Vol. 53 No. 1 (2026)
Column

The Tooth of Time: Charles Darwin’s Prophecy Regarding Selective Survival on Snowball Earth

Paul F. Hoffman
1216 Montrose Avenue, Victoria, British Columbia, V8T 2K4, Canada

Published 2026-03-16

Keywords

  • Charles Darwin,
  • Cryogenian,
  • Ecosystem relocation,
  • Glacier biomes,
  • Mass extinction,
  • Microbial mats,
  • Selective survival,
  • Snowball Earth
  • ...More
    Less

How to Cite

Hoffman, P. F. (2026). The Tooth of Time: Charles Darwin’s Prophecy Regarding Selective Survival on Snowball Earth. Geoscience Canada, 53(1), 1–10. https://doi.org/10.12789/geocanj.2026.53.227

References

  1. Abbot, D.S., 2014, Resolved Snowball Earth clouds: Journal of Climate, v. 27, p. 4391−4402, https://doi.org/10.1175/JCLI-D-13-00738.1.
  2. Abbot, D.S., Voigt, A., Li, D.W., Le Hir, G., Pierrehumbert, R.T., Branson, M., Pollard, D., and Koll, D.D.B., 2013, Robust elements of Snowball Earth atmospheric circulation and oases for life: Journal of Geophysical Research: Atmospheres, v. 118, p. 6017−6027, https://doi.org/10.1002/jgrd.50540.
  3. Agassiz, L., 1967, Studies on Glaciers, Preceded by the Discourse of Neuchâtel, 1837: Translated and edited by A. Carozzi, Hafner, New York, 213 p., 18 plates.
  4. Alroy, J., 2001, A multispecies overkill simulation of the end-Pleistocene megafaunal mass extinction: Science, v. 292, p. 1893−1896, https://doi.org/10.1126/science.1059342.
  5. Anesio, A.M., Hodson, A.J., Fritz, A., Psenner, R., and Sattler, B., 2009, High microbial activity on glaciers: importance to the global carbon cycle: Global Change Biology, v. 15, p. 955–960, https://doi.org/10.1111/j.1365-2486.2008.01758.x.
  6. Bao, H.M., Lyons, J.R., and Zhou, C.M., 2008, Triple oxygen isotope evidence for elevated CO2 levels after a Neoproterozoic glaciation: Nature, v. 453, p. 504−506, https://doi.org/10.1038/nature06959.
  7. Bar-On, Y.M., Phillips, R., and Milo, R., 2018, The biomass distribution on Earth: Proceedings of the National Academy of Sciences (USA), v. 115, p. 6506−6511, https://doi.org/10.1073/pnas.1711842115.
  8. Barrett, P.H., 1973, Darwin's 'gigantic blunder': Journal of Geological Education, v. 21, p. 19−28, https://doi.org/10.5408/0022-1368-21.1.19.
  9. Bernhardi, A., 1832, Wie kamen die aus den Norden stammenden Felsbruchstücke und Geschiebe, welche man in Nordeutschland und den benachbarten Ländern findet, an ihre gegenwärtigen Fundorte?: Jahrbuch für Mineralogie, Geognosie, und Petrafaktenkunde, v. 3, p. 257−267.
  10. Bernhardi, A., 1939, How did the rock fragments and boulders of northern origin found in northern Germany and neighbouring countries get to their present positions?, in Mather, K.F., and Mason, S.L., eds., A Sourcebook in Geology: McGraw-Hill, New Yok, NY, p. 327−328 (English translation of Bernhardi 1832).
  11. Beulig, F., Schubert, F., Adhikari, R.R., Glombitza, C., Heuer, V.B., Hinrichs, K.-U., Homola, K.L., Inagaki, F., Jørgensen, B.B., Kallmeyer, J., Krause, S.J.E., Morono, Y., Sauvage, J., Spivak, A.J., and Treude, T., 2022, Rapid metabolism fosters microbial survival in the deep, hot subseafloor biosphere: Nature Communications, v. 13, 312, https://doi.org/10.1038/s41467-021-27802-7.
  12. Bo, S., Siegert, M.J., Mudd, S.M., Sugden, D., Fujita, S., Cui, X.B., Jiang, Y.Y., Tang, X.Y., and Li, Y.S,, 2009, The Gamburtsev mountains and the origin and early evolution of the Antarctic Ice Sheet: Nature, v. 459, p. 690−693, https://doi.org/10.1038/nature08024.
  13. Bøggild, C.E., Brandt, R.E., Brown, K.J., and Warren, S.G., 2010, The ablation zone in northeast Greenland: ice types, albedos and impurities: Journal of Glaciology, v. 56, p. 101−113, https://doi.org/10.3189/002214310791190776.
  14. Boulton, G.S., 1992, Quaternary, in Duff, P.McL.D., and Smith, A.J., eds., Geology of England and Wales: Geological Society of London, UK, p. 413−444.
