Capitanian mass extinction event

896:, with the widespread demise of reefs in particular being linked to this marine regression. The Guadalupian-Lopingian boundary coincided with one of the most prominent first-order marine regressions of the Phanerozoic. Evidence for abrupt sea level fall at the terminus of the Guadalupian comes from evaporites and terrestrial facies overlying marine carbonate deposits across the Guadalupian-Lopingian transition. Additionally, a tremendous unconformity is associated with the Guadalupian-Lopingian boundary in many strata across the world. The closure of the Sino-Mongolian Seaway at the end of the Capitanian has been invoked as a potential driver of 884:, show that the large scale decrease in terrestrial vertebrate diversity coincided with volcanism in the Emeishan Traps, although robust evidence for a causal relationship between these two events remains elusive. A 2015 study called into question whether the Capitanian mass extinction event was global in nature at all or merely a regional biotic crisis limited to South China and a few other areas, finding no evidence for terrestrial or marine extinctions in eastern Australia linked to the Emeishan Traps or to any proposed extinction triggers invoked to explain the biodiversity drop in low-latitudes of the Northern Hemisphere. 807:. Euxinia may have been exacerbated even further by the increasing sluggishness of ocean circulation resulting from volcanically driven warming. The initial hydrothermal nature of the Emeishan Traps meant that local marine life around South China would have been especially jeopardised by anoxia due to hyaloclastite development in restricted, fault-bounded basins. Expansion of oceanic anoxia has been posited to have occurred slightly before the Capitanian extinction event itself by some studies, though it is probable that upwelling of anoxic waters prior to the mass extinction was a local phenomenon specific to South China. 119: 24: 666:. The brachiopod-mollusc transition that characterised the broader shift from the Palaeozoic to Modern evolutionary faunas has been suggested to have had its roots in the Capitanian mass extinction event, although other research has concluded that this may be an illusion created by taphonomic bias in silicified fossil assemblages, with the transition beginning only in the aftermath of the more cataclysmic end-Permian extinction. After the Capitanian mass extinction, disaster taxa such as 659:

patterns. A study examining foraminiferal extinctions in particular found that the Central and Western Palaeotethys experienced taxonomic losses of a lower magnitude than the Northern and Eastern Palaeotethys, which had the highest extinction magnitude. The same study found that Panthalassa's overall extinction magnitude was similar to that of the Central and Western Palaeotethys, but that it had a high magnitude of extinction of endemic taxa.

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the most precipitous in the entire geological history of the Earth. The rate of carbon dioxide emissions during the Capitanian mass extinction, though extremely abrupt, was nonetheless significantly slower than that during the end-Permian extinction, during which carbon dioxide levels rose five times faster according to one study. Significant quantities of

378:. Additionally, there is a dispute regarding the severity of the extinction and whether the extinction in China happened at the same time as the extinction in Spitsbergen. According to one study, the Capitanian mass extinction was not one discrete event but a continuous decline in diversity that began at the end of the 2907: 819:

absorbing atmospheric carbon dioxide, it is likely that the excessive volcanic emissions of carbon dioxide resulted in marine hypercapnia, which would have acted in conjunction with other killing mechanisms to further increase the severity of the biotic crisis. The dissolution of volcanically emitted

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global warming. The Emeishan Traps discharged between 130 and 188 teratonnes of carbon dioxide in total, doing so at a rate of between 0.08 to 0.25 gigatonnes of carbon dioxide per year, making them responsible for an increase in atmospheric carbon dioxide that was both one of the largest and one of

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piles from which currently cover an area of 250,000 to 500,000 km, although the original volume of the basalts may have been anywhere from 500,000 km to over 1,000,000 km. The age of the extinction event and the deposition of the Emeishan basalts are in good alignment. Reefs and other

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Whether and to what degree latitude affected the likelihood of taxa to go extinct remains disputed amongst palaeontologists. Whereas some studies conclude that the extinction event was a regional one limited to tropical areas, others suggest that there was little latitudinal variation in extinction

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After the recognition of a separate marine mass extinction at the end of the Guadalupian, the dinocephalian extinction was seen to represent its terrestrial correlate. Though it was subsequently suggested that because the Russian Ischeevo fauna, which was considered the youngest dinocephalian fauna

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intruding into coal-rich deposits has been implicated as an additional driver of warming, though this idea has been challenged by studies that instead conclude that the extinction was precipitated directly by the Emeishan Traps or by their interaction with platform carbonates. The emissions of the

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made up 61.2% of the individuals found in similar environments after the extinction. 87% of brachiopod species and 82% of fusulinacean foraminifer species in South China were lost. Although severe for brachiopods, the Capitanian extinction's impact on their diversity was nowhere near as strong as

803:. Two anoxic events, the middle Capitanian OAE-C1 and the end-Capitanian OAE-C2, occurred thanks to Emeishan volcanic activity. Volcanic greenhouse gas release and global warming increased continental weathering and mineral erosion, which in turn has been propounded as a factor enhancing oceanic 236:

mass extinctions, respectively, while being the fifth worst in terms of ecological severity. The global nature of the Capitanian mass extinction has been called into question by some palaeontologists as a result of some analyses finding it to have affected only low-latitude taxa in the Northern

