The DOI matches its Crossref record; 54 of the 54 references it cites were found in a registry or an index; 3 of the 31 sources its claims cite were checked for support.
Oxygen requirements of the earliest animals
Open accessAbstract
A rise in the oxygen content of the atmosphere and oceans is one of the most popular explanations for the relatively late and abrupt appearance of animal life on Earth. In this scenario, Earth's surface environment failed to meet the high oxygen requirements of animals up until the middle to late Neoproterozoic Era (850-542 million years ago), when oxygen concentrations sufficiently rose to permit the existence of animal life for the first time. Although multiple lines of geochemical evidence support an oxygenation of the Ediacaran oceans (635-542 million years ago), roughly corresponding with the first appearance of metazoans in the fossil record, the oxygen requirements of basal animals remain unclear. Here we show that modern demosponges, serving as analogs for early animals, can survive under low-oxygen conditions of 0.5-4.0% present atmospheric levels. Because the last common ancestor of metazoans likely exhibited a physiology and morphology similar to that of a modern sponge, its oxygen demands may have been met well before the enhanced oxygenation of the Ediacaran Period. Therefore, the origin of animals may not have been triggered by a contemporaneous rise in the oxygen content of the atmosphere and oceans. Instead, other ecological and developmental processes are needed to adequately explain the origin and earliest evolution of animal life on Earth.
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Claims the paper makes
3 of 31 cited sources checked · 1 supported · 0 flagged · accuracy not yet measured
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Internal anoxia, however, does appear to be a common feature of sponge physiology (34,35).”
§ Oxygen requirements of the earliest animals > Results and Discussion
What the sources say
- Not checked — no full text of the sourcecites [34] 34. Hoffmann F, et al. (2005) An anaerobic world in sponges. Geomicrobiol J 22(1-2):1-10. source
- Partly supportedcites [35] 35. Hoffmann F, et al. (2008) Oxygen dynamics and transport in the Mediterranean sponge Aplysina aerophoba. Mar Biol 153(6):1257-1264. sourceExperimental — accuracy not yet measured
The Abstract and the Discussion section on oxygen dynamics report internal anoxia in non-pumping A. aerophoba, with tissue becoming anoxic within 15 minutes after pumping stops. They do not establish that internal anoxia is common across sponges; the Discussion also notes that prolonged non-pumping was not observed in an in situ study of A. aerophoba unless the sponge was disturbed. (LLM, conf=0.88)
Load-bearingStrength:hedged· not yet calibratedTests✓ 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
“Overall, the patterns of oxygen uptake observed during the initial drawdown to anoxia followed Michaelis-Menten kinetics, with a V max of 2.42% air saturation/h•g dry weight , or 7.93 μM O 2 /h•g dry weight , and a K m of 4.86% of air saturation, or 15.9 μM O 2 , at the water salinity and temperature of our experiments (Fig. 3).”
§ Oxygen requirements of the earliest animals > Results and Discussion
Tests✓ 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
“Here we show that modern demosponges, serving as analogs for early animals, can survive under low-oxygen conditions of 0.5-4.0%present atmospheric levels.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Sponges were also observed on the Volcano 7 seamount in the eastern tropical Pacific, where oxygen concentrations ranged from 4.5 to 7.2 μM O 2 (31) .”
§ Oxygen requirements of the earliest animals > Significance
What the sources say
- Not checked — no full text of the sourcecites [31] 31. Levin LA, Huggett CL, Wishner KF (1991) Control of deep-sea benthic community structure by oxygen and organic matter gradients in the eastern Pacific Ocean. J Mar Res 49(4):763-800. source
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ 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
“Therefore, the origin of animals may not have been triggered by a contemporaneous rise in the oxygen content of the atmosphere and oceans.”
Load-bearingStrength:hedged· 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
“Because the last common ancestor of metazoans likely exhibited a physiology and morphology similar to that of a modern sponge, its oxygen demands may have been met well before the enhanced oxygenation of the Ediacaran Period.”
Tests✓ 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
“It would appear, then, that LMA sponges, such as H. panicea, may be better adapted to lowoxygen conditions than HMA sponges.”
§ Oxygen requirements of the earliest animals > Results and Discussion
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Numerous phylogenetic analyses conclude that sponges are the earliest diverging extant metazoan group (11,12,(20)(21)(22).”
What the sources say
- Not matchedcites (11,12,(20)(21)(22)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes- Not matched
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
“However, although these theoretical efforts provide some guidance, they are based on highly simplified, idealized organisms that do not reflect the complexity of living animals.”
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Choanoflagellates, in turn, are the closest living relatives of animals (24)(25)(26)(27)(28).”
What the sources say
- Not matchedcites (24)(25)(26)(27)(28)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passes- Not matched
- It may state two separate claims as one. 15
- Probably not a substantial claim. 3
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Indeed, oxygen levels this high may have been reached as early as the Great Oxidation Event some 2.3-2.4 billion years ago (43,45) and may have persisted at these levels through much of the intervening time (43,44), with some likely swings to lower levels, however, around 1.8 billion years ago (46).”
§ Oxygen requirements of the earliest animals > Sponge Evolution and Environmental Oxygen Levels.
What the sources say
- Not checked — no full text of the sourcecites [43] 43. Kump LR (2008) The rise of atmospheric oxygen. Nature 451(7176):277-278. source
- Not checked — no full text of the sourcecites [45] 45. Bekker A, Holland HD (2012) Oxygen overshoot and recovery during the early Pale- oproterozoic. Earth Planet Sci Lett 317-318:295-304. source
- Not checked — no full text of the sourcecites [44] 44. Canfield DE (2014) Proterozoic Atmospheric Oxygen. Treatise on Geochemistry, eds Holland HD, Turekian KK (Elsevier, Oxford), 2nd Ed., pp 197-216. source
- Partly supportedcites [46] 46. Canfield DE, et al. (2013) Oxygen dynamics in the aftermath of the Great Oxidation of Earth's atmosphere. Proc Natl Acad Sci USA 110(42):16736-16741. sourceExperimental — accuracy not yet measured
The Significance section (page unavailable) places a rise in oxygen at the Great Oxidation Event about 2.3 billion years ago, followed by elevated levels for roughly 200 million years and then a collapse. The Results and Discussion section (page unavailable) describes later low-oxygen episodes, but the paper does not establish the cited 0.5–4% PAL range or its persistence through much of the intervening time. (LLM, conf=0.85)
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
The sources with a verdict support this claim; the rest weren’t checked yet, have no full text we could read, or their verdicts aren’t rated yet.
