Mass Extinction and Survival during the Permian-Triassic Crisis
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摘要: 当今人类正在面临大气二氧化碳浓度升高、全球变暖、海洋酸化等一系列气候环境问题,有科学家提出这可能导致第6次生物大灭绝.类似的灾难事件在地质历史上多次发生,因此以史为鉴、以古示今才能更好地认识、应对和解决这些问题.显生宙最大的一次生物灭绝事件发生在2.52亿年前的二叠纪-三叠纪之交,超过90%的海洋物种永久消失.此次生物灭绝的过程和原因一直是科学家关注和致力解决的关键科学问题之一.近年来的研究表明当前人类面临的这些极端的气候环境事件在2.52亿年前也都有发生,而且更为严重.本文重点围绕近年来有关二叠纪-三叠纪之交的生物和环境事件研究进展,结合化石和环境指标的地质记录以及生物与环境之间的相互作用关系,总结生物灭绝的过程和形式及相关环境因子的贡献,并探讨残存生物能够躲过这次灾难事件得以延续和发展的内在机制和外界原因.Abstract: Today we are facing a series of climatic problems such as elevation of PCO2, global warming, and ocean acidification, which may lead to the sixth mass extinction. The similar extreme climate has been happened repeatedly in the Earth history. Therefore, taking history as mirror will help us to better understand the nature of these problems and resolve them. The most severe extinction happened during the Permian-Triassic transition (252 Ma), eliminating over 90% species. In the past few decades, the process and cause about this extinction event have become the key scientific questions that need to be uncovered. Recent studies show the extreme climatic events we are facing now have been occurred near the Permian-Triassic boundary. The main purpose of this paper is to summarize recent studies on biotic and environmental events during the Permian-Triassic transition, including extinction process, pattern, and causes of the extinction, and survival mechanisms in its aftermath.
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Key words:
- Permian /
- Triassic /
- mass extinction /
- survival /
- paleoenvironment /
- paleontology /
- stratigraphy.
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图 1 显生宙海洋生物多样性曲线
修改自Sepkoski(1978, 1984).a.科级多样性;b.属级多样性
Fig. 1. Marine biodiversity curves during the Phanerozoic
图 2 二叠纪-三叠纪之交生物的两幕式、选择性灭绝
Fig. 2. Episodic and selective extinction of biota during the Permian-Triassic crisis
图 4 古、中生代之交生物类群的转变
a, c.修改自Bambach et al.(2002);b, d, f.修改自Song et al.(2013b);e.数据来源于Sepkoski Online Data (http://strata.geology.wisc.edu/jack/start.php)
Fig. 4. Biotic turnover during the Paleozoic-Mesozoic transition
图 5 不同生物门类对溶氧量和高温的响应
Fig. 5. Biological effects of decreased oxygen concentration and increased temperature
表 1 二叠纪-三叠纪灭绝事件中科、属、种级灭绝率
Table 1. Extinction rates of family, genus, and species during the Permian-Triassic crisis
科级灭绝率
(%)属级灭绝率
(%)种级灭绝率
(%)文献来源 52.0 64.8 96.0 Raup, 1979 57.0 - - Sepkoski, 1984 - - 91.4 殷鸿福等, 1984 51.0 82.0 95.0 Jablonski, 1994 - - 94.0 Jin et al., 2000 - - 90.1 Song et al., 2013b - - 62.0 Wang et al., 2014 表 2 西伯利亚火山年龄与二叠纪-三叠纪之交灭绝事件年龄对比
Table 2. Ages of Siberian volcano and Permian-Triassic extinction event
方法 西伯利亚火山喷发年龄 二叠纪-三叠纪之交生物灭绝年龄 文献来源 40Ar/39Ar 249±4 Ma Rampino and Stothers, 1988 40Ar/39Ar 248.4±2.4 Ma Renne and Basu, 1991 206Pb/238U 248±4 Ma 251±4 Ma Campbell et al., 1992 40Ar/39Ar 250.0±0.3 Ma 250.0±0.2 Ma Renne et al., 1995 40Ar/39Ar 249±2 Ma Dalrymple et al., 1995 40Ar/39Ar 248.3±1.7 Ma Venkatesan et al., 1997 206Pb/238U 251.4±0.3 Ma Bowring et al., 1998 40Ar/39Ar 249.40±0.51 Ma Reichow et al., 2002 206Pb/238U 251.7±0.4 Ma Kamo et al., 2003 206Pb/238U 252.6±0.2 Ma Mundil et al., 2004 40Ar/39Ar 250.30±1.10 Ma 249.25±0.14 Ma Reichow et al., 2009 206Pb/238U 252.28±0.08 Ma Shen et al., 2011 206Pb/238U 251.941±0.037 Ma Burgess et al., 2014 -
[1] Alvarez, L.W., Alvarez, W., Asaro, F., et al., 1980.Extraterrestrial Cause for the Cretaceous-Tertiary Extinction.Science, 208(4448):1095-1108.doi: 10.1126/science.208.4448.1095 [2] Angiolini, L., Carabelli, L., 2010.Upper Permian Brachiopods from the Nesen Formation, North Iran.Special Papers in Palaeontology, (84):41-90. https://www.researchgate.net/publication/287563855_Upper_Permian_brachiopods_from_the_Nesen_formation_North_Iran [3] Bambach, R.K., Knoll, A.H., Sepkoski, J.J., 2002.Anatomical and Ecological Constraints on Phanerozoic Animal Diversity in the Marine Realm.Proceedings of the National Academy of Sciences of the United States of America, 99(10):6854-6859.doi: 10.1073/pnas.092150999 [4] Barnosky, A.D., Matzke, N., Tomiya, S., et al., 2011.Has the Earth/'s Sixth Mass Extinction already Arrived?Nature, 471(7336):51-57.doi: 10.1038/nature09678 [5] Basu, A.R., Petaev, M.I., Poreda, R.J., et al., 2003.Chondritic Meteorite Fragments Associated with the Permian-Triassic Boundary in Antarctica.Science, 302(5649):1388-1392.doi: 10.1126/science.1090852 [6] Becker, L., Poreda, R.J., Hunt, A.G., et al., 2001.Impact Event at the Permian-Triassic Boundary:Evidence from Extraterrestrial Noble Gases in Fullerenes.Science, 