  15. Boyce, C.K., and Nelsen, M.P., 2025, Terrestrialization: toward a shared framework for ecosystem evolution: Paleobiology, v. 51, p. 174−194, https://doi.org/10.1017/pab.2024.15.
  16. Brady, P.V., and Gíslason, S.R., 1997, Seafloor weathering controls on atmospheric CO2 and global climate: Geochimica et Cosmochimica Acta, v. 61, p. 965−973, https://doi.org/10.1016/S0016-7037(96)00385-7.
  17. Brown, M., Johnson, T., and Spencer, C.J., 2022, Secular changes in metamorphism and metamorphic cooling rates track the evolving plate-tectonic regime on Earth: Journal of the Geological Society, London, v. 179, jgs2022-050, https://doi.org/10.1144/jgs2022-050.
  18. Browne, J., 1995, Charles Darwin: Voyaging: Princeton University Press, Princeton, New Jersey, 605 p.
  19. Buckland, W., 1842, On Diluvio-Glacial Phaenomena in Snowdonia and the adjacent parts of North Wales: Proceedings of the Geological Society, London, v. 3, no. 2, p. 579−584.
  20. Campbell, A.J., Waddington, E.D., and Warren, S.G., 2011, Refugium for surface life on Snowball Earth in a nearly-enclosed sea? A first simple model for sea-glacier invasion: Geophysical Research Letters, v. 38, L19502, https://doi.org/10.1029/2011GL048846.
  21. Campbell, A.J., Waddington, E.D., and Warren, S.G., 2014, Refugium for surface life in a nearly enclosed sea? A numerical solution for sea-glacier invasion through a narrow strait: Journal of Geophysical Research: Oceans, v. 119, p. 2679−2690, https://doi.org/10.1002/2013JC009703.
  22. Campbell, A.J., Massarano, B., Waddington, E.D., and Warren, S.G., 2017, Could promontories have restricted sea-glacier penetration into marine embayments during Snowball Earth events?: The Cryosphere, v. 11, p. 1141−1148, https://doi.org/10.5194/tc-11-1141-2017.
  23. Campbell, S., and Bowen, D.Q., 1989, The Quaternary of Wales: Geological Conservation Review Series, No. 2, JNCC, Peterborough, ISBN 0 86139 570 0.
  24. Cao, X.B., and Bao, H.M., 2013, Dynamic model constraints on oxygen-17 depletion in atmospheric O2 after a snowball Earth: Proceedings of the National Academy of Sciences (USA), v. 110, p. 14546−14550, https://doi.org/10.1073/pnas.1302972110.
  25. Carozzi, A.V., 1966, Agassiz's amazing geological speculation: the Ice Age: Studies in Romanticism, v. 5, p. 57−83, https://doi.org/10.2307/25599657.
  26. Charpentier, J. de, 1837, Some conjectures regarding the great revolutions which have changed the surface of Switzerland, and particularly that of the Canton of Vaud, as to give rise to its present aspect: Edinburgh New Philosophical Journal, v. 22, p. 27−36.
  27. Charpentier, J. de, 1841, Essai sur les Glaciers et sur le Terrain Érratique du Bassin du Rhône: Ducloux, Lausanne, 363 p.
  28. Chatters, J.C., Potter, B.A., Fiedel, S.J., Morrow, J.E., Jass, C.N., and Wooller, M.J., 2024, Mammoth featured heavily in Western Clovis diet: Science Advances, v. 10, eadr3814, https://doi.org/10.1126/sciadv.adr3814.
  29. Clarke, A., Morris, G.J., Fonseca, F., Murray, B.J., Acton, E., and Price, H.C., 2013, A low temperature limit for life on Earth: PLoS ONE, v. 8, e66207, https://doi.org/10.1371/journal.pone.0066207.
  30. Coffey, N.B., MacAyeal, D.R., Copland, L., Mueller, D.R., Sergienko, O.V., Banwell, A.F., and Lai, C.Y., 2022, Enigmatic surface rolls of the Ellesmere Ice Shelf: Journal of Glaciology, v. 68, p. 867−878, https://doi.org/10.1017/jog.2022.3.
  31. Colman, D.R., Poudel, S., Stamps, B.W., Boyd, E.S., and Spear, J.R., 2017, The deep, hot biosphere: twenty-five years of retrospection: Proceedings of the National Academy of Sciences (USA), v. 114, p. 6895−6903, https://doi.org/10.1073/pnas.1701266114.
  32. Coogan, L.A., and Gillis, K.M., 2018, Low-temperature alteration of the seafloor: impacts on ocean chemistry: Annual Review of Earth and Planetary Sciences, v. 46, p. 21−45, https://doi.org/10.1146/annurev-earth-082517-010027.
  33. Cooper, A., Turney, C., Hughen, K.A., Brook, B.W., McDonald, H.G., and Bradshaw, C.J.A., 2015, Abrupt warming events drove Late Pleistocene Holarctic megafaunal turnover: Science, v. 349, p. 602−606, https://doi.org/10.1126/science.aac4315.