In the paper
“Indeed, according to present evidence, the earliest proposed signs of animals in the rock record (9,10), as well as molecular clock divergence estimates (11,12), predate the earliest evidence for deep ocean oxygenation (1), potentially by tens of millions of years.”
What the sources say
- Not checked — no full text of the sourcecites [9] 9. Love GD, et al. (2009) Fossil steroids record the appearance of Demospongiae during the Cryogenian period. Nature 457(7230):718-721. source
- Not checked — no full text of the sourcecites [10] 10. Maloof AC, et al. (2010) Possible animal-body fossils in pre-Marinoan limestones from South Australia. Nat Geosci 3(9):653-659. source
- Supportedcites [11] 11. Peterson KJ, Butterfield NJ (2005) Origin of the Eumetazoa: Testing ecological pre- dictions of molecular clocks against the Proterozoic fossil record. Proc Natl Acad Sci USA 102(27):9547-9552. source
The Results (page unavailable) give molecular-clock estimates for early animal divergences: the minimum-evolution analysis dates the origin of Metazoa to 664 Ma and brackets eumetazoan evolution between 634 and 604 Ma. This paper supports the molecular-clock dates attributed to citations (11,12); the separate citation (1), not this paper, is offered for the timing of deep-ocean oxygenation. (LLM, conf=0.86)
- Not checked — no full text of the sourcecites [12] 12. Erwin DH, et al. (2011) The Cambrian conundrum: Early divergence and later eco- logical success in the early history of animals. Science 334(6059):1091-1097. source
- Not checked — no full text of the sourcecites [1] 1. Sahoo SK, et al. (2012) Ocean oxygenation in the wake of the Marinoan glaciation. Nature 489(7417):546-549. 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
“We show that although sensitive to oxygen, H. panicea can both tolerate and grow at low oxygen levels from 0.5% to 4% PAL.”
- Also stated in: § Results and Discussion · plain
“Overall, the sponges were able to both respire and feed to oxygen levels cycling between 0.5% and 4% PAL.”
§ Oxygen requirements of the earliest animals > Significance
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Also stated in: § Results and Discussion · plain
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
“functional gene and 16S rRNA analyses reveal a significant population of ammonium-oxidizing Archaea and betaproteobacteria, as well as other aerobic organisms (Fig. S6 and Table S1).”
§ Oxygen requirements of the earliest animals > Results and Discussion
Strength:plain· 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
“Therefore, the majority of the oxygen uptake and Rhodomonas consumption observed during the experiments can be attributed to the sponges, with the background rates likely attributed to free-living microbes in our experimental system.”
§ Oxygen requirements of the earliest animals > Results and Discussion
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
“Our results suggest that H. panicea is sensitive to both anoxia and prolonged exposure to relatively stagnant seawater.”
§ Oxygen requirements of the earliest animals > Results and Discussion
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A claim is checked only against the cited paper’s full text, never its abstract, and none is available for its sources.
In the paper
“This K m is less than half of the 10% PAL serving as the estimated minimum level of atmospheric oxygen thought to have permitted the oxygenation of the deep oceans around 580 Ma (2), as well as the level estimated to permit the existence of millimeter-width, diffusionlimited animals (15).”
§ Oxygen requirements of the earliest animals > Results and Discussion
What the sources say
- Not checked — no full text of the sourcecites [2] 2. Canfield DE, Poulton SW, Narbonne GM (2007) Late-Neoproterozoic deep-ocean oxygenation and the rise of animal life. Science 315(5808):92-95. source
- Not checked — no full text of the sourcecites [15] 15. Catling DC, Glein CR, Zahnle KJ, McKay CP (2005) Why O2 is required by complex life on habitable planets and the concept of planetary "oxygenation time" Astrobiology 5(3):415-438. source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A claim is checked only against the cited paper’s full text, never its abstract, and none is available for its sources.
In the paper
“Although levels of atmospheric oxygen may have reached 10% PAL by 580 Ma, the geochemical record does not preclude oxygen levels of 0.5-4% PAL during the Cryogenian Period (43,44).”
§ Oxygen requirements of the earliest animals > Sponge Evolution and Environmental Oxygen Levels.
What the sources say
- Not checked — no full text of the sourcecites [43] 43. Kump LR (2008) The rise of atmospheric oxygen. Nature 451(7176):277-278. source
- Not checked — no full text of the sourcecites [44] 44. Canfield DE (2014) Proterozoic Atmospheric Oxygen. Treatise on Geochemistry, eds Holland HD, Turekian KK (Elsevier, Oxford), 2nd Ed., pp 197-216. source
Strength:hedged· 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
“Such evidence can reasonably be obtained by studying modern animals that resemble the earliest metazoans.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✗ Based: failsProbably not a substantial claim. The paper may give no reason why it holds.
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
“Given the complete absence of necrosis in the sponges maintained at both low and high oxygen levels during this run, the necrosis observed during experiment 1 was likely due to the infrequent exchange of fresh seawater and not necessarily the low oxygen levels.”
§ Oxygen requirements of the earliest animals > Results and Discussion
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
“This suggests that although 3-4% PAL oxygen may not compromise H. panicea's growth or maintenance of healthy tissue, anoxia still does.”
§ Oxygen requirements of the earliest animals > Results and Discussion
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“This suggests that the similarities between choanocytes and choanoflagellates were inherited from a common ancestor and that the choanocytebased feeding mode, as retained by sponges, is likely ancestral to all animals (29).”
What the sources say
- Not checked — no full text of the sourcecites [29] 29. Nichols SA, Dayel MJ, King N (2009) Genomic, phylogenetic, and cell biological in- sights into metazoan origins. Animal Evolution: Genes, Genomes, Fossils and Trees, eds Telford MJ, Littlewood DTJ (Oxford University Press, Oxford), pp 24-32. source
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
“Furthermore, to establish that low levels of atmospheric oxygen, in fact, delayed the origin of animal life on Earth, it is necessary to know the minimum oxygen requirements of early animals.”
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim.
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
“Therefore, to more conclusively determine the minimum oxygen requirements of metazoans, direct experimental evidence is required.”
Load-bearingStrength:boosted· not yet calibratedTests✓ Asserted: passes✗ Checkable: fails✓ Load-bearing: passes✓ Based: passesProbably not a substantial claim. It may be too vague to check against evidence.