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enrichment at the Guadalupian-Lopingian boundary further confirms the existence of massive volcanic activity; coronene can only form at extremely high temperatures created either by extraterrestrial impacts or massive volcanism, with the former being ruled out because of an absence of iridium

320:

in terms of the proportion of marine invertebrate genera lost; a different study found the Capitanian extinction event to be only the ninth worst in terms of taxonomic severity (number of genera lost) but found it to be the fifth worst with regard to its ecological impact (i.e., the degree of

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of East Greenland is similar to that of Spitsbergen; the faunal losses in Canada's Sverdrup Basin are comparable to the extinctions in Spitsbergen and East Greenland, but the post-extinction recovery that happened in Spitsbergen and East Greenland did not occur in the Sverdrup Basin. Whereas

144:

that are present in each interval of time but do not exist in the following interval) vs time in the past for marine genera. Geological periods are annotated (by abbreviation and colour) above. The Capitanian extinction event occurred 260–259 million years ago, ~7 million years before the

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Although it is known that the Capitanian mass extinction occurred after Olson's Extinction and before the Permian–Triassic extinction event, the exact age of the Capitanian mass extinction remains controversial. This is partly due to the somewhat circumstantial age of the

871:

shows a positive δ13C excursion and concludes that the end of the Capitanian was marked by massive aridification in the region, although the temperature remained largely the same, suggesting that global climate change did not account for the extinction event. Analysis of

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marine sediments interbedded among basalt piles indicate Emeishan volcanism initially developed underwater; terrestrial outflows of lava occurred only later in the large igneous province's period of activity. These eruptions would have released high doses of toxic

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and photoautotrophic bacteria dominated marine microbial communities. Significant turnovers in microbial ecosystems occurred during the Capitanian mass extinction, though they were smaller in magnitude than those associated with the end-Permian extinction.

1142:

Bond, D. P. G., Wignall, P. B., Wang, W., Izon, G., Jiang, H. S., Lai, X. L., Sund, Y.-D., Newtona, R.J., Shaoe, L.-Y., Védrinea, S. & Cope, H. (2010). "The mid-Capitanian (Middle Permian) mass extinction and carbon isotope record of South China".

836:

and a decline of terrestrial infaunal invertebrates. Some researchers have cast doubt on whether significant acidification took place globally, concluding that the carbon cycle perturbation was too small to have caused a major worldwide drop in

2488:"A remarkable sea-level drop and relevant biotic responses across the Guadalupian–Lopingian (Permian) boundary in low-latitude mid-Panthalassa: Irreversible changes recorded in accreted paleo-atoll limestones in Akasaka and Ishiyama, Japan" 697:

that lived during the late Guadalupian was cut in half by the Capitanian mass extinction. Terrestrial survivors of the Capitanian extinction event were generally 20 kg (44 lb) to 50 kg (110 lb) and commonly found in

427:

directly constraining the extinction horizons themselves in the marine sections, most recent studies refrain from placing a number on its age, but based on extrapolations from the Permian timescale an age of approximately 260–262

361:, Stevens and colleagues found an extinction of 56% of plant species recorded in the mid-Upper Shihhotse Formation in North China, which was approximately mid-Capitanian in age. 24% of plant species in South China went extinct. 382:. Another study examining fossiliferous facies in Svalbard found no evidence for a sudden mass extinction, instead attributing local biotic changes during the Capitanian to the southward migration of many taxa through the 3134: 3354:

Shen, Shu-zhong; Cao, Chang-qun; Zhang, Hua; Bowring, Samuel A.; Henderson, Charles M.; Payne, Jonathan L.; Davydov, Vladimir I.; Chen, Bo; Yuan, Dong-xun; Zhang, Yi-zhun; Wang, Wei; Zheng, Quan-feng (1 August 2013).

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has been estimated; this fits broadly with radiometric ages from the terrestrial realm, assuming the two events are contemporaneous. Plant losses occurred either at the same time as the marine extinction or after it.

676:

became abundant in what is now South China. The initial recovery of reefs consisted of non-metazoan reefs: algal bioherms and algal-sponge reef buildups. This initial recovery interval was followed by an interval of

2779:

Ota, A. & Isozaki, Y. (March 2006). "Fusuline biotic turnover across the Guadalupian–Lopingian (Middle–Upper Permian) boundary in mid-oceanic carbonate buildups: Biostratigraphy of accreted limestone in Japan".

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stage, well before the end of the Guadalupian, this constraint applied to the type locality only. The recognition of a younger dinocephalian fauna in Russia (the Sundyr Tetrapod Assemblage) and the retrieval of

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relationship; many species with poorly buffered respiratory physiologies also became extinct. The extinction event led to a collapse of the reef carbonate factory in the shallow seas surrounding South China.

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may have also played a role in the extinction. Potential drivers of extinction proposed as causes of end-Guadalupian reef decline include fluctuations in salinity and tectonic collisions of microcontinents.

2026:

Wignall, Paul B.; Bond, David P. G.; Haas, János; Wang, Wei; Jiang, Haishui; Lai, Xulong; Altiner, Demir; Védrine, Stéphanie; Hips, Kinga; Zajzon, Norbert; Sun, Yadong; Newton, Robert J. (1 February 2012).

2088:

Wignall, Paul B.; Sun, Yadong; Bond, David P. G.; Izon, Gareth; Newton, Robert J.; Védrine, Stéphanie; Widdowson, Mike; Ali, Jason R.; Lai, Xulong; Jiang, Haishui; Cope, Helen; Bottrell, Simon H. (2009).