  34. Cox, S.E., Thomson, S.N., Reiners, P.W., Hemming, S.R., and van de Flierdt, T., 2010, Extremely low long-term erosion rates around the Gamburtsev Mountains in interior East Antarctica: Geophysical Research Letters, v. 37, L22307, https://doi.org/10.1029/2010GL045106.
  35. Creyts, T.T., Ferraccioli, F., Bell, R.E., Wolovick, M., Corr, H., Rose, K.C., Frearson, N., Damaske, D., Jordan, T., Braaten, D., and Finn, C., 2014, Freezing of ridges and water networks preserves the Gamburtsev Subglacial Mountains for millions of years: Geophysical Research Letters, v. 41, p. 8114–8122, https://doi.org/10.1002/2014GL061491.
  36. Darwin, C., 1839a, Note on a rock seen on an iceberg in 61° South latitude: The Journal of the Royal Geographical Society of London, v. 9, p. 528−529, https://doi.org/10.2307/1797747.
  37. Darwin, C., 1839b, IV. Observations on the parallel roads of Glen Roy, and of other parts of Lochaber in Scotland, with an attempt to prove that they are of marine origin: Philosophical Transactions of the Royal Society, London, v. 139, p. 39−81, https://doi.org/10.1098/rstl.1839.0005.
  38. Darwin, C., 1840, Journal of Researches into the Geology and Natural History of the Various Countries Visited by H.M.S. Beagle, Under the Command of Captain Fitzroy, R.N. from 1832 to 1836: Henry Colburn, London, 637 p.
  39. Darwin, C., 1841, XXVII. —On the distribution of the erratic boulders and on the contemporaneous unstratified deposits of South America: Transactions of the Geological Society of London, v. 6, p. 415−431, https://doi.org/10.1144/transgslb.6.2.415.
  40. Darwin, C., 1842, Notes on the effects produced by the ancient glaciers of Caernarvonshire, and on the boulders transported by floating ice: Edinburgh New Philosophical Journal, v. 33, p. 352−363.
  41. Darwin, C., 1848, On the transport of erratic boulders from a lower to a higher level: Quarterly Journal of the Geological Society, v. 4, p. 315−323, https://doi.org/10.1144/GSL.JGS.1848.004.01-02.44.
  42. Darwin, C., 1849, Geology, in Herschel, J.F.W., ed., A Manual of Scientific Enquiry; Prepared for the Use of Her Majesty's Navy: and Adapted for Travellers in General: John Murray, London, p. 156−195, https://doi.org/10.1017/CBO9780511996580.008.
  43. Darwin, C., 1855, XII. On the power of icebergs to make rectilinear, uniformly-directed grooves across a submarine undulatory surface: The London, Edinburgh, and Dublin Philosophical Magazine and Journal of Science, v. 10, p. 96−98, https://doi.org/10.1080/14786445508641938.
  44. Darwin, C., 1964, On the Origin of Species: A Facsimile of the First 1859 Edition, with an Introduction by Ernst Mayr: Harvard University Press, Cambridge, MA, 513 p.
  45. Davies, G.L., 1968, The tour of the British Isles made by Louis Agassiz in 1840: Annals of Science, v. 24, p. 131−146, https://doi.org/10.1080/00033796800200101.
  46. de Vrese, P., Stacke, T., Rugenstein, J.C., Goodman, J., and Brovkin, V., 2021, Snowfall-albedo feedbacks could have led to deglaciation of snowball Earth starting from mid-latitudes: Nature Communications Earth & Environment, v. 2, 91, https://doi.org/10.1038/s43247-021-00160-4.
  47. Del Cortona, A., Jackson, C.J., Bucchini, F., Van Bel, M., D'hondt, S., Škaloud, P., Delwiche, C.F., Knoll, A.H., Raven, J.A., Verbruggen, H., Vendepoele, K., De Clerck, O., and Leliaert, F., 2020, Neoproterozoic origin and multiple transitions to macroscopic growth in green seaweeds: Proceedings of the National Academy of Sciences (USA), v. 117, p. 2551−2559, https://doi.org/10.1073/pnas.1910060117.
  48. Dutkiewicz, A., Merdith, A.S., Collins, A.S., Mather, B., Llano, L., Zahirovic, S., and Müller, R.D., 2024, Duration of Sturtian "Snowball Earth" glaciation linked to exceptionally low mid-ocean ridge outgassing: Geology, v. 52, p. 292−296, https://doi.org/10.1130/G51669.1.
  49. Esmark, J., 1824, Bidrag til vor jordlkodes historie: Magazin for Naturvidenskaberne, v. 2, no. 1, p. 28−49.
  50. Esmark, J., 1826, Remarks tending to explain the geological history of the Earth: The Edinburgh New Philosophical Journal, v. 2, p. 107−121.