None of the sources cited for this claim have been checked yet.
In the paper
“LMA sponges are known to have shorter and wider water canals (38), as well as larger choanoflagellate chambers (39), than high-microbial abundance (HMA) sponges.”
§ Oxygen requirements of the earliest animals > Results and Discussion
What the sources say
- Not checked — no full text of the sourcecites [38] 38. Vacelet J, Donadey C (1977) Electron-microscope study of association between some sponges and bacteria. J Exp Mar Biol Ecol 30(3):301-314. source
- Not matchedcites [39] 39. Boury-Esnault N, de Vos L, Donadey C, Vacelet J (1990) Ultrastructure of Choanosome and Sponge Classification (Smithsonian Institution Press, Washington, DC), pp 237-244.
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
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
“Under this scenario, the origin of animal life was not a response to atmospheric oxygenation but part of the ecological reworking of marine biogeochemical cycles.”
§ Oxygen requirements of the earliest animals > Sponge Evolution and Environmental Oxygen Levels.
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
A claim is checked only against the cited paper’s full text, never its abstract, and none is available for its sources.
In the paper
“Recent molecular clock estimates place the last common ancestor of all sponges at nearly 800 Ma (12), whereas putative sponge-like body fossils are found in rocks dated between 660 and 635 Ma (10) in the Cryogenian Period (850-635 Ma).”
§ Oxygen requirements of the earliest animals > Sponge Evolution and Environmental Oxygen Levels.
What the sources say
- Not checked — no full text of the sourcecites [12] 12. Erwin DH, et al. (2011) The Cambrian conundrum: Early divergence and later eco- logical success in the early history of animals. Science 334(6059):1091-1097. source
- Not checked — no full text of the sourcecites [10] 10. Maloof AC, et al. (2010) Possible animal-body fossils in pre-Marinoan limestones from South Australia. Nat Geosci 3(9):653-659. source
Load-bearingStrength:hedged· not yet calibratedTests✓ 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
“We provide experimental evidence suggesting that the last common ancestor of animals could have thrived in oxygen levels as low as 0.5% to 4% of present atmospheric levels, which were likely met on Earth well before animals evolved.”
§ Oxygen requirements of the earliest animals > Significance
Load-bearingStrength:hedged· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
The paper relies on this without support.
The minimum oxygen requirements of modern sponges and their metabolic response to low oxygen had not been investigated prior to this study.
“Despite these observations, however, the minimum oxygen requirements of modern sponges and their metabolic response to low oxygen have not, to our knowledge, been investigated.”
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?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Indeed, under most conditions, the estimated tolerance range for basal metazoan organisms (12) is only safely exceeded throughout the year at p O2 on the order of ∼ 5 -10% PAL.”
From Earth's oxygen cycle and the evolution of animal life (2016) · cites this paper as [12]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Experimental work with the modern demosponge Halichondria panacea suggests an empirical O2 threshold of ∼ 0.5 -4.0% of the present atmospheric level (PAL; corresponding to an equilibrium concentration of ∼ 1 -10 μ mol · kg -1 for air-saturated water at 25 °C and a salinity of 35 ‰ ) for basal metazoan life (12)”
From Earth's oxygen cycle and the evolution of animal life (2016) · cites this paper as [12]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Finally, in the greater context of what triggered the emergence of complex animal ecosystems on Earth, our isotope data and models suggest that there were ample amounts of O2 for animals in the surface ocean at the onset of the Cambrian (69).”
From Atmosphere-ocean oxygen and productivity dynamics during early animal radiations (2019) · cites this paper as [69]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“The early originating suspension-feeding sponges had low O2 requirements63, and may have helped consume the large dissolved organic carbon reservoir that acted as a major redox buffer in the Proterozoic ocean12.”
From Rise to modern levels of ocean oxygenation coincided with the Cambrian radiation of animals (2015) · cites this paper as [63]
The check wasn’t confident either way, so it gives no verdict — never disputed.
In the citing paper
“The BACE pre-dates the first appearance of Treptichnus pedum (31,32)”
From Integrated records of environmental change and evolution challenge the Cambrian Explosion (2019) · cites this paper as [31]
Experimental — accuracy not yet measured
In the citing paper
“Recent work by Mills et al. (2014), however, suggests that primitive metazoans, such as sponges, may have needed very little oxygen in the water column in order to thrive.”
From Trace elements at the intersection of marine biological and geochemical evolution (2016) · cites this paper as Mills et al., 2014
In the citing paper
“This billion-year stagnation in eukaryotic diversification is often attributed to the late rise of atmospheric oxygen (e.g., Nursall, 1959;Knoll and Carrol, 1999;Fedonkin, 2003;Planavksy et al., 2014a) to levels required by metazoans (i.e.0.5% to 4% PAL; Mills et al., 2014).”
From Trace elements at the intersection of marine biological and geochemical evolution (2016) · cites this paper as Mills et al., 2014
In the citing paper
“This level of dissolved O2 in surface waters is generally consistent with the 0.1-1% PAL proposed as necessary for early animals”
From Trace elements at the intersection of marine biological and geochemical evolution (2016) · cites this paper as Mills et al., 2014
In the citing paper
“The recognition that modern sponges can survive at very low oxygen levels (e.g.Mills et al., 2014;though it has not yet been demonstrated that they can complete their entire life cycle under such conditions) suggests that the initial evolution of animals may have taken place at lower oxygen levels than are generally assumed necessary.”
From Remarkable insights into the paleoecology of the Avalonian Ediacaran macrobiota (2015) · cites this paper as Mills et al., 2014
Citations
Cites
Kasting, J. F. (1993) · Science · doi:10.1126/science.11536547 · Cited by 1,374
Erwin, D. H., Laflamme, M., Tweedt, S. M., Sperling, E. A., Pisani, D., & Peterson, K. J. (2011) · Science · doi:10.1126/science.1206375 · Cited by 1,336 · cited 3×
“Recent molecular clock estimates place the last common ancestor of all sponges at nearly 800 Ma (12), whereas putative sponge-like body fossils are found in rocks dated between 660 and 635 Ma (10) in the Cryogenian Period (850–635 Ma).”