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species emerged after the extinction, the dominant position of the brachiopods was taken over by the bivalves. Approximately 70% of other species found at the Kapp Starostin Formation also vanished. The

3624:"Submarine palaeoenvironments during Emeishan flood basalt volcanism, SW China: Implications for plume–lithosphere interaction during the Capitanian, Middle Permian ('end Guadalupian') extinction event" 2670:"Stratigraphy of the Guadalupian (Permian) siliceous deposits from central Guizhou of South China: Regional correlations with implications for carbonate productivity during the Middle Permian biocrisis" 1460: 3517:"Age and duration of the Emeishan flood volcanism, SW China: Geochemistry and SHRIMP zircon U–Pb dating of silicic ignimbrites, post-volcanic Xuanwei Formation and clay tuff at the Chaotian section" 4262:

Song, Huyue; Algeo, Thomas J.; Song, Haijun; Tong, Jinnan; Wignall, Paul B.; Bond, David P. G.; Zheng, Wang; Chen, Xinming; Romaniello, Stephen J.; Wei, Hengye; Anbar, Ariel D. (15 May 2023).

1098:"Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis" 465:

The existence of change in tetrapod faunas in the mid-Permian has long been known in South Africa and Russia. In Russia, it corresponded to the boundary between what became known as the

2266: 1507:

Marchetti, Lorenzo; Logghe, Antoine; Mujal, Eudald; Barrier, Pascal; Montenat, Christian; Nel, André; Pouillon, Jean-Marc; Garrouste, Romain; Steyer, J. Sébastien (1 August 2022).

554:, and brachiopods. It was more severe in restricted marine basins than in the open oceans. It appears to have been particularly selective against shallow-water taxa that relied on 2908:"Earliest Wuchiapingian (Lopingian, late Permian) brachiopods in southern Hunan, South China: implications for the pre-Lopingian crisis and onset of Lopingian recovery/radiation" 1731:

McGhee, G.R., Sheehan, P.M., Bottjer, D.J. & Droser, M.L., 2004. "Ecological ranking of Phanerozoic biodiversity crises: Ecological and taxonomic severities are decoupled".

445:

in Southwest China was originally dated to the end of the Guadalupian, but studies published in 2009 and 2010 dated the extinction of these fusulinaceans to the mid-Capitanian.

4391:"Development of a high-productivity and anoxic-euxinic condition during the late Guadalupian in the Lower Yangtze region: Implications for the mid-Capitanian extinction event" 693:, which were one of the most common elements of tetrapod fauna of the Guadalupian; only one dinocephalian genus survived the Capitanian extinction event. The diversity of the 2840: 273:

began recovery immediately after the Capitanian extinction event, rebuilding complex trophic structures and refilling guilds, diversity and disparity fell further until the

4856:"The appearance of an oxygen-depleted condition on the Capitanian disphotic slope/basin in South China: Middle–Upper Permian stratigraphy at Chaotian in northern Sichuan" 791:

Emeishan Traps may also have contributed to the downfall of the ozone shield, exposing the Earth's surface to a vastly increased flux of high-frequency solar radiation.

3918:

Wang, Wen-qian; Zheng, Feifei; Zhang, Shuang; Cui, Ying; Zheng, Quan-feng; Zhang, Yi-chun; Chang, Dong-xun; Zhang, Hua; Xu, Yi-gang; Shen, Shu-zhong (15 January 2023).

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boundary itself, which is currently estimated to be approximately 259.1 million years old, but is subject to change by the Subcommission on Permian Stratigraphy of the

1963: 3023:"Middle Permian (Capitanian) seawater 87Sr/86Sr minimum coincided with disappearance of tropical biota and reef collapse in NE Japan and Primorye (Far East Russia)" 1357:"High-precision U-Pb CA-TIMS calibration of Middle Permian to Lower Triassic sequences, mass extinction and extreme climate-change in eastern Australian Gondwana" 993:

De la Horra, R.; Galán-Abellán, A. B.; López-Gómez, José; Sheldon, Nathan D.; Barrenechea, J. F.; Luque, F. J.; Arche, A.; Benito, M. I. (August–September 2012).

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bivalves. By virtue of the greater solubility of carbon dioxide in colder waters, ocean acidification was especially lethal in high latitude waters. Furthermore,

4600:

Buatois, Luis A.; Borruel-Abadía, Violeta; De la Horra, Raúl; Galán-Abellán, Ana Belén; López-Gómez, José; Barrenechea, José F.; Arche, Alfredo (25 March 2021).