  51. Evans, D.A.D., 2021, Meso−Neoproterozoic Rodinia supercycle, in Pesonen, L.J., Salminen, J., Elming, S.-Å., Evans, D.A.D., and Veikkolainen, T., eds., Ancient Supercontinents and the Paleogeography of Earth: Elsevier, Amsterdam, p. 549−576, https://doi.org/10.1016/B978-0-12-818533-9.00006-0.
  52. Feistel, R., and Marion, G.M., 2007, A Gibbs−Pitzer function for high-salinity seawater thermodynamics: Progress in Oceanography, v. 74, p. 515−539, https://doi.org/10.1016/j.pocean.2007.04.020.
  53. Ferraccioli, F., Finn, C.A., Jordan, T.A., Bell, R.E., Anderson, L.M., and Damaske, D., 2011, East Antarctic rifting triggers uplift of the Gamburtsev Mountains: Nature, v. 479, p. 388−392, https://doi.org/10.1038/nature10566.
  54. Fielding, C.R., Frank, T.D., and Birgenheier, L.P., 2023, A revised, late Paleozoic glacial time-space framework for eastern Australia, and comparison with other regions and events: Earth-Science Reviews, v. 236, 104263, https://doi.org/10.1016/j.earscirev.2022.104263.
  55. Forbes, E., 1846, On the connexion between the distribution of existing fauna and flora of the British Isles, and the geological changes which have affected their area, especially during the epoch of the Northern Drift: Memoirs of the Geological Survey of Great Britain, v. 1, p. 336−403.
  56. Gaidos, E.J., Nealson, K.H., and Kirschvink, J.L., 1999, Life in ice-covered oceans: Science, v. 284, p. 1631−1633, https://doi.org/10.1126/science.284.5420.1631.
  57. Goodman, J.C., 2006, Through thick and thin: marine and meteoric ice in a "Snowball Earth" climate: Geophysical Research Letters, v. 33, L16701, https://doi.org/10.1029/2006GL026840.
  58. Goodman, J.C., and Pierrehumbert, R.T., 2003, Glacial flow of floating marine ice in "Snowball Earth": Journal of Geophysical Research: Oceans, v. 108, 3308, https://doi.org/10.1029/2002JC001471.
  59. Goodman, J.C., and Strom, D.C., 2013, Feedbacks in a coupled ice−atmosphere−dust model of the glacial Neoproterozoic "Mudball Earth": Journal of Geophysical Research: Atmospheres, v. 118, p. 11546–11557, https://doi.org/10.1002/jgrd.50849.
  60. Halverson, G.P., Porter, S., and Shields, G., 2020, The Tonian and Cryogenian Periods, in Gradstein, F.M., Ogg, J.G., Schmitz, M.D., and Ogg, G.M., eds., Geologic Time Scale 2020: Elsevier, Amsterdam, v. 1, p. 495−519, https://doi.org/10.1016/B978-0-12-824360-2.00017-6.
  61. Hawes, I., Jungblut, A.D., Matys, E.D., and Summons, R.E., 2018, The 'dirty ice" of the McMurdo Ice Shelf: analogues for biological oases during the Croygenian: Geobiology, v. 16, p. 369−377, https://doi.org/10.1111/gbi.12280.
  62. Herbert, S., 2005, Charles Darwin, Geologist: Cornell University Press, Ithaca, New York, 485 p.
  63. Herries-Davies, G.L., 1969, The Earth in Decay: a History of British Geomorphology 1578−1878: Macdonald & Co., London, 390 p.
  64. Hestmark, G., 2018, Jens Esmark's mountain glacier traverse 1823 - the key to his discovery of Ice Ages: Boreas, v. 47, p. 1−10, https://doi.org/10.1111/bor.12260.
  65. Hoffman, P.F., 2015, The Tooth of Time: James Smith of Jordanhill: Geoscience Canada, v. 42, p. 7−26, https://doi.org/10.12789/geocanj.2015.42.068.
  66. Hoffman, P.F., 2016, Cryoconite pans on Snowball Earth: supraglacial oases for Cryogenian eukaryotes?: Geobiology, v. 14, p. 531−542, https://doi.org/10.1111/gbi.12191.
  67. Hoffman, P.F., 2023, Glacial erosion on a snowball Earth: testing for bias in flux balance, geographic setting, and tectonic regime: Canadian Journal of Earth Sciences, v. 60, p. 765−777, https://doi.org/10.1139/cjes-2022-0004.
  68. Hoffman, P.F., 2025, Ecosystem relocation on Snowball Earth: polar−alpine ancestry of the extant surface biosphere?: Proceedings of the National Academy of Sciences (USA), v. 122, e2414059122, https://doi.org/10.1073/pnas.2414059122.
  69. Husson, J.M., and Peters, S.E., 2017, Atmospheric oxygenation driven by unsteady growth of the continental sedimentary reservoir: Earth and Planetary Science Letters, v. 460, p. 68−75, https://doi.org/10.1016/j.epsl.2016.12.012.