291(5508):1530-1533.doi: 10.1126/science.1057243 [7] Becker, L., Poreda, R.J., Basu, A.R., et al., 2004.Bedout:A Possible End-Permian Impact Crater Offshore of Northwestern Australia.Science, 304(5676):1469-1476.doi: 10.1126/science.1093925 [8] Benton, M.J., Zhang, Q.Y., Hu, S.X., et al., 2013.Exceptional Vertebrate Biotas from the Triassic of China, and the Expansion of Marine Ecosystems after the Permo-Triassic Mass Extinction.Earth-Science Reviews, 125:199-243.doi: 10.1016/j.earscirev.2013.05.014 [9] Bond, D.P.G., Wignall, P.B., 2010.Pyrite Framboid Study of Marine Permian-Triassic Boundary Sections:A Complex Anoxic Event and Its Relationship to Contemporaneous Mass Extinction.Geological Society of America Bulletin, 122:1265-1279.doi: 10.1130/B30042.1 [10] Bond, D.P.G., Wignall, P.B., 2014.Large Igneous Provinces and Mass Extinctions:An Update.Geological Society of America Special Papers, 505:SPE505-502.doi: 10.1130/2014.2505(02) [11] Bowring, S.A., Erwin, D.H., Jin, Y.G., et al., 1998.U/ Pb Zircon Geochronology and Tempo of the End-Permian Mass Extinction.Science, 280(5366):1039-1045.doi: 10.1126/science.280.5366.1039 [12] Brennecka, G.A., Herrmann, A.D., Algeo, T.J., et al., 2011.Rapid Expansion of Oceanic Anoxia Immediately before the End-Permian Mass Extinction.Proceedings of the National Academy of Sciences, 108(43):17631-17634.doi: 10.1073/pnas.1106039108 [13] Burgess, S.D., Bowring, S., Shen, S.Z., 2014.High-Precision Timeline for Earth's most Severe Extinction.Proceedings of the National Academy of Sciences, 111(9):3316-3321.doi: 10.1073/pnas.1317692111 [14] Campbell, I.H., Czamanske, G.K., Fedorenko, V.A., et al., 1992.Synchronism of the Siberian Traps and the Permian-Triassic Boundary.Science, 258:1760-1763.doi: 10.1126/science.258.5089.1760 [15] Cao, C.Q., Love, G.D., Hays, L.E., et al., 2009.Biogeochemical Evidence for Euxinic Oceans and Ecological Disturbance Presaging the End-Permian Mass Extinction Event.Earth and Planetary Science Letters, 281(3-4):188-201.doi: 10.1016/j.epsl.2009.02.012 [16] Chen, J., Shen, S.Z., Li, X.H., et al., 2016.High-Resolution SIMS Oxygen Isotope Analysis on Conodont Apatite from South China and Implications for the End-Permian Mass Extinction.Palaeogeography, Palaeoclimatology, Palaeoecology, 448:26-38.doi: 10.1016/j.palaeo.2015.11.025 [17] Chen, J., Tong, J.N., Song, H.J., et al., 2015.Recovery Pattern of Brachiopods after the Permian-Triassic Crisis in South China.Palaeogeography, Palaeoclimatology, Palaeoecology, 433:91-105.doi: 10.1016/j.palaeo.2015.05.020 [18] Chu, D.L., Tong, J.N., Song, H.J., et al., 2015.Lilliput Effect in Freshwater Ostracods during the Permian-Triassic Extinction.Palaeogeography, Palaeoclimatology, Palaeoecology, 435:38-52.doi:doi: 10.1016/j.palaeo.2015.06.003 [19] Clarkson, M.O., Kasemann, S.A., Wood, R.A., et al., 2015.Ocean Acidification and the Permo-Triassic Mass Extinction.Science, 348(6231):229-232.doi: 10.1126/science.aaa0193 [20] Cuvier, G., 1812.Recherches sur les Ossements Fossiles Discours Préliminaire:Anatomie des Catastrophes.The National Scientific and Technical Research Council, Buenos Aires (in French). [21] Dalrymple, G.B., Czamanske, G.K., Fedorenko, V.A., et al., 1995.A Reconnaissance 40Ar/39Ar Geochronologic Study of Ore-Bearing and Related Rocks, Siberian Russia.Geochimica et Cosmochimica Acta, 59(10):2071-2083.doi: 10.1016/0016-7037(95)00127-1 [22] Erwin, D.H., 1994.The Permo-Triassic Extinction.Nature, 367(6460):231-236.doi: 10.1038/367231a0 [23] Fang, Z.J., 2003.Discussion on the Study of the Genera Claraia and Claraioides (Bivalvia) of Late Permian Age.Acta Palaeontologica Sinica, 42(4):613-619 (in Chinese with English abstract). [24] Farley, K.A., Mukhopadhyay, S., 2001.An Extraterrestrial Impact at the Permian-Triassic Boundary?Science, 293(5539):2343.doi: 10.1126/science.293.5539.2343a [25] Forel, M.B., Crasquin, S., Chitnarin, A., et al., 2015.Precocious Sexual Dimorphism and the Lilliput Effect in Neo-Tethyan Ostracoda (Crustacea) through the Permian-Triassic Boundary.Palaeontology, 58(3):409-454.doi: 10.1111/pala.12151 [26] Fraiser, M.L., Twitchett, R.J., Bottjer, D.J., 2005.Unique Microgastropod Biofacies in the Early Triassic:Indicator of Long-Term Biotic Stress and the Pattern of Biotic Recovery after the End-Permian Mass Extinction.Comptes Rendus Palevol, 4(6-7):543-552.doi: 10.1016/j.crpv.2005.04.006 [27] Grasby, S., Beauchamp, B., Embry, A., et al., 2013.Recurrent Early Triassic Ocean Anoxia.Geology, 41(2):175-178.doi: 10.1130/G33599.1 [28] Grice, K., Cao, C., Love, G.D., et al., 2005.Photic Zone Euxinia during the