2243:

The Distribution of the Karroo Vertebrate Fauna: With Special Reference to Certain Genera and the Bearing of this Distribution on the Zoning of the Beaufort Beds

498:

and the overlying assemblages. In both Russia and South Africa, this transition was associated with the extinction of the previously dominant group of therapsid

3747:"The Emeishan large igneous province eruption triggered coastal perturbations and the Capitanian mass extinction: Insights from mercury in Permian bauxite beds" 3671: 3405:"The Emeishan large igneous province eruption triggered coastal perturbations and the Capitanian mass extinction: Insights from mercury in Permian bauxite beds" 550:

In the oceans, the Capitanian extinction event led to high extinction rates among ammonoids, corals and calcareous algal reef-building organisms, foraminifera,

261:. The Capitanian mass extinction greatly reduced disparity (the range of different guilds); eight guilds were lost. It impacted the diversity within individual 4447:"Middle–Upper Permian carbon isotope stratigraphy at Chaotian, South China: Pre-extinction multiple upwelling of oxygen-depleted water onto continental shelf" 2028: 4215: 3672:"Mercury anomalies associated with three extinction events (Capitanian Crisis, Latest Permian Extinction and the Smithian/Spathian Extinction) in NW Pangea" 1712: 4319: 2725: 2153: 1461:"Biodiversity across the Guadalupian-Lopingian Boundary: first results on the ostracod (Crustacea) fauna, Chaotian section (Sichuan Province, South China)" 681:-dominated reefs, which in turn was followed by a return of metazoan, sponge-dominated reefs. Overall, reef recovery took approximately 2.5 million years. 3857:

Jiang, Qiang; Jourdan, Fred; Olierook, Hugo K. H.; Merle, Renaud E.; Bourdet, Julien; Fougerouse, Denis; Godel, Belinda; Walker, Alex T. (25 July 2022).

4216:"The Capitanian (Guadalupian, Middle Permian) mass extinction in NW Pangea (Borup Fiord, Arctic Canada): A global crisis driven by volcanism and anoxia" 1509:"Vertebrate tracks from the Permian of Gonfaron (Provence, Southern France) and their implications for the late Capitanian terrestrial extinction event" 611:

brachiopods made up 99.1% of the individuals found in tropical carbonates in the Western United States, South China and Greece prior to the extinction,

4395: 4263: 3807: 3628: 3465: 3027: 2669: 2626: 2029:"CAPITANIAN (MIDDLE PERMIAN) MASS EXTINCTION AND RECOVERY IN WESTERN TETHYS: A FOSSIL, FACIES, AND δ13C STUDY FROM HUNGARY AND HYDRA ISLAND (GREECE)" 1513: 1308: 1046: 502:, the dinocephalians, which led to its later designation as the dinocephalian extinction. Post-extinction origination rates remained low through the 3079:"Estimating spatial variation in origination and extinction in deep time: a case study using the Permian–Triassic marine invertebrate fossil record" 4894:

Shu-Zhong, Shen; Shi, G. R. (8 April 2016). "Paleobiogeographical extinction patterns of Permian brachiopods in the Asian–western Pacific region".

2892: 4131: 1896: 1866:

CA-TIMS zircon U–Pb dating of felsic ignimbrite from the Binchuan section: Implications for the termination age of Emeishan large igneous province

537:

Assemblage Zone of the Karoo Basin demonstrated that the dinocephalian extinction did occur in the late Capitanian, around 260 million years ago.

2427:"High-precision temporal calibration of Late Permian vertebrate biostratigraphy: U-Pb zircon constraints from the Karoo Supergroup, South Africa" 3182: 2859: 832:

would have arisen as yet another biocidal consequence of the intense sulphur emissions produced by Emeishan Traps volcanism. This resulted in

4656:

Jost, Adam B.; Mundil, Roland; He, Bin; Brown, Shaun T.; Altiner, Demir; Sun, Yadong; DePaolo, Donald J.; Payne, Jonathan L. (15 June 2014).

1916:"The double mass extinction revisited: reassessing the severity, selectivity, and causes of the end-Guadalupian biotic crisis (Late Permian)" 1040:

Huang, Yuangeng; Chen, Zhong-Qiang; Zhao, Laishi; Stanley Jr., George D.; Yan, Jiaxin; Pei, Yu; Yang, Wanrong; Huang, Junhua (1 April 2019).

2668:

Meng, Qi; Xue, Wuqiang; Chen, Fayao; Yan, Jiaxin; Cai, Jiahua; Sun, Yadong; Wignall, Paul B.; Liu, Ke; Liu, Zhichen; Chen, Deng (May 2022).

1042:"Restoration of reef ecosystems following the Guadalupian–Lopingian boundary mass extinction: Evidence from the Laibin area, South China" 3622:

Jerram, Dougal A.; Widdowson, Mike; Wignall, Paul B.; Sun, Yadong; Lai, Xulong; Bond, David P. G.; Torsvik, Trond H. (1 January 2016).

3295:"Global Taxonomic Diversity of Anomodonts (Tetrapoda, Therapsida) and the Terrestrial Rock Record Across the Permian-Triassic Boundary" 3563:

Huang, Hu; Cawood, Peter A.; Hou, Ming-Cai; Yang, Jiang-Hai; Ni, Shi-Jun; Du, Yuan-Sheng; Yan, Zhao-Kun; Wang, Jun (1 November 2016).

2245:. Memoir No. 1. Johannesburg: Bernard Price Institute for Palaeontological Research, University of the Witwatersrand. pp. 1–131. 2228: 1759:

When and how did the terrestrial mid-Permian mass extinction occur? Evidence from the tetrapod record of the Karoo Basin, South Africa

995:"Paleoecological and paleoenvironmental changes during the continental Middle–Late Permian transition at the SE Iberian Ranges, Spain" 3077:

Allen, Bethany J.; Clapham, Matthew E.; Saupe, Erin E.; Wignall, Paul B.; Hill, Daniel J.; Dunhill, Alexander M. (10 February 2023).

