, B., Modjewski, L., Kapust, N., Mizrahi, I., & Martin, W. F. (2022) · Frontiers in Physiology · doi:10.3389/fphys.2022.977391 · Cited by 7
“, 2019); during the Phanerozoic period (541–0 Ma), the atmospheric oxygen content increased constantly to the modern level (Lenton et al., 2016; Krause et al., 2018).”
BackgroundThe record of sea water chemistry evolution during the Ediacaran–Cambrian from early marine cements
Xiong, Y., Wood, R. A., & Pichevin, L. (2022) · The Depositional Record · doi:10.1002/dep2.211 · Cited by 10
“…levels comparable to the modern (average of all post- middle Devonian Ceanom = 0.55), which may be related to the rise of the land plants which probably increased weathering regimes and the extent of organic matter burial (Algeo & Scheckler, 1998; Lenton et al., 2016; Wallace et al., 2017).”
BackgroundInfluentialEnvironmental gradients reveal stress hubs predating plant terrestrializationPreprint
Dadras, A., Fürst‐Jansen, J. M. R., Darienko, T., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.10.17.512551 · Cited by 9
A reassessment of the “hard-steps” model for the evolution of intelligent life
Mills, D. B., Macalady, J. L., Frank, A., & Wright, J. T. (2025) · Science Advances · doi:10.1126/sciadv.ads5698 · Cited by 14
“…of dynamic and increasing p O 2 levels ( 117 – 119 ), p O 2 stabilized near present values ( ~ 100% PAL, or ~ 21% of the atmosphere by volume) by 420 to 400 Ma during the so-called Paleozoic Oxygenation Event (POE) ( 117 , 120 – 122 ), and have been maintained near these levels ever since (Fig.”
Repeated occurrences of marine anoxia under high atmospheric O 2 and icehouse conditions
Chen, J., Li, S., Zhang, S., Isson, T. T., Dahl, T. W., Planavsky, N. J., Zhang, F., Wang, X., Shen, S., & Montañez, I. P. (2025) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2420505122 · Cited by 10
Self-citationLenton, T. M. (2025) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2024.0095 · Cited by 9
“Finally in the Palaeozoic, the rise of land plants and associated resource recycling roughly doubled global productivity to the modern level, increasing oxygen levels and cooling the climate [83,84].”
Self-citationThe Interplay between Enucleated Sieve Elements and Companion Cells
Matilla, A. J. (2023) · Plants · doi:10.3390/plants12173033 · Cited by 8
“Taken together, sessile life and terrestrial colonization have been attributed to a series of major improvements in plant body plans, anatomy, and biochemistry, resulting in a substantial decrease in atmospheric CO2 levels and progressive oxygenation of the environment [9].”
BackgroundThe Power of a Systems Approach to Mineral and Petroleum Exploration in Sedimentary Basins
McCuaig, T. C., Scarselli, S., O’Connor, T., Busuttil, S., & McCormack, N. (2018) · doi:10.5382/sp.21.03 · Cited by 9
Zhang, X., Liu, J., Wang, Y., & Xu, H. (2020) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2020.109772 · Cited by 8
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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