Permian-Triassic Superanoxic Event.Science, 307(5710):706-709.doi: 10.1126/science.1104323 [29] Groves, J.R., Rettori, R., Payne, J.L., et al., 2007.End-Permian Mass Extinction of Lagenide Foraminifers in the Southern Alps (Northern Italy).Journal of Paleontology, 81(3):415-434.doi: 10.1666/05123.1 [30] Hallam, A., Wignall, P.B., 1997.Mass Extinctions and Their Aftermath.Oxford University Press, Oxford. [31] Haq, B.U., Hardenbol, J., Vail, P.R., 1987.Chronology of Fluctuating Sea Levels since the Triassic.Science, 235(4793):1156-1167.doi: 10.1126/science.235.4793.1156 [32] Haq, B.U., Schutter, S.R., 2008.A Chronology of Paleozoic Sea-Level Changes.Science, 322(5898):64-68.doi: 10.1126/science.1161648 [33] Harries, P.J., Kauffman, E.G., Hansen, T.A., 1996.Models for Biotic Survival Following Mass Extinction.Geological Society Special Publication, 102(1):41-60. doi: 10.1144/GSL.SP.1996.001.01.03 [34] Hayami, I., 1998.Ecology of Mass Extinctions:The Diversity and Shell Size of Bivalves through Time.Iden, 52:38-44 (in Japanese). http://evolution.berkeley.edu/evolibrary/news/120901_afterextinction [35] Hayami, I., 1997.Size Changes of Bivalves and a Hypothesis about the Cause of Mass Extinction.Fossils, 62:24-36 (in Japanese). [36] He, W.H., Shi, G.R., Twitchett, R.J., et al., 2015.Late Permian Marine Ecosystem Collapse Began in Deeper Waters:Evidence from Brachiopod Diversity and Body Size Changes.Geobiology, 13(2):123-138.doi: 10.1111/gbi.12119 [37] He, W.H., Feng, Q.L., Weldon, E.A., et al., 2007a.A Late Permian to Early Triassic Bivalve Fauna from the Dongpan Setion, Southern Guangxi, South China.Journal of Paleontology, 81(5):1009-1019.doi:?10.1666/pleo05-158 [38] He, W.H., Shi, G.R., Feng, Q.L., et al., 2007b.Brachiopod Miniaturization and Its Possible Causes during the Permian-Triassic Crisis in Deep Water Environments, South China.Palaeogeography, Palaeoclimatology, Palaeoecology, 252(1-2):145-163.doi: 10.1016/j.palaeo.2006.11.040 [39] Heydari, E., Hassanzadeh, J., 2003.Deev Jahi Model of the Permian-Triassic Boundary Mass Extinction:A Case for Gas Hydrates as the Main Cause of Biological Crisis on Earth.Sedimentary Geology, 163(1-2):147-163.doi: 10.1016/j.sedgeo.2003.08.002 [40] Hinojosa, J.L., Brown, S.T., Chen, J., et al., 2012.Evidence for End-Permian Ocean Acidification from Calcium Isotopes in Biogenic Apatite.Geology, 40(8):743-746.doi: 10.1130/G33048.1 [41] Hoegh-Guldberg, O., 1999.Climate Change, Coral Bleaching and the Future of the World's Coral Reefs.Marine and Freshwater Research, 50(8):839-866.doi: 10.1071/MF99078 [42] Huey, R.B., Ward, P.D., 2005.Hypoxia, Global Warming, and Terrestrial Late Permian Extinctions.Science, 308(5720):398-401.doi: 10.1126/science.1108019 [43] Huttenlocker, A.K., 2014.Body Size Reductions in Nonmammalian Eutheriodont Therapsids (Synapsida) during the End-Permian Mass Extinction.PloS One, 9(2):e87553.doi: 10.1371/journal.pone.0087553 [44] Huttenlocker, A.K., Botha-Brink, J., 2013.Body Size and Growth Patterns in the Therocephalian Moschorhinus Kitchingi (Therapsida:Eutheriodontia) before and after the End-Permian Extinction in South Africa.Paleobiology, 39(2):253-277.doi: 10.1666/12020 [45] Isozaki, Y., 1997.Permo-Triassic Boundary Superanoxia and Stratified Superocean:Records from Lost Deep Sea.Science, 276(5310):235-238.doi: 10.1126/science.276.5310.235 [46] Isozaki, Y., 2001.An Extraterrestrial Impact at the Permian-Triassic Boundary?Science, 293(5539):2343.doi: 10.1126/science.293.5539.2343a [47] Jablonski, D., 1994.Extinctions in the Fossil Record.Philosophical Transactions of the Royal Society of London B:Biological Sciences, 344(1307):11-17.doi: 10.1098/rstb.1994.0045 [48] Jiang, H.S., Lai, X.L., Sun, Y.D., et al., 2014.Permian-Triassic Conodonts from Dajiang (Guizhou, South China) and Their Implication for the Age of Microbialite Deposition in the Aftermath of the End-Permian Mass Extinction.Journal of Earth Science, 25(3):413-430.doi: 10.1007/s12583-014-0444-4 [49] Jin, Y.G., Wang, Y., Wang, W., et al., 2000.Pattern of Marine Mass Extinction near the Permian-Triassic Boundary in South China.Science, 289(5478):432-436.doi: 10.1126/science.289.5478.432 [50] Jin, Y.G., Zhang, J., Shang, Q., 1994.Two Phases of the End-Permian Mass Extinction.Canadian Society of Petroleum Geologists, 17:813-822. http://archives.datapages.com/data/cspg_sp/data/017/017001/813_cspgsp0170813.htm [51] Joachimski, M.M., Lai, X., Shen, S., et al., 2012.Climate Warming in the Latest Permian and the Permian-Triassic Mass Extinction.Geology, 40(3):195-198.doi: 10.1130/G32707.1 [52] Kaiho, K., Kajiwara, Y., Nakano, T., et al., 2001.End-Permian Catastrophe by a Bolide Impact:Evidence of a Gigantic Release of Sulfur from the Mantle.Geology, 29(9):815-818.doi: 10.1130/0091-7613(2001) [53] Kamo, S.L., Czamanske, G.K., Amelin, Y., et al., 2003.Rapid Eruption of Siberian Flood-Volcanic Rocks and Evidence for Coincidence with the Permian-Triassic Boundary and Mass Extinction at 251 Ma.Earth and Planetary Science Letters, 214(1):75-91.doi: 10.1016/S0012-821X(03)00347-9 [54] Kauffman, E.G., Erwin, D.H., 1995.Surviving Mass Extinctions.Geotimes, 14(3):14-17. https://www.researchgate.net/publication/284046016_Surviving_mass_extinctions [55] Kershaw, S., Zhang, T.S., Lan, G.Z., 1999.A?Microbialite Carbonate Crust at the Permian-Triassic Boundary in South China, and Its Palaeoenvironmental Significance.Palaeogeography, Palaeoclimatology, Palaeoecology, 146:1-18.doi: 10.1016/S0031-0182(98)00139-4 [56] Knoll, A.H., Bambach, R.K., Canfield, D.E., et al., 1996.Comparative Earth History and Late Permian Mass Extinction.Science, 273(5274):452-457.doi: 10.1126/science.273.5274.452 [57] Koeberl, C., Farley, K.A., Peucker-Ehrenbrink, B., et al., 2004.Geochemistry of the End-Permian Extinction Event in Austria and Italy:No Evidence for an Extraterrestrial Component.Geology, 32(12):1053-1056.doi: 10.1130/G20907.1 [58] Korte, C., Kozur, H.W., Joachimski, M.M., et al., 2004.Carbon, Sulfur, Oxygen and Strontium Isotope Records, Organic Geochemistry and Biostratigraphy Across the Permian/Triassic Boundary in Abadeh, Iran.International Journal of Earth Sciences, 93:565-581.doi: 10.1007/s00531-004-0406-7 [59] Korte, C., Pande, P., Kalia, P., et al., 2010.Massive Volcanism at the Permian-Triassic Boundary and Its Impact on the Isotopic Composition of the Ocean and Atmosphere.Journal of Asian Earth Sciences, 37(4):293-311.doi: 10.1016/j.jseaes.2009.08.012 [60] Kozur, H.W., Weems, R.E., 2011.Detailed Correlation and Age of Continental Late Changhsingian and Earliest Triassic Beds:Implications for the Role of the Siberian Trap in the Permian-Triassic Biotic Crisis.Palaeogeography, Palaeoclimatology, Palaeoecology, 308(1-2):22-40.doi: 10.1016/j.palaeo.2011.02.020 [61] Kump, L.R., Pavlov, A., Arthur, M.A., 2005.Massive Release of Hydrogen Sulfide to the Surface Ocean and Atmosphere during Intervals of Oceanic Anoxia.Geology, 33(5):397-400.doi: 10.1130/G21295.1 [62] Lehrmann, D.J., 1999.Early Triassic Calcimicrobial Mounds and Biostromes of the Nanpanjiang Basin, South China.Geology, 27(4):359-362.doi: 10.1130/0091-7613(1999) [63] Li, Y.F., Liang, H.D., Yin, H.F., et al., 2005.Determination of Fullerenes (C60/C70) from the Permian-Triassic Boundary in the Meishan Section of South China.Acta Geologica Sinica, 79(1):11-15.doi: 10.1111/j.1755-6724.2005.tb00862.x [64] Liang, H.D., 2002.End-Permian Catastrophic Event of Marine Acidification by Hydrated Sulfuric Acid:Mineralogical Evidence from Meishan Section of South China.Chinese Science Bulletin, 47(10):784-788 (in Chinese). http://www.cqvip.com/QK/86894X/200216/1004290117.html [65] Liao, W., Wang, Y., Kershaw, S., et al., 2010.Shallow-Marine Dysoxia Across the Permian-Triassic Boundary:Evidence from Pyrite Framboids in the Microbialite in South China.Sedimentary Geology, 232(1-2):77-83.doi: 10.1016/j.sedgeo.2010.09.019 [66] Luo, G.M., Kump, L.R., Wang, Y.B., et al., 2010.Isotopic Evidence for an Anomalously Low Oceanic Sulfate Concentration Following End-Permian Mass Extinction.Earth and Planetary Science Letters, 300(1-2):101-111.doi: 10.1016/j.epsl.2010.09.041 [67] Luo, G.M., Lai, X.L., Shi, G.R., et al., 2008.Size Variation of Conodont Elements of the Hindeodus-Isarcicella Clade during the Permian-Triassic Transition in South China and Its Implication for Mass Extinction.Palaeogeography, Palaeoclimatology, Palaeoecology, 264(1-2):176-187.doi: 10.1016/j.palaeo.2008.04.015 [68] Luo, G.M., Wang, Y.B., Yang, H., et al., 2011.Stepwise and Large-Magnitude Negative Shift in Δ13Ccarb Preceded the Main Marine Mass Extinction of the Permian-Triassic Crisis Interval.Palaeogeography, Palaeoclimatology, Palaeoecology, 299(1-2):70-82.doi: 10.1016/j.palaeo.2010.10.035 [69] MacArthur, R.H., Wilson, E.O., 1967.The Theory of Island Biogeography.Princeton University Press, Princeton. [70] Mundil, R., Ludwig, K.R., Metcalfe, L., et al., 2004.Age and Timing of the Permian Mass Extinctions:U/Pb Dating of Closed-System Zircons.Science, 305(5691):1760-1763.doi: 10.1126/science.1101012 [71] Mutter, R.J., Neuman, A.G., 2009.Recovery from the End-Permian Extinction Event:Evidence from "Lilliput