2620:

Lai, Xulong; Wang, Wei; Wignall, Paul B.; Bond, David P. G.; Jiang, Haishui; Ali, J. R.; John, E. H.; Sun, Yadong (4 November 2008).

313: 75: 4335:"Collapsed upwelling and intensified euxinia in response to climate warming during the Capitanian (Middle Permian) mass extinction" 4078:"Recycled carbon degassed from the Emeishan plume as the potential driver for the major end-Guadalupian carbon cycle perturbations" 3239:"Delayed recovery of metazoan reefs on the Laibin-Heshan platform margin following the Middle Permian (Capitanian) mass extinction" 1626:

Sepkoski Jr., J. J. (1996). "Patterns of Phanerozoic Extinction: a Perspective from Global Data Bases". In: Walliser, O. H. (Ed.),

578:, 77.8% of coral genera and 82.2% of coral species that were in Permian China were lost during the Capitanian mass extinction. The 394:

show no sign of an extinction event at the end of the Capitanian; the extinction event there is recorded in the middle Capitanian.

375: 224:

extinction event, and only viewed as separate relatively recently, this mass extinction is believed to be the third largest of the

2154:

An abrupt extinction in the Middle Permian (Capitanian) of the Boreal Realm (Spitsbergen) and its link to anoxia and acidification

744:; increased mercury concentrations are coincident with the negative carbon isotope excursion, indicating a common volcanic cause. 2227:

Ivakhnenko, M. F., Golubev, V. K., Gubin, Yu. M., Kalandadze, N. N., Novikov, I. V., Sennikov, A. G. & Rautian, A. S. 1997. "

4220: 1968: 82: 197:

epoch. It is often called the end-Guadalupian extinction event because of its initial recognition between the Guadalupian and

2604: 2189: 1849: 655:

suggests that mid-latitude marine life became affected earlier by the extinction event than marine organisms of the tropics.

266: 146: 89: 3727: 2956:"Contrasting microbial community changes during mass extinctions at the Middle/Late Permian and Permian/Triassic boundaries" 2954:

Xie, Shucheng; Algeo, Thomas J.; Zhou, Wenfeng; Ruan, Xiaoyan; Luo, Genming; Huang, Junhua; Yan, Jiaxin (15 February 2017).

1868: 4662: 4268: 3924: 3521: 3361: 2960: 2452: 205:

study suggests that extinction peaks in many taxonomic groups occurred within the Guadalupian, in the latter half of the

2152:

Bond, D.P.G., Wignall, P.B., Joachimski, M.M., Sun, Y., Savov, I., Grasby, S.E., Beauchamp, B. and Blomeier, D.P. 2015.

4707:

Kévin Rey; Michael O. Day; Romain Amiot; Jean Goedert; Christophe Lécuyer; Judith Sealy; Bruce S. Rubidge (July 2018).

4546:"The implications of the giant bivalve family Alatoconchidae for the end-Guadalupian (Middle Permian) extinction event" 4195: 2448:"Impacts of basin restriction on geochemistry and extinction patterns: A case from the Guadalupian Delaware Basin, USA" 1808:

Radiation and extinction patterns in Permian floras from North China as indicators for environmental and climate change

1566:

Villier, L.; Korn, D. (October 2004). "Morphological Disparity of Ammonoids and the Mark of Permian Mass Extinctions".

1414: 4498:"Permian reefs re-examined: extrinsic control mechanisms of gradual and abrupt changes during 40 my of reef evolution" 3357:"High-resolution δ13Ccarb chemostratigraphy from latest Guadalupian through earliest Triassic in South China and Iran" 2742:

Wang, X.-D. & Sugiyama, T. (December 2000). "Diversity and extinction patterns of Permian coral faunas of China".

2250: 2181: 1841: 1302:

McGhee Jr., George R.; Clapham, Matthew E.; Sheehan, Peter M.; Bottjer, David J.; Droser, Mary L. (15 January 2013).

126: 31: 3003: 513:

in that region, was constrained to below the Illawarra magnetic reversal and therefore had to have occurred in the

405:

of the Maokou Formation, are unique for preserving a mass extinction and the cause of that mass extinction. Large

4451: 2782: 2492: 2329: 503: 233: 110: 4909: 2426: 1662: 1166:"Relationship between extinction magnitude and climate change during major marine and terrestrial animal crises" 3745:

Ling, Kunyue; Wen, Hanjie; Grasby, Stephen E.; Zhao, Haonan; Deng, Changzhou; Yin, Runsheng (5 February 2023).

3461:"End-Guadalupian extinction of larger fusulinids in central Iran and implications for the global biotic crisis" 3403:

Ling, Kunyue; Wen, Hanjie; Grasby, Stephen E.; Zhao, Haonan; Deng, Changzhou; Yin, Runsheng (5 February 2023).

492: 4658:"Constraining the cause of the end-Guadalupian extinction with coupled records of carbon and calcium isotopes" 3515:

He, Bin; Xu, Yi-Gang; Huang, Xiao-Long; Luo, Zhen-Yu; Shi, Yu-Ruo; Yun, Qi-Jun; Yu, Song-Yue (30 March 2007).