  70. Izaguirre, I., Allende, L., and Schiaffino, M.R., 2021, Phytoplankton in Antarctic lakes: biodiversity and main ecological features: Hydrobiologia, v. 848, p. 177−207, https://doi.org/10.1007/s10750-020-04306-x.
  71. Jacobsen, S.B., and Kaufman, A.J., 1999, The Sr, C and O isotopic evolution of Neoproterozoic seawater: Chemical Geology, v. 161, p. 37−57, https://doi.org/10.1016/S0009-2541(99)00080-7.
  72. Jungblut, A.D., Vincent, W.F., and Lovejoy, C., 2012, Eukaryotes in Arctic and Antarctic cyanobacterial mats: FEMS Microbiological Ecology, v. 82, p. 416−428, https://doi.org/10.1111/j.1574-6941.2012.01418.x.
  73. Keller, C.B., Husson, J.M., Mitchell, R.N., Bottke, W.F., Gernon, T.M., Boehnke, P., Bell, E.A., Swanson-Hysell, N.L., and Peters, S.E., 2019, Neoproterozoic glacial origin of the Great Unconformity: Proceedings of the National Academy of Sciences (USA), v. 116, p. 1136−1145, https://doi.org/10.1073/pnas.1804350116.
  74. Laybourn-Parry, J., and Wadham, J.L., 2014, Antarctic Lakes: Oxford University Press, Oxford, UK, 346 p., https://doi.org/10.1093/acprof:oso/9780199670499.001.0001.
  75. Laybourn-Parry, J., Tranter, M., and Hodson, A.J., 2012, The Ecology of Snow and Ice Environments: Oxford University Press, Oxford, UK, 190 p., https://doi.org/10.1093/acprof:oso/9780199583072.001.0001.
  76. Lea, E.J., Jamieson, S.S.R., and Bentley, M.J., 2024, Alpine topography of the Gamburtsev Subglacial Mountains, Antarctica, mapped from ice sheet surface morphology: The Cryosphere, v. 18, p. 1733−1751, https://doi.org/10.5194/tc-18-1733-2024.
  77. Li, D.D., Luo, G.M., Tang, Q., She, Z.B., and Xiao, S.H., 2023, New record of the green algal fossil Proterocladus and coexisting microfossils from the Meso-Neoproterozoic Diaoyutai Formation in southern Liaoning, North China: Precambrian Research, v. 393, 107104, https://doi.org/10.1016/j.precamres.2023.107104.
  78. Li, D.W., and Pierrehumbert, R.T., 2011, Sea glacier flow and dust transport on Snowball Earth: Geophysical Research Letters, v. 38, L17501, https://doi.org/10.1029/2011GL048991.
  79. Li, Z.-X., Liu, Y.B., and Ernst, R., 2023, A dynamic 2000−540 Ma Earth history: from cratonic amalgamation to the age of supercontinent cycle: Earth-Science Reviews, v. 238, 104336, https://doi.org/10.1016/j.earscirev.2023.104336.
  80. Lyell, C., 1833, The Principles of Geology, Vol. 3: John Murray, London, 509 p. (reprinted in 1990 by University of Chicago Press).
  81. Lyell, C., 1836, Anniversary address of the President, February 19, 1836: Proceedings of the Geological Society, London, v. 2, p. 357−390.
  82. Lyell, C., 1841, Elements of Geology, 1st Edition: John Murray, London, 570 p.
  83. Lyell, C., 1863, The Antiquity of Man: John Murray, London, 551 p.
  84. Macdonald, F.A., Renger, E., Tasistro-Hart, A.R., Byerly, B.L., Jackson, M.G., Bergmann, K.D., Horner, T.J., and Crockford, P.W., 2025, Mantle-like Sr isotopes in a Sturtian cap carbonate in Oman: Geology, v. 53, p. 753−756, https://doi.org/10.1130/G53385.1.
  85. Montross, S.N., Skidmore, M., Tranter, M., Kivimäki, A.-L., and Parkes, R.J., 2013, A microbial driver of chemical weathering in glaciated systems: Geology, v. 41, p. 215−218, https://doi.org/10.1130/G33572.1.
  86. Moret, P., Muriel, P., Jaramillo, R., and Dangles, O., 2019, Humboldt's Tableau Physique revisited: Proceedings of the National Academy of Sciences (USA), v. 116, p. 12889−12894, https://doi.org/10.1073/pnas.1904585116.
  87. Morueta-Holme, N., Engemann, K., Sandoval-Acuña, P., Jonas, J.D., Segnitz, R.M., and Svenning, J.-C., 2015, Strong upslope shifts in Chimborazo's vegetation over two centuries since Humboldt: Proceedings of the National Academy of Sciences (USA), v. 112, p. 12741−12745, https://doi.org/10.1073/pnas.1509938112.
  88. Mueller, D.R., and Vincent, W.F., 2006, Microbial habitat dynamics and ablation control on the Ward Hunt Ice Shelf: Hydrologic Processes, v. 20, p. 857−876, https://doi.org/10.1002/hyp.6113.