Listracanthus".Palaeogeography, Palaeoclimatology, Palaeoecology, 284(1-2):22-28.doi: 10.1016/j.palaeo.2009.08.024 [72] Payne, J.L., 2005.Evolutionary Dynamics of Gastropod Size across the End-Permian Extinction and through the Triassic Recovery Interval.Paleobiology, 31(2):269-290.doi:10.1666/0094-8373(2005)031[0269:EDOGSA]2.0.CO; 2 [73] Payne, J.L., Lehrmann, D.J., Follett, D., et al., 2007.Erosional Truncation of Uppermost Permian Shallow-Marine Carbonates and Implications for Permian-Triassic Boundary Events.Geological Society of America Bulletin, 119(7-8):771-784.doi: 10.1130/B26091.1 [74] Payne, J.L., Lehrmann, D.J., Wei, J., et al., 2004.Large Perturbations of the Carbon Cycle during Recovery from the End-Permian Extinction.Science, 305(5683):506-509.doi: 10.1126/science.1097023 [75] Payne, J.L., Turchyn, A.V., Paytan, A., et al., 2010.Calcium Isotope Constraints on the End-Permian Mass Extinction.Proceedings of the National Academy of Sciences, 107(19):8543-8548.doi: 10.1073/pnas.0914065107 [76] Peng, Y.Q., Shi, G.R., Gao, Y.Q., et al., 2007.How and why did the Lingulidae (Brachiopoda) not only Survive the End-Permian Mass Extinction but also Thrive in Its Aftermath?Palaeogeography, Palaeoclimatology, Palaeoecology, 252(1-2):118-131.doi: 10.1016/j.palaeo.2006.11.039 [77] Phillips, J., 1860.Life on Earth—Its Origin and Succession.MacMillan, Cambridge. [78] Pruss, S.B., Bottjer, D.J., Corsetti, F.A., et al., 2006.A Global Marine Sedimentary Response to the End-Permian Mass Extinction:Examples from Southern Turkey and the Western United States.Earth-Science Reviews, 78(3-4):193-206.doi: 10.1016/j.earscirev.2006.05.002 [79] Rampino, M.R., Adler, A.C., 1998.Evidence for Abrupt Latest Permian Mass Extinction of Foraminifera:Results of Tests for the Signor-Lipps Effect.Geology, 26(5):415-418.doi: 10.1130/0091-7613 [80] Rampino, M.R., Stothers, R.B., 1988.Flood Basalt Volcanism during the Past 250 Million Years.Science, 241(4866):663-668.doi: 10.1126/science.241.4866.663 [81] Raup, D.M., 1979.Size of the Permo-Triassic Bottleneck and Its Evolutionary Implications.Science, 206(4415):217-218.doi: 10.1126/science.206.4415.217 [82] Raup, D.M., Sepkoski, J.J., 1982.Mass Extinctions in the Marine Fossil Record.Science, 215(4539):1501-1503.doi: 10.1126/science.215.4539.1501 [83] Raup, D.M., Sepkoski, J.J., 1984.Periodicity of Extinctions in the Geologic Past.Proceedings of the National Academy of Sciences, 81(3):801-805.doi: 10.1073/pnas.81.3.801 [84] Reichow, M.K., Pringle, M.S., Al'Mukhamedov, A.I., et al., 2009.The Timing and Extent of the Eruption of the Siberian Traps Large Igneous Province:Implications for the End-Permian Environmental Crisis.Earth and Planetary Science Letters, 277(1-2):9-20.doi: 10.1016/j.epsl.2008.09.030 [85] Reichow, M.K., Saunders, A.D., White, R.V., et al., 2002.40Ar/39Ar Dates from the West Siberian Basin:Siberian Flood Basalt Province Doubled.Science, 296(5574):1946-1849.doi: 10.1126/science.1071671 [86] Renne, P.R., Basu, A.R., 1991.Rapid Eruption of the Siberian Traps Flood Basalts at the Permo-Triassic Boundary.Science, 253(5016):176-179.doi: 10.1126/science.253.5016.176 [87] Renne, P.R., Zhang, Z.C., Richards, M.A., et al., 1995.Synchrony and Causal Relations between Permian-Triassic Boundary Crises and Siberian Flood Volcanism.Science, 269(5229):1413-1416.doi: 10.1126/science.269.5229.1413 [88] Retallack, G.J., Jahren, A.H., 2008.Methane Release from Igneous Intrusion of Coal during Late Permian Extinction Events.The Journal of Geology, 116(1):1-20.doi: 10.1086/524120 [89] Retallack, G.J., Seyedolali, A., Krull, E.S., et al., 1998.Search for Evidence of Impact at the Permian-Triassic Boundary in Antarctica and Australia.Geology, 26(11):979-982.doi: 10.1130/0091-7613 [90] Rodland, D.L., Bottjer, D.J., 2001.Biotic Recovery from the End-Permian Mass Extinction:Behavior of the Inarticulate Brachiopod Lingula as a Disaster Taxon.Palaios, 16(1):95-101.doi:10.1669/0883-1351(2001)016<0095:BRFTEP>2.0.CO; 2 [91] Romano, C., Goudemand, N., Vennemann, T.W., et al., 2013.Climatic and Biotic Upheavals Following the End-Permian Mass Extinction.Nature Geoscience, 6(1):57-60.doi: 10.1038/ngeo1667 [92] Rong, J.Y., Huang, B., 2014.Study of Mass Extinction over the Past Thirty Years:A Synopsis.Science in China (Series D).44(3):377-404 (in Chinese). https://www.sciencedaily.com/releases/2014/07/140716141302.htm [93] Ryskin, G., 2003.Methane-Driven Oceanic Eruptions and Mass Extinctions:Comment and Reply.Geology, 32(1):e42-e43.doi: 10.1130/0091-7613-32.1.e42 [94] Schobben, M., Joachimski, M.M., Korn, D., et al., 2014.Palaeotethys Seawater Temperature Rise and an Intensified Hydrological Cycle Following the End-Permian Mass Extinction.Gondwana Research, 26(2):675-683.doi: 10.1016/j.gr.2013.07.019 [95] Schubert, J.K., Bottjer, D.J., 1992.Early Triassic Stromatolites as Post-Mass Extinction Disaster Forms.Geology, 20(10):883-886.doi:10.1130/0091-7613(1992)?020<0883:ETSAPM>2.3.CO; 2 [96] Schubert, J.K., Bottjer, D.J., 1995.Aftermath of the Permian-Triassic Mass Extinction Event:Paleoecology of Lower Triassic Carbonates in the Western USA.Palaeogeography, Palaeoclimatology, Palaeoecology, 116(1-2):1-39.doi: 10.1016/0031-0182(94)00093-N [97] Sepkoski, J.J., 1978.A Kinetic Model of Phanerozoic Taxonomic Diversity.