485:

Superassemblage, respectively. In South Africa, this corresponded to the boundary between the variously named

4854:

Saitoh, Masamufi; Isozaki, Yukio; Yao, Jianxin; Ji, Zhansheng; Ueno, Yuichiro; Yoshida, Naohiro (June 2015).

2381:

Golubev, V. K. (2015). "Dinocephalian stage in the history of the Permian tetrapod fauna of Eastern Europe".

1757:

Day, M.O., Ramezani, J., Bowring, S.A., Sadler, P.M., Erwin, D.H., Abdala, F. and Rubidge, B.S., July 2015. "

799:

Global warming resulting from the large igneous province's activity has been implicated as a cause of marine

152: 118: 23: 4445:

Saitoh, Masafumi; Isozaki, Yukio; Ueno, Yuichiro; Yoshida, Naohiro; Yao, Jianxin; Ji, Zhansheng (May 2013).

2536:"Guadalupian (Middle Permian) ocean redox evolution in South China and its implications for mass extinction" 312:. The loss of marine invertebrates during the Capitanian mass extinction was comparable in magnitude to the 4709:"Stable isotope record implicates aridification without warming during the late Capitanian mass extinction" 1711:

Retallack, G. J., Metzger, C. A., Greaver, T., Jahren, A. H., Smith, R. M. H. & Sheldon, N. D. (2006).

130: 35: 4855: 4807: 4708: 4657: 4497: 4446: 4390: 4334: 4022:

Sheldon, Nathan D.; Chakrabarti, Ramananda; Retallack, Gregory J.; Smith, Roger M. H. (20 February 2014).

3919: 3803:"High-temperature combustion event spanning the Guadalupian−Lopingian boundary terminated by soil erosion" 3802: 3746: 3623: 3564: 3516: 3460: 3404: 3356: 3238: 3022: 2955: 2621: 2535: 2487: 1508: 1356: 1303: 1097: 1041: 994: 509:

for at least 1 million years, which suggests that there was a delayed recovery of Karoo Basin ecosystems.

185:

that predated the end-Permian extinction event. The mass extinction occurred during a period of decreased

4860: 3243: 3183:"Ecological consequences of the Guadalupian extinction and its role in the brachiopod-mollusk transition" 999: 123: 28: 2327:

Lucas, S. G. (2009). "Timing and magnitude of tetrapod extinctions across the Permo-Triassic boundary".

131: 36: 4945: 4769: 1264:"The Hirnantian (Late Ordovician) and end-Guadalupian (Middle Permian) mass-extinction events compared" 4077: 2208: 824:, which probably contributed to the demise of various calcareous marine organisms, particularly giant 2229:

Permskiye I Triasovyye tetrapody vostochnoi Evropy (Permian and Triassic Tetrapods of Eastern Europe)

1215: 406: 129: 128: 34: 33: 4132:"Climate changes caused by degassing of sediments during the emplacement of large igneous provinces" 2871: 296:

genera between 35 and 47%, while an estimate published in 2016 suggested a loss of 33–35% of marine

125: 30: 4765:"The Late Capitanian Mass Extinction of Terrestrial Vertebrates in the Karoo Basin of South Africa" 4076:

Zhu, Jiang; Zhang, Zhaochong; Santosh, M.; Tan, Shucheng; Deng, Yinan; Xie, Qiuhong (1 July 2021).

2622:"Palaeoenvironmental change during the end-Guadalupian (Permian) mass extinction in Sichuan, China" 2367:

The Pristerognathus AZ and the aftermath of the Capitanian extinction event in the main Karoo Basin

2207:

Bond, D.P.G.; Hilton, J.; Wignall, P.B.; Ali, J.R.; Steven, L.G.; Sun, Y.; Lai, X. September 2010.

590: 124: 122: 103: 29: 27: 132: 127: 37: 32: 3670:

Grasby, Stephen E.; Beauchamp, Benoit; Bond, David P. G.; Wignall, Paul B.; Sanei, Hamed (2016).

3021:

Kani, Tomomi; Isozaki, Yukio; Hayashi, Ryutaro; Zakharov, Yuri; Popov, Alexander (15 June 2018).

2912: 2447: 2383: 2304:

Watson, D. M. S. May 1914. "The Zones of the Beaufort Beds of the Karoo System in South Africa."

1784: 1459:

Zazzali, Sindbad; Crasquin, Sylvie; Deconinck, Jean-François; Feng, Qinglai (25 September 2015).

877: 800: 526: 305: 1758: 54: 4896: 4024:"Contrasting geochemical signatures on land from the Middle and Late Permian extinction events" 3980: 3187: 3139: 3083: 2534:

Wei, Hengye; Tang, Zhanwen; Yan, Detian; Wang, Jianguo; Roberts, Andrew P. (30 December 2019).

1964:"Mass extinction or extirpation: Permian biotic turnovers in the northwestern margin of Pangea" 1920: 1649: 916: 732: 3859:"Volume and rate of volcanic CO2 emissions governed the severity of past environmental crises" 2267:

Vertebrate biozonation of the Beaufort Group with special reference to the Western Karoo Basin

1962:

Lee, Sangmin; Shi, Guang R.; Nakrem, Hans A.; Woo, Jusun; Tazawa, Jun-Ichi (4 February 2022).