BackgroundThe genome of the choanoflagellate Monosiga brevicollis and the origin of metazoans
King, N., Westbrook, M. J., Young, S. L., Kuo, A., Abedin, M., Chapman, J., Fairclough, S., Hellsten, U., Isogai, Y., Letunic, I., et al. (2008) · Nature · doi:10.1038/nature06617 · Cited by 1,139
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Late-Neoproterozoic Deep-Ocean Oxygenation and the Rise of Animal Life
Canfield, D. E., Poulton, S. W., & Narbonne, G. M. (2007) · Science · doi:10.1126/science.1135013 · Cited by 1,000 · cited 5×
“G evidence points to the general oxygenation of the oceans around 635 million years ago (Ma) (1), with an oxygenation of the deep oceans around 580 Ma, likely requiring a minimum of 10% present atmospheric levels (PAL) (2).”
BackgroundInfluentialSelf-citationMicroelectrodes: Their Use in Microbial Ecology
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Fossil steroids record the appearance of Demospongiae during the Cryogenian period
Love, G. D., Grosjean, E., Stalvies, C., Fike, D. A., Grotzinger, J. P., Bradley, A. S., Kelly, A. E., Bhatia, M., Meredith, W., Snape, C. E., et al. (2009) · Nature · doi:10.1038/nature07673 · Cited by 731 · cited 2×
“Indeed, according to present evidence, the earliest proposed signs of animals in the rock record (9, 10), as well as molecular clock divergence estimates (11, 12), predate the earliest evidence for deep ocean oxygenation (1), potentially by tens of millions of years.”
BackgroundThe Early Evolution of Eukaryotes: A Geological Perspective
Knoll, A. H. (1992) · Science · doi:10.1126/science.1585174 · Cited by 684
EVOLUTIONARY AND ECOLOGIC SIGNIFICANCE OF OXYGEN‐DEFICIENT MARINE BASINS
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The Multiple Origins of Complex Multicellularity
Knoll, A. H. (2011) · Annual Review of Earth and Planetary Sciences · doi:10.1146/annurev.earth.031208.100209 · Cited by 562
Flagged first, then by citations
Research works printed but not linked: 1
- N Boury-Esnault, L de Vos, C Donadey, J Vacelet Ultrastructure of Choanosome and Sponge Classification (Smithsonian Institution Press, Washington, DC), pp. 237–244 (1990).
Cited by
Sorted by citations
Items 201–300 of 354
On the use of models in understanding the rise of complex life
Lenton, T. M. (2020) · Interface Focus · doi:10.1098/rsfs.2020.0018 · Cited by 12
“Recently, it became apparent that oxygen levels would have to have been very low to have constrained the evolution or ecological expansion of sponges (the basal animals) [9] and several other (subsequently evolved) types of animal life [2].”
BackgroundAnimal survival strategies in Neoproterozoic ice worlds
Griffiths, H. J., Whittle, R. J., & Mitchell, E. G. (2022) · Global Change Biology · doi:10.1111/gcb.16393 · Cited by 14
“Early animals may also have been more tolerant of anoxic and hypoxic environments, requiring far lower oxygen levels than later taxa (Danovaro et al., 2010; Mills et al., 2014).”
BackgroundStrehlow, B. W., Schuster, A., Francis, W. R., et al. (2022) · BMC Research Notes · doi:10.1186/s13104-022-06013-3 · Cited by 12
Self-citationThe Ediacaran‐Cambrian Transition
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Wang, 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
Respiration kinetics and allometric scaling in the demosponge Halichondria panicea
Kumala, L., Thomsen, M., & Canfield, D. E. (2023) · BMC Ecology and Evolution · doi:10.1186/s12862-023-02163-5 · Cited by 12
“This question is not new (e.g. [108, 112]) but our results, combined with other physiological studies on the low-oxygen tolerance of sponges [48, 113], suggest that oxygen availability was not likely the reason.”
BackgroundResultSelf-citationPhysical constraints during Snowball Earth drive the evolution of multicellularity
Crockett, W. W., Shaw, J. O., Simpson, C., & Kempes, C. P. (2024) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2023.2767 · Cited by 13
“…primary productivity (NPP) as a function of temperature and intensity of photosynthetically active radiation (PAR) [28]: where V is the volume of water, n a is the number of autotrophic cells, e is the efficiency of production of organic matter and P i is the productivity of each autotrophic cell.”
BackgroundGrimes, C. J., Paiva, P. C., Petersen, L. H., et al. (2020) · Marine Biology · doi:10.1007/s00227-020-03756-0 · Cited by 11
Cole, D., Ozaki, K., & Reinhard, C. (2021) · Global Biogeochemical Cycles · doi:10.1029/2021gb007052 · Cited by 7
Wang, Y., Botting, J., Tan, J.-Q., Li, M., & Wang, W.-H. (2023) · Journal of Palaeogeography · doi:10.1016/j.jop.2023.03.005 · Cited by 9
A Bayesian Analysis of Technological Intelligence in Land and Oceans
Lingam, M., Balbi, A., & Mahajan, S. M. (2023) · The Astrophysical Journal · doi:10.3847/1538-4357/acb6fa · Cited by 11
“…(Cole et al. 2020; Lyons et al. 2021; Mills et al. 2022; Sperling et al. 2022); in particular, some animals comfortably survive under low-oxygen (e.g., Demospongiae) or completely anoxic (e.g., Henneguya salminicola) conditions (Danovaro et al. 2010; Mills et al. 2014; Yahalomi et al. 2020).”
BackgroundAuthigenic chromium enrichments in Proterozoic ironstones
Cole, D. B., O’Connell, B., & Planavsky, N. J. (2018) · Sedimentary Geology · doi:10.1016/j.sedgeo.2018.05.002 · Cited by 10
Early Prosperity of Iron Bacteria at the End of the Paleoproterozoic Era
Qiu, Y., Qin, L., Huang, F., Zhao, T., & Li, Y. (2022) · Geophysical Research Letters · doi:10.1029/2022gl097877 · Cited by 13
“5%–4% PAL) for the prosperity of complex life (Mills et al., 2014), but is the favorable condition for the thriving of FeOB (Chan et al.”
BackgroundDid high temperature rather than low O2 hinder the evolution of eukaryotes in the Precambrian?