Ⅰ.Analysis of Marine Orders.Paleobiology, 4:223-251.doi: 10.1017/S0094837300005972 [98] Sepkoski, J.J., 1979.A Kinetic Model of Phanerozoic Taxonomic Diversity:Ⅱ.Early Phanerozoic Families and Multiple Equilibria.Paleobiology, 5:222-251.doi: 10.1017/S0094837300006539 [99] Sepkoski, J.J., 1981.A Factor Anaytic Description of the Phanerozoic Marine Fossil Record.Paleobiology, 7(1):36-53.doi: 10.1017/S0094837300007006 [100] Sepkoski, J.J., 1984.A Kinetic Model of Phanerozoic Taxonomic Diversity, Ⅲ.Post-Paleozoic Families and Mass Extinctions.Paleobiology, 10(2):246-267.doi: 10.1017/S0094837300008186 [101] Shen, J., Algeo, T.J., Zhou, L., et al., 2012.Volcanic Perturbations of the Marine Environment in South China Preceding the Latest Permian Mass Extinction and Their Biotic Effects.Geobiology, 10(1):82-103.doi: 10.1111/j.1472-4669.2011.00306.x [102] Shen, S.Z., Cao, C.Q., Henderson, C.M., et al., 2006.End-Permian Mass Extinction Pattern in the Northern Peri-Gondwanan Region.Palaeoworld, 15(1):3-30.doi: 10.1016/j.palwor.2006.03.005 [103] Shen, S.Z., Archbold, N.W., Shi, G.R., 2001.A Lopingian (Late Permian) Brachiopod Fauna from the Qubuerga Formatin at Shengmi in the Mount Qomolangma Region of Southern Xizang (Tibet), China.Journal of Paleontology, 75(2):274-283.doi:10.1666/0022-3360(2001)075<0274:ALLPBF>2.0.CO; 2 [104] Shen, S.Z., Crowley, J.L., Wang, Y., et al., 2011.Calibrating the End-Permian Mass Extinction.Science, 334(6061):1367-1372.doi: 10.1126/science.1213454 [105] Shen, W., Lin, Y., Xu, L., et al., 2007.Pyrite Framboids in the Permian-Triassic Boundary Section at Meishan, China:Evidence for Dysoxic Deposition.Palaeogeography, Palaeoclimatology, Palaeoecology, 253(3-4):323-331.doi: 10.1016/j.palaeo.2007.06.005 [106] Song, H.J., Tong, J.N., 2010.Size Variation of Foraminifers during the Permian-Triassic Transition at Meishan Section, South China.Journal of Earth Science, 21(S1):154-157.doi: 10.1007/s12583-010-0197-7 [107] Song, H.J., Tong, J.N., Chen, Z.Q., 2009a.Two Episodes of Foraminiferal Extinction near the Permian-Triassic Boundary at the Meishan Section, South China.Australian Journal of Earth Sciences, 56:765-773.doi: 10.1080/08120090903002599 [108] Song, H.J., Tong, J.N., Chen, Z.Q., et al., 2009b.End-Permian Mass Extinction of Foraminifers in the Nanpanjiang Basin, South China.Journal of Paleontology, 83(5):718-738.doi: 10.1666/08-175.1 [109] Song, H.J., Tong, J.N., Chen, Z.Q., 2011.Evolutionary Dynamics of the Permian-Triassic Foraminifer Size:Evidence for Lilliput Effect in the End-Permian Mass Extinction and Its Aftermath.Palaeogeography, Palaeoclimatology, Palaeoecology, 308(1-2):98-110.doi: 10.1016/j.palaeo.2010.10.036 [110] Song, H.J., Tong, J.N., Wignall, P.B., et al., 2016.Early Triassic Disaster and Opportunistic Foraminifers in South China.Geological Magazine, 153(2):298-315.doi: 10.1017/S0016756815000497 [111] Song, H.J., Wignall, P.B., Tong, J.N., et al., 2012.Geochemical Evidence from Bio-Apatite for Multiple Oceanic Anoxic Events during Permian-Triassic Transition and the Link with End-Permian Extinction and Recovery.Earth and Planetary Science Letters, 353-354:12-21.doi: 10.1016/j.epsl.2012.07.005 [112] Song, H.Y., Tong, J.N., Algeo, T.J., et al., 2013a.Large Vertical δ13CDIC Gradients in Early Triassic Seas of the South China Craton:Implications for Oceanographic Changes Related to Siberian Traps Volcanism.Global and Planetary Change, 105:7-20.doi: 10.1016/j.gloplacha.2012.10.023 [113] Song, H.J., Wignall, P.B., Tong, J.N., et al., 2013b.Two Pulses of Extinction during the Permian-Triassic Crisis.Nature Geoscience, 6(1):52-56.doi: 10.1038/ngeo1649 [114] Song, H.Y., Tong, J.N., Algeo, T.J., et al., 2014a.Early Triassic Seawater Sulfate Drawdown.Geochimica et Cosmochimica Acta, 128:95-113.doi: 10.1016/j.gca.2013.12.009 [115] Song, H.J., Wignall, P.B., Chu, D., et al., 2014b.Anoxia/High Temperature Double Whammy