662:

This mass extinction marked the beginning of the transition between the Palaeozoic and Modern

209:

age. The extinction event has been argued to have begun around 262 million years ago with the

4812: 4601: 4313: 2886: 2719: 2674: 338: 301: 246: 4675: 4281: 3975: 3937: 3534: 3374: 2973: 2091:"Volcanism, Mass Extinction, and Carbon Isotope Fluctuations in the Middle Permian of China" 1897:"Sixth extinction, rivaling that of the dinosaurs, should join the big five, scientists say" 1263: 4955: 4821: 4778: 4722: 4671: 4615: 4606: 4559: 4511: 4456: 4404: 4389:

Zhang, Bolin; Yao, Suping; Hu, Wenxuan; Ding, Hai; Liu, Bao; Ren, Yongle (1 October 2019).

4348: 4277: 4145: 4089: 3933: 3872: 3816: 3760: 3685: 3637: 3578: 3530: 3474: 3418: 3370: 3308: 3252: 3196: 3092: 3036: 2969: 2791: 2753: 2683: 2635: 2549: 2501: 2446:

Smith, Benjamin P.; Larson, Toti; Martindale, Rowan C.; Kerans, Charles (15 January 2020).

2392: 2338: 2104: 2042: 1977: 1577: 1522: 1370: 1317: 1179: 1055: 1004: 948: 720:

It is believed that the extinction, which coincided with the beginning of a major negative

4264:"Global oceanic anoxia linked with the Capitanian (Middle Permian) marine mass extinction" 8: 4950: 3676: 2425:

Rubidge, B. S., Erwin, D. H., Ramezani, J., Bowring, S. A. & De Klerk, W. J. (2013).

821: 766: 762: 293: 262: 66: 4825: 4782: 4726: 4619: 4563: 4515: 4460: 4408: 4352: 4149: 4093: 4023: 3876: 3820: 3764: 3689: 3641: 3582: 3478: 3422: 3312: 3256: 3200: 3096: 3040: 2821: 2795: 2757: 2687: 2639: 2553: 2505: 2396: 2342: 2108: 2046: 1981: 1581: 1526: 1374: 1321: 1183: 1059: 1008: 952: 4921: 4913: 4738: 4631: 4575: 4550: 4420: 4364: 4293: 4237: 4188:

The Worst of Times: How Life on Earth Survived Eighty Million Years of Mass Extinctions

4051: 3997: 3949: 3895: 3858: 3832: 3776: 3701: 3490: 3434: 3331: 3294: 3268: 3212: 3156: 3135:"Latitudinal selectivity of foraminifer extinctions during the late Guadalupian crisis" 3108: 3052: 2929: 2699: 2565: 2408: 2128: 2058: 2001: 1937: 1666: 1609: 1538: 1482: 1436: 1409: 1237: 1220: 1071: 972: 897: 881: 863: 833: 663: 644: 632: 608: 424: 402: 213:, though its most intense pulse occurred 259 million years ago in what is known as the 4833: 4545: 4523: 3976:"Methane Release from Igneous Intrusion of Coal during Late Permian Extinction Events" 2860:"Decoupled diversity and ecology during the end-Guadalupian extinction (late Permian)" 2090: 1216:"The end-Guadalupian (259.8 Ma) biodiversity crisis: the sixth major mass extinction?" 849:

Not all studies, however, have supported the volcanic warming hypothesis; analysis of

453:

losses occurred in the middle of the Capitanian stage. The extinction suffered by the

409:

occurred when the Emeishan Traps first started to erupt, leading to the extinction of

4742: 4713: 4635: 4579: 4424: 4368: 4339: 4297: 4241: 4191: 4161: 4136: 4105: 4055: 3953: 3900: 3836: 3780: 3565:"Silicic ash beds bracket Emeishan Large Igneous province to < 1 m.y. at ~ 260 Ma" 3494: 3438: 3336: 3272: 3160: 3112: 3056: 2933: 2703: 2600: 2569: 2412: 2365: 2246: 2185: 2132: 2120: 2095: 2062: 2005: 1845: 1806:

Stevens, L.G., Hilton, J., Bond, D.P.G., Glasspool, I.J. & Jardine, P.E., 2011. "

1601: 1593: 1568: 1542: 1486: 1441: 1361: 1241: 1075: 964: 893: 741: 575: 574:, suffered a selective extinction pulse at the end of the Capitanian. 75.6% of coral 250: 4925: 4873: 4333:

Zhang, Bolin; Wignall, Paul B.; Yao, Suping; Hu, Wenxuan; Liu, Biao (January 2021).

4001: 3705: 3264: 3216: 2695: 1941: 1807: 1670: 1613: 1016: 976: 4905: 4869: 4829: 4786: 4730: 4679: 4623: 4567: 4519: 4464: 4412: 4356: 4285: 4229: 4153: 4097: 4043: 4035: 3989: 3941: 3890: 3880: 3824: 3768: 3751: 3693: 3645: 3594: 3586: 3569: 3538: 3482: 3426: 3409: 3378: 3326: 3316: 3260: 3204: 3148: 3100: 3044: 2977: 2921: 2799: 2761: 2691: 2643: 2557: 2540: 2509: 2461: 2400: 2346: 2112: 2050: 1993: 1985: 1929: 1658: 1585: 1530: 1474: 1431: 1423: 1378: 1325: 1277: 1229: 1187: 1165: 1111: 1063: 1012: 956: 939: 868: 640: 597:

disappeared over a period of tens of thousands of years; though new brachiopod and