  89. Mueller, D.R., Vincent, W.F., Pollard, W.H., and Fritsen, C.H., 2001, Glacial cryoconite ecosystems: a bipolar comparison of algal communities and habitats: Nova Hedwigia, v. 123, p. 173−197.
  90. Nealson, K.H., 1997, The limits of life on Earth and searching for life on Mars: Journal of Geophysical Research: Planets, v. 102, p. 23675−23686, https://doi.org/10.1029/97JE01996.
  91. Ockenden, H., Bingham, R.G., Goldberg, D., Curtis, A., and Morlighem, M., 2026, Complex mesoscale landscapes beneath Antarctica mapped from space: Science, v. 391, p. 314−319, https://doi.org/10.1126/science.ady2532.
  92. Partin, C.A., and Sadler, P.M., 2016, Slow net sediment accumulation sets snowball Earth apart from younger glacial episodes: Geology, v. 44, p. 1019−1022, https://doi.org/10.1130/G38350.1.
  93. Prates, L., Medina, M.E., and Perez, S.I., 2025, Extinct megafauna dominated human subsistence in southern South America before 11,600 years ago: Science Advances, v. 11, eadx2615, https://doi.org/10.1126/sciadv.adx2615.
  94. Rooney, A.D., Strauss, J.V., Brandon, A.D., and Macdonald, F.A., 2015, A Cryogenian chronology: two long-lasting synchronous Neoproterozoic glaciations: Geology, v. 43, p. 459−462, https://doi.org/10.1130/G36511.1.
  95. Roos, J.C., and Vincent, W.F., 1998, Temperature dependence of UV radiation effects on Antarctic cyanobacteria: Journal of Phycology, v. 34, p. 118−125, https://doi.org/10.1046/j.1529-8817.1998.340118.x.
  96. Rudwick, M., 1974, Darwin and Glen Roy: A "great failure" in scientific method?: Studies in the History and Philosophy of Science, Part A, v. 5, p. 97−185, https://doi.org/10.1016/0039-3681(74)90024-7.
  97. Rupke, N.A., 1983, The Great Chain of History: William Buckland and the English School of Geology (1814−1849): Clarendon Press, Oxford, 322 p.
  98. Sánchez-Baracaldo, P., 2015, Origin of marine planktonic cyanobacteria: Scientific Reports, v. 5, 17418, https://doi.org/10.1038/srep17418.
  99. Sánchez-Baracaldo, P., Ridgwell, A., and Raven, J.A., 2014, A Neoproterozoic transition in the marine nitrogen cycle: Current Biology, v. 24, p. 652−657, https://doi.org/10.1016/j.cub.2014.01.041.
  100. Sánchez-Baracaldo, P., Raven, J.A., Pisani, D., and Knoll, A.H., 2017, Early photosynthetic eukaryotes inhabited low-salinity habitats: Proceedings of the National Academy of Sciences (USA), v. 114, p. E7737−E7745, https://doi.org/10.1073/pnas.1620089114.
  101. Smith, J., 1836, On indications of changes in the relative levels of sea and land in the west of Scotland: Proceedings of the Geological Society of London, v. 2, p. 427−429.
  102. Smith, J., 1839a, On the last changes in the relative levels of the land and sea in the British Isles: Memoirs of the Wernerian Natural History Society of Edinburgh, v. 8, p. 49−88, 108−113.
  103. Smith, J., 1839b, On the climate of the Newer Pliocene Tertiary Period: Proceedings of the Geological Society of London, v. 3, p. 118−119 (reprinted in Smith, J., 1962, Researches in New Pliocene and Post-Tertiary Geology: John Gray, Glasgow, 199 p. A collection of Smith's geological papers.).
  104. Soares, A., Edwards, A., An, D., Bagnoud, A., Bradley, J., Barnhart, E., Bomberg, M., Budwill, K., Caffrey, S.M., Fields, M., Gralnick, J., Kadnikov, V., Momper, L., Osburn, M., Mu, A., Moreau, J.W., Moser, D., Purkamo, L., Rassner, S.M., Sheik, C.S., Sherwood Lollar, B., Toner, B.M., Voordouw, G., Wouters, K., and Mitchell, A.C., 2023, A global perspective on bacterial diversity in the terrestrial deep subsurface: Microbiology, v. 169, 001172, https://doi.org/10.1099/mic.0.001172.
  105. Stoeck, T., Kasper, J., Bunge, J., Leslin, C., Ilyin, V., and Epstein, S., 2007, Protistan diversity in the Arctic: a case of paleoclimate shaping modern biodiversity?: PLoS ONE, v. 2, e728, https://doi.org/10.1371/journal.pone.0000728.
  106. Sugden, D.E., 1978, Glacial erosion by the Laurentide Ice Sheet: Journal of Glaciology, v. 20, p. 367−391, https://doi.org/10.3189/S0022143000013915.