during the Permian-Triassic Marine Crisis and Its Aftermath.Scientific Reports, 4(4132).doi: 10.1038/srep04132 [116] Song, H.J., Tong, J.N., Xiong, Y.L., et al., 2012.The Large Increase of δ13Ccarb-Depth Gradient and the End-Permian Mass Extinction.Science in China (Series D), 42(8):1182-1191 (in Chinese). http://kns.cnki.net/KCMS/detail/detail.aspx?filename=jdxg201207005&dbname=CJFD&dbcode=CJFQ [117] Stanley, S.M., Yang, X., 1994.A Double Mass Extinction at the End of the Paleozoic Era.Science, 266(5189):1340-1344.doi: 10.1126/science.266.5189.1340 [118] Stramma, L., Johnson, G.C., Sprintall, J., et al., 2008.Expanding Oxygen-Minimum Zones in the Tropical Oceans.Science, 320(5876):655-658.doi: 10.1126/science.1153847 [119] Sun, Y., Joachimski, M.M., Wignall, P.B., et al., 2012.Lethally Hot Temperatures during the Early Triassic Greenhouse.Science, 338(6105):366-370.doi: 10.1126/science.1224126 [120] Tian, L., Tong, J.N., Algeo, T.J., et al., 2014.Reconstruction of Early Triassic Ocean Redox Conditions Based on Framboidal Pyrite from the Nanpanjiang Basin, South China.Palaeogeography, Palaeoclimatology, Palaeoecology, 412:68-79.doi: 10.1016/j.palaeo.2014.07.018 [121] Tong, J.N., Zhang, S., Zuo, J., et al., 2007a.Events during Early Triassic Recovery from the End-Permian Extinction.Global and Planetary Change, 55(1-3):66-80.doi: 10.1016/j.gloplacha.2006.06.015 [122] Tong, J.N., Zuo, J.X., Chen, Z.Q., 2007b.Early Triassic Carbon Isotope Excursions from South China:Proxies for Devastation and Restoration of Marine Ecosystems Following the End-Permian Mass Extinction.Geological Journal, 42:371-389.doi: 10.1002/gj.1084 [123] Tong, J.N., Yin, H.F., 2015.Triassic Chronostratigraphy and Chinese Stages.Earth Science, 40(2):189-197 (in Chinese with English abstract). [124] Twitchett, R.J., 2007.The Lilliput Effect in the Aftermath of the End-Permian Extinction Event.Palaeogeography, Palaeoclimatology, Palaeoecology, 252(1-2):132-144.doi: 10.1016/j.palaeo.2006.11.038 [125] Venkatesan, T.R., Kumar, A., Gopalan, K., et al., 1997.40Ar-39Ar Age of Siberian Basaltic Volcanism.Chemical Geology, 138(3-4):303-310.doi: 10.1016/S0009-2541(97)00006-5 [126] Wang, Y., Sadler, P.M., Shen, S.Z., et al., 2014.Quantifying the Process and Abruptness of the End-Permian Mass Extinction.Paleobiology, 40(1):113-129.doi: 10.1666/13022 [127] Wang, Y.B., Tong, J.N., Wang, J.S., et al., 2005.Calcimicrobialite after End-Permian Mass Extinction in South China and Its Palaeoenvironmental Significance.Chinese Science Bulletin, 50(6):552-558 (in Chinese). http://www.cqvip.com/qk/86894X/200507/15596221.html [128] Wignall, P.B., 2001.Large Igneous Provinces and Mass Extinctions.Earth-Science Reviews, 53(1-2):1-33.doi: 10.1016/S0012-8252(00)00037-4 [129] Wignall, P.B., 2005.The Link between Large Igneous Province Eruptions and Mass Extinctions.Elements, 1(5):293-297.doi: 10.2113/gselements.1.5.293 [130] Wignall, P.B., Bond, D.P.G., Kuwahara, K., et al., 2010.An 80 Million Year Oceanic Redox History from Permian to Jurassic Pelagic Sediments of the Mino-Tamba Terrane, SW Japan, and the Origin of Four Mass Extinctions.Global and Planetary Change, 71(1-2):109-123.doi: 10.1016/j.gloplacha.2010.01.022 [131] Wignall, P.B., Hallam, A., 1992.Anoxia as a Cause of the Permian/Triassic Mass Extinction:Facies Evidence from Northern Italy and the Western United States.Palaeogeography, Palaeoclimatology, Palaeoecology, 93(1-2):21-46.doi: 10.1016/0031-0182(92)90182-5 [132] Wignall, P.B., Kershaw, S., Collin, P.Y., et al., 2009.Erosional Truncation of Uppermost Permian Shallow-Marine Carbonates and Implications for Permian-Triassic Boundary Events:Comment.Geological Society of America Bulletin, 121(5-6):954-956.doi: 10.1130/B26424.1 [133] Wignall, P.B., Twitchett, R.J., 1996.Oceanic Anoxia and the End Permian Mass Extinction.Science, 272(5265):1155-1158.doi: 10.1126/science.272.5265.1155 [134] Wignall, P.B., Twitchett, R.J., 2002.Extent, Duration, and Nature of the Permian-Triassic Superanoxic Event.Geological Society of America Special Publication, 356:395-413. https://www.researchgate.net/publication/279396802_Extent_duration_and_nature_of_the_Permian-Triassic_superanoxic_event [135] Wu, Y.S., Jiang, H.X., Fan, J.S., 2010.Evidence for Sea-Level Falls in the Permian-Triassic Transition in the Ziyun Area, South China.Geological Journal, 45(2-3):170-185.doi: 10.1002/gj.1168 [136] Xie, S., Pancost, R.D., Wang, Y., et al., 2010.Cyanobacterial Blooms Tied to Volcanism during the 5 M.y.Permo-Triassic Biotic Crisis.Geology, 38(5):447-450.doi: 10.1130/G30769.1 [137] Xu, G.R., Tong, J.N., 