Zhang, F., Wang, H., Ye, Y., Liu, Y., Lyu, Y., Deng, Y., Lyu, D., Wang, X., Wu, H., Deng, S., et al. (2022) · Precambrian Research · doi:10.1016/j.precamres.2022.106755 · Cited by 11
Red crust: evidence for an early Paleozoic oceanic anoxic event
Langsford, N. R., Raimondo, T., & JAGO, J. B. (2019) · Australian Journal of Earth Sciences · doi:10.1080/08120099.2018.1563827 · Cited by 10
Hu, R., Tan, J., Dick, J. M., Wang, Y., Li, G., & Liu, C. (2023) · Marine and Petroleum Geology · doi:10.1016/j.marpetgeo.2023.106307 · Cited by 9
Life in Changing Fluids: A Critical Appraisal of Swimming Animals Before the Cambrian
Gold, D. A. (2018) · Integrative and Comparative Biology · doi:10.1093/icb/icy015 · Cited by 9
“Many sponge species can grow and reproduce under extremely cold temperatures, low oxygen concentrations, and can subsist on a diet of dissolved organic matter (De Goeij et al. 2013; Mills et al. 2014; Sperling et al. 2015).”
BackgroundSouthern Ocean carbonate dissolution paced by Antarctic Ice-Sheet expansion in the early Miocene
Tagliaro, G. T., Fulthorpe, C. S., Watkins, D. K., Brumsack, H., & Jovane, L. (2021) · Global and Planetary Change · doi:10.1016/j.gloplacha.2021.103510 · Cited by 9
Actual and Model-Predicted Growth of Sponges—With a Bioenergetic Comparison to Other Filter-Feeders
Riisgård, H. U., & Larsen, P. S. (2022) · Journal of Marine Science and Engineering · doi:10.3390/jmse10050607 · Cited by 9
Meinhold, G., Willbold, M., Karius, V., Jensen, S., Agić, H., Ebbestad, J. O. R., Palacios, T., Högström, A. E. S., Høyberget, M., & Taylor, W. L. (2022) · Precambrian Research · doi:10.1016/j.precamres.2022.106845 · Cited by 9
“However, it has to be factored in that early metazoans – some found among the Ediacara biota – may have had a high tolerance to low oxygen concentrations under which they evolved, as do some modern animals (for instance, sponges; Mills et al., 2014).”
BackgroundA 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
“Geobiologists have theoretically ( 129 , 130 ) and experimentally ( 131 , 132 ) constrained the minimum oxygen requirements of animals to around 0.1 to 1% PAL O 2 ( 133 ), based primarily on the physiology of living animals.”
InfluentialSelf-citationOceanic Ba removal improved marine habitability for the oldest-known animals at ca. 600 Ma
Sui, P., Wei, W., Zhang, S., Zhao, Y.-Y., & Huang, F. (2025) · Global and Planetary Change · doi:10.1016/j.gloplacha.2025.104790 · Cited by 8
Fu, F., Zou, C., Zhao, Z., Pan, S., Wang, W., Luo, C., Hua, G., Lu, G., Yuan, M., Yin, J., et al. (2025) · Marine and Petroleum Geology · doi:10.1016/j.marpetgeo.2025.107438 · Cited by 8
Fru, E. C., Aubineau, J., Bankole, O. M., Ghnahalla, M., Tamehe, L. S., & Albani, A. E. (2024) · Precambrian Research · doi:10.1016/j.precamres.2024.107453 · Cited by 9
Bezerra, B. S., Belato, F. A., Mello, B., Brown, F. D., Coates, C. J., Leme, J. D. M., Trindade, R. I. F., & Costa-Paiva, E. M. (2021) · Scientific Reports · doi:10.1038/s41598-021-95094-4 · Cited by 8
Molecular dating of the blood pigment hemocyanin provides new insight into the origin of animals
Costa-Paiva, E. M., Mello, B., Bezerra, B. S., Coates, C. J., Halanych, K. M., Brown, F. D., Leme, J. D. M., & Trindade, R. I. F. (2021) · Geobiology · doi:10.1111/gbi.12481 · Cited by 8
Geological evidences and mechanisms for oceanic anoxic events during the Early Paleozoic
Zhang, J., Li, C., & Zhang, Y. (2021) · Chinese Science Bulletin (Chinese Version) · doi:10.1360/tb-2021-0535 · Cited by 8
Deng, T., Li, Y., & Hu, X. (2022) · Journal of Asian Earth Sciences · doi:10.1016/j.jseaes.2022.105143 · Cited by 8
Breathing room for early animals
Fischer, W. W. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1525100113 · Cited by 8
Palaeoecology of Ediacaran metazoan reefs
Wood, R. A. (2016) · Geological Society London Special Publications · doi:10.1144/sp448.1 · Cited by 8
“Experimental work has also shown that the oxygen levels necessary to support small, primitive metazoans such as sponges (Porifera) are far lower (Mills et al., 2014) than those required for large, active, and ecologically-important animals such as carnivorous predators (Sperling et al., 2013).”
BackgroundRytkönen, K. T. (2018) · eLife · doi:10.7554/elife.34756 · Cited by 8
“Both phyla seem to survive extended periods of low oxygen, whereas the placozoans do not (Mills et al., 2014; Purcell et al., 2001; Loenarz et al., 2011).”
BackgroundEarth: Atmospheric Evolution of a Habitable Planet
Olson, S. L., Schwieterman, E. W., Reinhard, C. T., & Lyons, T. W. (2018) · Springer eBooks · doi:10.1007/978-3-319-55333-7_189 · Cited by 24
“…weathering at a particular temperature has necessarily, but not straightforwardly, changed through time as the balance of seafloor and continental weathering has evolved and as Earth’s subaerially exposed crust has become colonized by progressively more complex ecosystems (B Mills et al 2014).”
BackgroundInfluentialMa, Q., Feng, Q., Ye, Y., & Shen, J. (2020) · Palaeogeography Palaeoclimatology Palaeoecology · doi:10.1016/j.palaeo.2020.110193 · Cited by 8
Coupling of shallow marine redox change and multi-stage animal radiations during the early Cambrian
Zheng, Y., Chang, C., Li, J., Ding, Y., Tang, X., & Liu, J. (2025) · Gondwana Research · doi:10.1016/j.gr.2025.01.003 · Cited by 6
Sub-millimeter molybdenum and uranium isotopes track millennial redox events in the Cambrian ocean
Zhao, Z., Hougård, I. W., Zou, C., Dickson, A. J., Dai, S., Jing, Z., Nielsen, A. T., Ju, P., Guo, Z., Basu, A., et al. (2025) · Communications Earth & Environment · doi:10.1038/s43247-025-02722-2 · Cited by 7
“In addition, the oxygen demands of Cambrian organisms were likely lower 70 .”