  107. Swanson-Hysell, N.L., Zhang, Y.M., Macdonald, F.A., Koran, I., Tasistro-Hart, A.R., and Jay, A.F., 2025, Oman was on the northern margin of a wide late Tonian Mozambique Ocean: Geology, v. 53, p. 909−913, https://doi.org/10.1130/G53450.1.
  108. Tang, Q., Pang, K., Yuan, X.L., and Xiao, S.H., 2020, A one-billion-year-old multicellular chlorophyte: Nature Ecology & Evolution, v. 4, p. 543−549, https://doi.org/10.1038/s41559-020-1122-9.
  109. Tasistro-Hart, A.R., and Macdonald, F.A., 2023, Phanerozoic flooding of North America and the Great Unconformity: Proceedings of the National Academy of Sciences (USA), v. 120, e2309084120, https://doi.org/10.1073/pnas.2309084120.
  110. Tasistro-Hart, A.R., Macdonald, F.A., Crowley, J.L., and Schmitz, M.D., 2025, Four-million-year Marinoan snowball shows multiple routes to deglaciation: Proceedings of the National Academy of Sciences (USA), v. 122, e2418281122, https://doi.org/10.1073/pnas.2418281122.
  111. Thomas, T.B., Macdonald, F.A., and Catling, D.C., 2026, Seafloor weathering can explain the disparate durations of Snowball glaciations: Geology, v. 54, p. 158−162, https://doi.org/10.1130/G53722.1.
  112. Thomson, J., 1871, On the stratified rocks of Islay, in Report of the 41st Meeting of the British Association for the Advancement of Science, Edinburgh: John Murray, London, p. 110−111.
  113. Thomson, J., 1877, XIII. On the geology of the island of Islay: Transactions of the Geological Society of Glasgow, v. 5, p. 200−222, https://doi.org/10.1144/transglas.5.2.200.
  114. Thomson, K., 2009, The Young Charles Darwin: Yale University Press, New Haven, CT, 276 p.
  115. Torres, M.A., Moosdorf, N., Hartmann, J., Adkins, J.F., and West, A.J., 2017, Glacial weathering, sulfide oxidation, and global carbon cycle feedbacks: Proceedings of the National Academy of Sciences (USA), v. 114, p. 8716−8721, https://doi.org/10.1073/pnas.1702953114.
  116. Tziperman, E., Abbot, D.S., Ashkenazy, Y., Gildor, H., Pollard, D., Schoof, C.G., and Schrag, D.P., 2012, Continental constriction and oceanic ice-cover thickness in a Snowball-Earth scenario: Journal of Geophysical Research: Oceans, v. 117, C05016, https://doi.org/10.1029/2011JC007730.
  117. van de Flierdt, T., Hemming, S.R., Goldstein, S.L., Gehrels, G.E., and Cox., S.E., 2008, Evidence against a young volcanic origin of the Gamburtsev Subglacial Mountains, Antarctica: Geophysical Research Letters: Solid Earth, v. 35, L21303, https://doi.org/10.1029/2008GL035564.
  118. Vérard, C., 2021, 888−444 Ma global plate tectonic reconstruction with emphasis on the formation of Gondwana: Frontiers in Earth Science, v. 9, 666153, https://doi.org/10.3389/feart.2021.666153.
  119. Vincent, W.F., and Laybourn-Parry, J., editors, 2008, Polar Lakes and Rivers: Limnology of Arctic and Antarctic Aquatic Ecosystems: Oxford University Press, Oxford, UK, 327 p., https://doi.org/10.1093/acprof:oso/9780199213887.001.0001.
  120. Vincent, W.F., Gibson, J.A.E., Pienitz, R., Villeneuve, V., Broady, P.A., Hamilton, P.B., and Howard-Williams, C., 2000, Ice shelf microbial ecosystems in the high Arctic and implications for life on Snowball Earth: Naturwissenschaften, v. 87, p. 137−141, https://doi.org/10.1007/s001140050692.
  121. Vincent, W.F., Mueller, D.R., and Bonilla, S., 2004, Ecosystems on ice: the microbial ecology of Markham Ice Shelf in the high Arctic: Cryobiology, v. 48, p. 103−112, https://doi.org/10.1016/j.cryobiol.2004.01.006.
  122. Voigt, A., and Abbot, D.S., 2012, Sea-ice dynamics strongly promote Snowball Earth initiation and destabilize tropical sea-ice margins: Climate of the Past, v. 8, p. 2079−2092, https://doi.org/10.5194/cp-8-2079-2012.
  123. von Humboldt, A., and Bonpland, A., 2009, Essay on the Geography of Plants-with a physical tableau of the equinoctial regions (1807), in Jackson, S.T., ed., Essay on the Geography of Plants: University of Chicago Press, Chicago, p. 61−155. (English translation of von Humbolt and Bonpland 1807).