1991.Age Correlation and Mass Extinction Event.In:Yang, Z.Y., Wu, S.B., Yin, H.F., et al., eds., Permo-Triassic Events of South China.Geological Publishing House, Beijing (in Chinese). [138] Yang, H., Chen, Z.Q., Wang, Y.B., et al., 2011.Composition and Structure of Microbialite Ecosystems Following the End-Permian Mass Extinction in South China.Palaeogeography, Palaeoclimatology, Palaeoecology, 308(1-2):111-128.doi: 10.1016/j.palaeo.2010.05.029 [139] Yang, L.R., Song, H.J., Tong, J.N., et al., 2013.The Extinction Pattern of Fusulinids during the Permian-Triassic Crisis at the Kangjiaping Section, Cili, Hunan Province.Acta Micropalaeontologica Sinica, 30(4):353-366 (in Chinese with English abstract). http://en.cnki.com.cn/Article_en/CJFDTotal-WSGT201304003.htm [140] Yang, Z.Y., Wu, S.B., Yin, H.F., et al., 1991.Permo-Triassic Events of South China.Geological Publishing House, Beijing (in Chinese). [141] Yin, H.F., Huang, S.J., Zhang, K.X., et al., 1992.The Effects of Volcanism on the Permo-Triassic Mass Extinction in South China.In:Sweet, W.C., Yang, Z.Y., Dickins, J.M., et al., eds., Permo-Triassic Events in the Eastern Tethys, Cambridge University Press, Cambridge, 146-157. [142] Yin, H.F., Feng, Q.L., Lai, X.L., et al., 2007.The Protracted Permo-Triassic Crisis and Multi-Episode Extinction around the Permian-Triassic Boundary.Global and Planetary Change, 55(1-3):1-20.doi: 10.1016/j.gloplacha.2006.06.005 [143] Yin, H.F., Jiang, H.S., Xia, W.C., et al., 2014.The End-Permian Regression in South China and Its Implication on Mass Extinction.Earth-Science Reviews, 137:19-33.doi: 10.1016/j.earscirev.2013.06.003 [144] Yin, H.F., Zhang, K.X., Tong, J.N., et al., 2001.The Global Stratotype Section and Point (GSSP) of the Permian-Triassic Boundary.Episodes, 24(2):102-114. http://en.cnki.com.cn/Article_en/CJFDTotal-DCXZ200702000.htm [145] Yin, H.F., Huang, S.J., Zhang, K.X., et al., 1989.Volcanism at the Permian-Triassic Boundary in South China and Its Effects on Mass Extinction.Acta Geologica Sinica, 63(2):169-180 (in Chinese with English abstract). doi: 10.1111/j.1755-6724.1989.mp2004007.x/abstract [146] Yin, H.F., Song, H.J., 2013.Mass Extinction and Pangea Integration during the Paleozoic-Mesozoic Transition.Science in China (Series D), 43(6):1539-1552 (in Chinese). http://earth.scichina.com:8080/sciDe/EN/abstract/abstract511745.shtml [147] Yin, H.F., Xu, G.R., Ding, M.H., 1984.Paleozoic-Mesozoic Fauna Change of South China.In:Editorial Office of Geology Department, ed., Scientific Papers of Geological International Exchange, Geological Publishing House, Beijing (in Chinese). [148] Zhang, K.X., Tong, J.N., Shi, G.R., et al., 2007.Early Triassic Conodont-Palynological Biostratigraphy of the Meishan D Section in Changxing, Zhejiang Province, South China.Palaeogeography, Palaeoclimatology, Palaeoecology, 252(1-2):4-23.doi: 10.1016/j.palaeo.2006.11.031 [149] Zhao, X.M., Tong, J.N., Yao, H.Z., et al., 2015.Early Triassic Trace Fossils from the Three Gorges Area of South China:Implications for the Recovery of Benthic Ecosystems Following the Permian-Triassic Extinction.Palaeogeography, Palaeoclimatology, Palaeoecology, 429:100-116.doi: 10.1016/j.palaeo.2015.04.008 [150] 方宗杰, 2003.关于晚二叠世克氏蛤类研究中的几个问题.古生物学报, 42(4): 613-619. http://www.cnki.com.cn/Article/CJFDTOTAL-GSWX200304013.htm [151] 梁汉东, 2002.二叠纪末期海洋硫酸化环境灾变事件:煤山剖面岩石矿物证据.科学通报, 47(10): 784-788. doi: 10.3321/j.issn:0023-074X.2002.10.013 [152] 戎嘉余, 黄冰, 2014.生物大灭绝研究三十年.中国科学(D辑), 44(3): 377-404. http://www.cnki.com.cn/Article/CJFDTOTAL-JDXK201403001.htm [153] 宋海军, 童金南, 熊炎林, 等, 2012.δ13Ccarb-深度梯度的剧增与二叠纪末生物大灭绝.中国科学(D辑), 42(8): 1182-1191. http://kns.cnki.net/KCMS/detail/detail.aspx?filename=jdxk201208005&dbname=CJFD&dbcode=CJFQ [154] 童金南, 殷鸿福, 2015.三叠纪年代地层与中国建阶.地球科学, 40(2): 189-197. http://earth-science.net/WebPage/Article.aspx?id=3052 [155] 王永标, 童金南, 王家生, 等, 2005.华南二叠纪末大绝灭后的钙质微生物岩及古环境意义.科学通报, 50(6): 552-558. http://www.cnki.com.cn/Article/CJFDTOTAL-KXTB200506008.htm [156] 徐桂荣, 童金南, 1991. 时间对比与集群灭绝事件. 见杨遵仪, 吴顺宝, 殷鸿福, 等, 编, 华南二叠-三叠纪过渡期地质事件, 北京: 地质出版社. [157] 杨利蓉, 宋海军, 童金南, 等, 2013.湖南慈利康家坪剖面二叠纪末簸类有孔虫的灭绝过程.微体古生物学报, 30(4): 353-366. http://www.cnki.com.cn/Article/CJFDTOTAL-WSGT201304003.htm [158] 杨遵仪, 吴顺宝, 殷鸿福, 等, 1991.华南二叠-三叠纪过渡时期地质事件.北京:地质出版社. [159] 殷鸿福, 黄思骥, 张克信, 等, 1989.华南二叠纪-三叠纪之交的火山活动及其对生物绝灭的影响.地质学报, 63(2): 169-180. http://cdmd.cnki.com.cn/Article/CDMD-10491-1015661028.htm [160] 殷鸿福, 宋海军, 2013.古、中生代之交生物大灭绝与泛大陆聚合.中国科学(D辑), 43(6): 1539-1552. http://www.cnki.com.cn/Article/CJFDTOTAL-JDXK201310002.htm [161] 殷鸿福, 徐桂荣, 丁梅华, 1984. 华南古、中生代之交海洋生物界的更替. 见: 地质部书刊编辑室编, 国际交流地质学术论文集. 北京: 地质出版社.