Zheng, Y., Tang, Q., Liu, P., Sun, J., Yao, X., Hao, W., Yang, X., Guo, J., Uesugi, K., Komiya, T., et al. (2021) · Journal of Geophysical Research Biogeosciences · doi:10.1029/2020jg006102 · Cited by 8
“Despite the causal relationship with the three-episode Cambrian explosion was still controversial (Butterfield, 2009; X. Chen et al., 2015; He et al., 2019; Knoll & Carroll, 1999; Li et al., 2017; Mills et al., 2014), the stepwise oxygenation of ocean in this period was widely accepted (Dong et al.”
Background张, 水., 王, 华., 王, 晓., & 叶, 云. (2021) · SCIENTIA SINICA Terrae · doi:10.1360/sste-2020-0373 · Cited by 6
Non-zero I/(Ca + Mg) recorded in Archean and Paleoproterozoic shallow marine Ca-carbonate sediments
Hao, F., Fralick, P. W., Ramsay, B., Tang, D., & Riding, R. (2024) · Precambrian Research · doi:10.1016/j.precamres.2024.107350 · Cited by 7
A billion years of geological drama – Boring or brilliant?
Mukherjee, I., Corkrey, R., Gregory, D. D., Large, R. R., & Poole, A. M. (2025) · Gondwana Research · doi:10.1016/j.gr.2025.02.018 · Cited by 8
Pei, Y., Duda, J., & Reitner, J. (2020) · Paläontologische Zeitschrift · doi:10.1007/s12542-020-00530-x · Cited by 7
Characteristics of aquatic biospheres on temperate planets around Sun-like stars and M dwarfs
Lingam, M., & Loeb, A. (2021) · Monthly Notices of the Royal Astronomical Society · doi:10.1093/mnras/stab611 · Cited by 7
Taphonomy of exceptional sponge preservation in the Hirnantian Anji Biota, China
Wu, X., Botting, J. P., Zhang, Y., Muir, L. A., & Ma, J. (2022) · Journal of the Geological Society · doi:10.1144/jgs2021-112 · Cited by 7
Li, S., Lang, X., Wang, H., Zhu, S., Zhao, K., & Hou, M. (2023) · Minerals · doi:10.3390/min13020227 · Cited by 7
New keratose sponges after the end‐Permian extinction provide insights into biotic recoveries
Wu, S., Joachim Reitner, A., Harper, D. A. T., Yu, J., & Chen, Z. (2023) · Geobiology · doi:10.1111/gbi.12582 · Cited by 7
“In fact, some modern dem-osponges can survive under low-oxygen conditions (Hoffmann et al., 2005; Mills et al., 2014) and the earliest crown-group sponges had already evolved a great tolerance of oxygen-deficient habitats (Mills et al., 2014).”
BackgroundMurphy, A., Penny, A., Zhuravlev, A. Y., & Wood, R. A. (2025) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2025.0968 · Cited by 6
“This pattern is consistent with the observation that some modern siliceous demosponges are able to tolerate remarkably low concentrations of dissolved oxygen [90].”
Chanchai, W., Nelson, L. L., Smith, E. F., Hardisty, D. S., Lonsdale, M. C., Burke, J. E., & Lau, K. (2025) · Geobiology · doi:10.1111/gbi.70039 · Cited by 6
“Although early animals may have been tolerant of low-oxygen conditions (Mills et al. 2014), dynamic redox conditions could have led to biological innovation by shaping behavioral complexity, physiological stress due to anoxia, and competition pressure from co-habiting species (e.g., Childress and…”
Ai, Y., Zhu, G., Li, T., & Zhu, Z. (2023) · Applied Geochemistry · doi:10.1016/j.apgeochem.2023.105588 · Cited by 7
Yun, H., Janussen, D., Zhang, X., & Joachim Reitner, A. (2023) · Lethaia · doi:10.18261/let.56.3.5 · Cited by 5
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
“In modern environments, nonquantitative respiration of Se oxyanions is observed at micromolar dissolved oxygen levels or higher (24), similar to thresholds for animal survival (25), making Se a useful proxy for tracking oxygen levels relevant to animal respiration.”
Fraga-Ferreira, P., Ader, M., Caetano-Filho, S., Sansjofre, P., Paula-Santos, G., Babinski, M., Guacaneme, C., Bedoya-Rueda, C., Rojas, V., Reis, H., et al. (2021) · Frontiers in Earth Science · doi:10.3389/feart.2021.692895 · Cited by 6
Therapeutic hypercapnia. Review
Tolstun, D. A., Muradian, K. K., & Bezrukov, V. V. (2022) · Ageing & Longevity · doi:10.47855/al9020-2022-3-5 · Cited by 5
“After all, life originated, and most of the evolution took place in atmospheres with an extremely low content of O 2 and a high content of CO 2 , while in the modern atmosphere, on the contrary, the content of O 2 is more than 500 times higher than the content of CO 2 [69–71].”
He, Y., Zhu, X., Qiu, Y., Pang, L., & Zhao, T. (2022) · Precambrian Research · doi:10.1016/j.precamres.2022.106901 · Cited by 5
Cordie, D. R. (2023) · Evolving Earth · doi:10.1016/j.eve.2023.100002 · Cited by 5
Costa-Paiva, E. M., & Coates, C. J. (2020) · Sub-cellular biochemistry/Subcellular biochemistry · doi:10.1007/978-3-030-41769-7_10 · Cited by 5
Earth: Atmospheric Evolution of a Habitable Planet
Olson, S. L., Schwieterman, E. W., Reinhard, C. T., & Lyons, T. W. (2018) · Springer eBooks · doi:10.1007/978-3-319-30648-3_189-1 · Cited by 5
Exploiting the hypoxia sensitive non-coding genome for organ-specific physiologic reprogramming
Bischof, C., & Krishnan, J. (2016) · Biochimica et Biophysica Acta (BBA) - Molecular Cell Research · doi:10.1016/j.bbamcr.2016.01.024 · Cited by 5
Feng, T., Abbatiello, J., Omran, A. P., Mehta, C., & Pasek, M. A. (2021) · Minerals · doi:10.3390/min11121394 · Cited by 5
A billion years of temperature variability: a key driver of Earth’s long-term habitabilityPreprint
Bergmann, K., Boekelheide, N., Clarke, J. W., Cantine, M. D., Wilcots, J., Anderson, N. T., Jost, A. B., Laub, O., Drozd, J. K., Goldberg, S. L., et al. (2022) · doi:10.1002/essoar.10511918.1 · Cited by 5
Tan, Y., Meng, G., Feng, Y., Liu, W., Wang, Q., Gao, P., & Xiao, X. (2025) · Journal of Marine Science and Engineering · doi:10.3390/jmse13061175 · Cited by 5
Ye, Y., Zhang, L., Chen, C., Chang, S., Yi, Y., & Feng, Q. (2025) · Science China Earth Sciences · doi:10.1007/s11430-024-1468-9 · Cited by 5
“Regarding sponges, studies have shown that they can survive under very low oxygen conditions, indicating a very low demand for oxygen (Mills et al., 2014, 2018; Leys and Kahn, 2018).”