  124. Wadham, J.L., Tranter, M., Skidmore, M., Hodson, A.J., Priscu, J., Lyons, W.B., Sharp, M., Wynn, P., and Jackson, M., 2010, Biogeochemical weathering under ice: size matters: Global Biogeochemical Cycles, v. 24, GB3025, https://doi.org/10.1029/2009GB003688.
  125. Warren, S.G., Brandt, R.E., Grenfell, T.C., and McKay, C.P., 2002, Snowball Earth: ice thickness on the tropical ocean: Journal of Geophysical Research: Oceans, v. 107, p. 31-1–31-18, https://doi.org/10.1029/2001JC001123.
  126. White, W.A., 1972, Deep erosion by continental ice sheets: Geological Society of America Bulletin, v. 83, p. 1037−1056, https://doi.org/10.1130/0016-7606(1972)835B1037:DEBCIS5D2.0.CO;2.
  127. Worsley, P., 2006, Jens Esmark, Vassryggen and early glacial theory in Britain: Mercian Geologist, v. 16, no. 3, p. 161−172.
  128. Worsley, P., 2008, Esmark's end moraine and the glacial theory from a British perspective: Earth Sciences History, v. 27, p. 12−30, https://doi.org/10.17704/eshi.27.1.6184m4v727vg0403.
  129. Xiao, S.H., Droser, M., Gehling, J.G., Hughes, I.V., Wan, B., Chen, Z., and Yuan, X.L., 2013, Affirming life aquatic for the Ediacara biota in China and Australia: Geology, v. 41, p. 1095−1098, https://doi.org/10.1130/G34691.1.
  130. Yan, M.Y., Yang, J., and Li, D.W., 2025, Simulating continental dust on a hard Snowball Earth: 2. climatic effect of dust: Journal of Geophysical Research: Atmospheres, v. 130, e2025JD043536, https://doi.org/10.1029/2025JD043536.
  131. Yang, C., Li, Y., Selby, D., Wan, B., Guan, C.G., Zhou, C.M., and Li, X.-H., 2022, Implications for Ediacaran biological evolution from the ca. 602 Ma Lantian biota in China: Geology, v. 50, p. 562−566, https://doi.org/10.1130/G49734.1.
  132. Ye, Q., Tong, J.N., Xiao, S.H., Zhu, S.X., An, Z.H., Tian, L., and Hu, J., 2015, The survival of benthic macroscopic phototrophs on a Neoproterozoic snowball Earth: Geology, v. 43, p. 507−510, https://doi.org/10.1130/G36640.1.
  133. Yuan, X.L., Li, J., and Cao, R.J., 1999, A diverse metaphyte assemblage from the Neoproterozoic black shales of South China: Lethaia, v. 32, p. 143−155, https://doi.org/10.1111/j.1502-3931.1999.tb00533.x.
  134. Yuan, X.L., Chen, Z., Xiao, S.H., Zhou, C.M., and Hua, H., 2011, An early Ediacaran assemblage of macroscopic and morphologically differentiated eukaryotes: Nature, v. 470, p. 390−393, https://doi.org/10.1038/nature09810.
  135. Yuan, X.L., Wan, B., Guan, C.G., Che, Z., Zhou, C.M., Xiao, S.H., Wang, W., Pang, K., Tang, Q., and Hua, H., 2016, Lantian Biota (in Chinese): Shanghai Scientific & Technical Publishers, Shanghai, China, 138 p.
  136. Zachos, J., Pagani, M., Sloan, L., Thomas, E., and Billups, K., 2001, Trends, rhythms, and aberrations in global climate 65 Ma to Present: Science, v. 292, p. 686−693, https://doi.org/10.1126/science.1059412.
  137. Žárský, J., Žárský, V., Hanáček, M., and Žárský, V., 2022, Cryogenian glacial habitats as a plant terrestrialisation cradle - the origin of the Anydrophytes and Zygnematophyceae split: Frontiers in Plant Science, v. 12, 735020, https://doi.org/10.3389/fpls.2021.735020.
  138. Zawierucha, K., Kolicka, M., Takeuchi, N., and Kaczmarek, Ł., 2015, What animals can live in cryoconite holes? A faunal review: Journal of Zoology, v. 295, p. 159−169, https://doi.org/10.1111/jzo.12195.
  139. Zhang, H., Sun, Y., Zeng, Q.L., Crowe, S.A., and Luo, H.W., 2021, Snowball Earth, population bottleneck and Prochlorococcus evolution: Proceedings of the Royal Society B, v. 288, 20211956, https://doi.org/10.1098/rspb.2021.1956.
  140. Zhang, H., Hellweger, F.L., and Luo, H.W., 2024, Genome reduction occurred in early Prochlorococcus with an unusually low effective population size: The ISME Journal, v. 18, wrad035, https://doi.org/10.1093/ismejo/wrad035.
  141. Zhou, C.M., Huyskens, M.H., Lang, X.G., Xiao, S.H., and Yin, Q.-Z., 2019, Calibrating the terminations of Cryogenian global glaciations: Geology, v. 47, p. 251−254, https://doi.org/10.1130/G45719.1.