Reexamination and reidentification of ocean oxygenation event in the wake of the Marinoan glaciation
Li, R., Zhou, X., Guo, C., Huang, T., Wang, Z., Huang, K., Peng, P., Wang, X., Du, T., Xue, D., et al. (2025) · Earth and Planetary Science Letters · doi:10.1016/j.epsl.2025.119312 · Cited by 5
Marine sponges in the once and future ocean
Mills, D. B., & Sperling, E. A. (2021) · Global Change Biology · doi:10.1111/gcb.16053 · Cited by 4
“Traditional p O 2 estimates for the mid-Proterozoic Eon (2.5– 0.541 billion years ago)— 1– 10% PAL— are shown in yellow (Mills et al., 2014).”
InfluentialSelf-citationAl‐Haddad, S., Caldwell, G. S., & Clare, A. S. (2024) · Aquatic Sciences · doi:10.1007/s00027-024-01132-4 · Cited by 4
“It is, therefore, a widely studied species in terms of its biology and ecophysiology (Mills et al. 2014; Riisgård et al. 2016).”
Neoproterozoic copper cycling, and the rise of metazoans
Parnell, J., & Boyce, A. J. (2019) · Scientific Reports · doi:10.1038/s41598-019-40484-y · Cited by 4
Garcia, A. K. (2019) · Precambrian Research · doi:10.1016/j.precamres.2019.105389 · Cited by 4
Application of the apatite oxygen paleobarometer (AOP) across the Neoproterozoic-Cambrian transition
Schopf, J. W., & Garcia, A. K. (2019) · Precambrian Research · doi:10.1016/j.precamres.2019.105404 · Cited by 4
Biotic vs abiotic origin of unusual features from Mesoproterozoic of Vindhyan Supergroup, India
Choudhuri, A., Albani, A. E., Mandal, S., & Sarkar, S. (2023) · Annales de Paléontologie · doi:10.1016/j.annpal.2023.102629 · Cited by 4
Belato, F. A., Mello, B., Coates, C. J., Halanych, K. M., Brown, F. D., Morandini, A. C., Leme, J. D. M., Trindade, R. I. F., & Costa-Paiva, E. M. (2023) · Geobiology · doi:10.1111/gbi.12577 · Cited by 4
“In support of this hypothe - sis, recent works have proposed that early animals were likely small, soft-bodied, collagen-poor, and restricted their use of oxygen to high-priority physiological functions (Cole et al., 2020 ; Mills et al., 2014 ).”
BackgroundInfluentialTranscriptomic responses of sponge holobionts to in situ, seasonal anoxia and hypoxia
Strehlow, B. W., Schuster, A., Francis, W. R., Eckford-Soper, L., Kraft, B., McAllen, R., Nielsen, R., Mandrup, S., & Canfield, D. E. (2024) · Peer Community Journal · doi:10.24072/pcjournal.410 · Cited by 4
“Indeed, these limits are above those at which mortality or substantial physiological changes have been observed for sponges (Micaroni et al. 2021b; Kumala et al. 2021; Mills et al. 2014).”
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Sponges as sensitive animals: sensory systems and energetics of filtration in demosponges
Ludeman, D. A. (2015) · ERA: Education and Research Archive (University of Alberta) · doi:10.7939/r3rx93n3m · Cited by 4
Mángano, M. G., & Buatois, L. A. (2020) · Elsevier eBooks · doi:10.1016/b978-0-08-102908-4.00142-9 · Cited by 4
Hou, A., Shi, X., Guilbaud, R., Xie, B., Sun, L., Xu, L., Zhou, L., Liu, W., Zhou, X., Wang, X., et al. (2025) · Journal of Geophysical Research Biogeosciences · doi:10.1029/2025jg008835 · Cited by 3
“Aerobic respiration is a fundamental characteristic of eukaryotes, and oxygen concentration is a key factor influencing their evolution (Mills et al., 2014; Runnegar, 1991; Stolper et al., 2010).”
Graziano, R., Raspini, A., & Bartiromo, A. (2023) · Sedimentology · doi:10.1111/sed.13146 · Cited by 3
Venugopal, A., Tripathy, G. R., Goswami, V., Khan, T., & Ackerman, L. (2025) · Geochimica et Cosmochimica Acta · doi:10.1016/j.gca.2025.09.013 · Cited by 3
Li, D., Luo, G., Shen, B., & Xiao, S. (2025) · Precambrian Research · doi:10.1016/j.precamres.2025.107960 · Cited by 3
Mesoproterozoic calcareous paleosols from Montana
Retallack, G. J. (2023) · Precambrian Research · doi:10.1016/j.precamres.2023.107134 · Cited by 3
Energy costs of the sponge pump
Riisgård, H. U., & Larsen, P. S. (2024) · Marine Ecology Progress Series · doi:10.3354/meps14684 · Cited by 3
Atmosphere, Metabolism and Longevity
Muradian, K. K. (2015) · Healthy ageing and longevity · doi:10.1007/978-3-319-18326-8_13 · Cited by 2
Carbon and sulfur biogeochemical cycles in the Brazilian Ediacaran record
Filho, S. C. (2020) · Digital Library of Theses and Dissertations (Universidade de São Paulo) · doi:10.11606/t.44.2020.tde-23122020-125734 · Cited by 4
Pei, Y. (2022) · eDiss (Georg-August-Universität Göttingen) · doi:10.53846/goediss-9160 · Cited by 3
Experimental Snowball Earth Viscosity Drives the Evolution of Motile MulticellularityPreprint
Halling, A., Goodson, B., Hirschmann, A., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.02.06.579218 · Cited by 3
Park, J. G., Hudgins, M., Fralick, P. W., Shelley, J. T., & Schaller, M. F. (2025) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2513030122 · Cited by 3
“Mesoproterozoic oxygen concentrations at least transiently exceeded the threshold necessary for early respiration and a UV - shielding ozone layer (6, 44).”
Noncanonical genetic markers resolve the pre-GOE emergence of aerobic bacteria in Earth’s history
Liao, T., Chen, S., Wang, S., Huang, Y. J. W., Tsui, S. K., Stüeken, E. E., Cao, Q., & Luo, H. (2026) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2515709123 · Cited by 2
“Te ecological impact of the NOE in particular is sometimes questioned, because the geochemical proxy record as well as O 2 requirements of early animals remain debated (61, 62).”
Revisiting the greatness of Earth’s great oxidation
Crockford, P. W., Sugiyama, I., Kipp, M. A., Hao, J., Nelson, L. L., Hemingway, J., Wimmer, S., & Fakhraee, M. (2026) · Communications Earth & Environment · doi:10.1038/s43247-026-03518-8 · Cited by 4
“Plausible marine oxygen levels through the GOE also span ranges above and below the thresholds for eukaryotic or even metazoan aerobic metabolism 55,56 , leaving it open for debate whether the later Paleoproter-ozoic appearance of eukaryotic microfossils re fl ects an immediate response to…”
Zhang, Z., Cheng, M., Wang, H., Chen, X., Guo, W., & Li, C. (2023) · Science China Earth Sciences · doi:10.1007/s11430-022-1116-3 · Cited by 2
“The reasons are: first, early animals such as sponges had extremely low oxygen demands, possibly as low as 0.5% PAL (Mills et al., 2014), which cannot be detected with the precision of I/(Ca+Mg) ratio; second, the response of early animals to oxygen was nonlinear, and an increase in oxygen within a small range may hav…”
BackgroundInfluentialOxygenation of the Proterozoic Earth's surface: An evolving story
Kendall, B., & Ostrander, C. M. (2024) · Elsevier eBooks · doi:10.1016/b978-0-323-99762-1.00058-9 · Cited by 3
Yang, H., Zhou, J., Ma, J., & Cui, X. (2025) · Organic Geochemistry · doi:10.1016/j.orggeochem.2025.104950 · Cited by 2
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
“Asnotedabove,thelossofMIF-Shasbeenusedasametricforde fi ning the timing of the GOE for over two decades 7 and constrains the point at whichatmosphericoxygenroseacrossthe10 − 6 PALthreshold 10,11 .”
InfluentialSegmentation and oxygen diffusion in the Ediacaran Dickinsonia: an applied analysis
Gooden, B. A. (2014) · Journal and proceedings of the Royal Society of New South Wales · doi:10.5962/p.361716 · Cited by 2
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Montalvão, S. I. G. H. M. (2016) · Työväentutkimus Vuosikirja · Cited by 2
Evolution of Eukaryotes with Respect to Atmosphere Oxygen Appearance and Rise
Ivanović, Z., & Vlaski‐Lafarge, M. (2015) · Elsevier eBooks · doi:10.1016/b978-0-12-800540-8.00008-9 · Cited by 1
Cao, H., Wang, Z., Chen, F., Wei, K., Huang, H., Chen, Q., & Chen, C. (2025) · Global and Planetary Change · doi:10.1016/j.gloplacha.2025.104958 · Cited by 1
Subarkah, D., Blades, M., Collins, A. S., Farkaš, J., & Virgo, G. M. (2025) · Australian Journal of Earth Sciences · doi:10.1080/08120099.2025.2559364 · Cited by 1
“This level of oxygen has been suggested to be sufficient for fuelling early evolved animal clades (Mills et al ., 2014; Zhang et al ., 2016).”
Choudhuri, A., Plummer, P. S., Ghnahalla, M., Albani, A. E., Sarkar, S., Bouougri, E. H., Reisberg, L. C., Kumar, A., & Bose, T. (2025) · Australian Journal of Earth Sciences · doi:10.1080/08120099.2025.2586733 · Cited by 1
“…ancestors of motile eukaryotes could survive at oxygen concentrations as low as 0.5 to 4%, and that bilaterian animals can sustain 3% oxygen concentration in modern ocean surface waters (Knoll & Sperling, 2014; Levin et al ., 2002; Mills et al ., 2014; Palma et al ., 2005; Sperling et al ., 2013).”
InfluentialHolland, 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
Evolutionary Origins of Stemness
Ivanović, Z., & Vlaski‐Lafarge, M. (2015) · Elsevier eBooks · doi:10.1016/b978-0-12-800540-8.00010-7 · Cited by 1
Metadata
- Authors
- Mills, D. B., Ward, L. M., Jones, C., Sweeten, B., Forth, M., Treusch, A. H., & Canfield, D. E.
- Affiliations
- University of Southern Denmark (DK)
- California Institute of Technology (US)
- University of British Columbia (CA)
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2014-02-18
- Volume
- 111
- Issue
- 11
- Pages
- 4168-4172
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC3964089
- OpenAlex
- W2029322624
- Semantic Scholar
- ac41b42cf0cc457afe8379a4d86261edcc53d9e5
- Cited by
- 355
- References
- 53
- Citations by year
- 2026: 15 · 2025: 29 · 2024: 24 · 2023: 24 · 2022: 39 · 2021: 40 · 2020: 29 · 2019: 29 · 2018: 33 · 2017: 23 · 2016: 27 · 2015: 32 · 2014: 11
- FWCI
- 22.57
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- University of Southern Denmark Research Portal (University of Southern Denmark) (submittedVersion)
- Funders
- Agouron Institute
- National Research Foundation
- Danmarks Grundforskningsfond (Grant DNRF 53, DNRF 53)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Stratigraphy of Fossils
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Physical Sciences › Physics and Astronomy › Astronomy and Astrophysics › Origins and Evolution of Life
- Keywords
- origin of animals, Ediacaran, oxygen requirement, demosponges, low oxygen tolerance, atmospheric oxygenation, Neoproterozoic, animal evolution
- MeSH terms
- Adaptation, Biological, Animals, Atmosphere, Base Sequence, Denmark, Biological Evolution, Kinetics, Molecular Sequence Data, Oxygen, Oxygen Consumption, Porifera, Polymorphism, Restriction Fragment Length, RNA, Ribosomal, 16S, Sequence Analysis, DNA, Microbiota
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
A language model drew the claims from the paper on 2026-10-11, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-10-11 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Cited sources are checked with a language model’s help (gpt-6-sol).
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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