326: 289: 258: 221: 186: 182: 3772: 3430: 2561: 1233: 1152: 292:

is still heavily debated by palaeontologists. Early estimates indicated a loss of

4468: 4416: 3828: 3649: 3590: 3486: 3321: 3048: 2803: 2647: 2513: 2350: 1872: 1534: 1329: 1115: 1067: 912: 628: 519: 317: 1631: 4706: 4683: 4289: 3945: 3864:

Proceedings of the National Academy of Sciences of the United States of America

3542: 3382: 3152: 2981: 2765: 2465: 2054: 1865: 1739: 1689:

Proceedings of the National Academy of Sciences of the United States of America

1685:"Estimates of the magnitudes of major marine mass extinctions in earth history" 1170: 825: 787: 783: 754: 750: 729: 570:, which had been in a long-term decline for a 30 million year period since the 559: 555: 398: 391: 354: 254: 190: 4791: 4764: 4101: 3920:"Ecosystem responses of two Permian biocrises modulated by CO2 emission rates" 3697: 2404: 4939: 4734: 4599: 4360: 4165: 4109: 1597: 1465: 1382: 908: 603: 579: 478: 467: 383: 371: 202: 3885: 2312: 2293: 2116: 1684: 1644: 1589: 1304:"A new ecological-severity ranking of major Phanerozoic biodiversity crises" 1192: 3904: 3340: 2124: 1605: 1445: 1427: 968: 758: 725: 487: 442: 4214:

Bond, David P. G.; Wignall, Paul B.; Grasby, Stephen E. (30 August 2019).

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The Middle Permian (Capitanian) mass extinction on land and in the oceans

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anomalies coeval with mercury and coronene anomalies. A large amount of

4917: 3598: 3104: 1914:

Clapham, Matthew E.; Shen, Shuzhong; Bottjer, David J. (8 April 2016).

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Broom, R. 1906. "On the Permian and Triassic Faunas of South Africa."

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location of the Emeishan Traps, leading to sudden global cooling and

694: 690: 668: 620: 583: 567: 454: 429: 322: 198: 4571: 1933: 1350: 1348: 1346: 345:, Day and colleagues suggested a 74–80% loss of generic richness in 149:, with just over 35% (according to this source) failing to survive. 4186:

Wignall, Paul B. (29 September 2015). "Extinction in the Shadows".

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Kaiho, Kunio; Grasby, Stephen E.; Chen, Zhong-Qiang (15 May 2023).

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Kofukuda, Daisuke; Isozaki, Yukio; Igo, Hisayoshi (15 March 2014).

745: 689:

Among terrestrial vertebrates, the main victims were dinocephalian

652: 551: 346: 276: 4544:

Chen, Fayao; Xue, Wuqiang; Yan, Jiaxin; Meng, Qi (19 April 2021).

1645:"Origination, extinction, and mass depletions of marine diversity" 4502: 2744: 2033: 1343: 1268: 858: 804: 779: 770: 636: 612: 598: 571: 514: 499: 387: 379: 229: 4021: 854: 850: 721: 1301: 736: 699: 316:. Some studies have considered it the third or fourth greatest 297: 1410:"Recovery from the most profound mass extinction of all time" 1354: 844: 648: 450: 417: 270: 141: 2596:

Volcanism, Impacts, and Mass Extinctions: Causes and Effects

2177:

Volcanism, Impacts, and Mass Extinctions: Causes and Effects

1837:

Volcanism, Impacts, and Mass Extinctions: Causes and Effects

1458: 876:

extinction rates in the Karoo Basin, specifically the upper

253:

rose during the Capitanian. This was probably the result of

4910:

10.1666/0094-8373(2002)028<0449:PEPOPB>2.0.CO;2

3174: 2445: 1663:

10.1666/0094-8373(2004)030<0522:OEAMDO>2.0.CO;2

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Retallack, Gregory J.; Jahren, A. Hope (1 October 2007).

3856: 3728:"Brachiopod die-off signaled mid-Permian mass extinction" 3621: 1214:

Rampino, Michael R.; Shen, Shu-Zhong (5 September 2019).

627:

Most of the marine victims of the extinction were either

4444: 4209: 4207: 3076: 1864:

Zhong, Y.-T., He, B., Mundil, R., and Xu, Y.-G. (2014).

1039: 4763:

Day, Michael O.; Rubidge, Bruce S. (18 February 2021).

3459:

Arefifard, Sakineh; Payne, Jonathan L. (15 July 2020).

2741: 1643:

Bambach, R. K.; Knoll, A. H. & Wang, S. C. (2004).

1628:

Global Events and Event Stratigraphy in the Phanerozoic

936: 838: 892:

The Capitanian mass extinction has been attributed to

4204: 4190:. Princeton: Princeton University Press. p. 38. 2864:

Geological Society of America Abstracts with Programs

4130:

Ganino, Clément; Arndt, Nicholas T. (1 April 2009).

3617: 3615: 3133:

Bond, David P. G.; Wignall, Paul B. (8 April 2016).

2822:"New mass extinction event identified by geologists" 2370:. 35th International Geological Congress. Cape Town. 1785:"South Africa's Great Karoo reveals mass extinction" 3353: 2025: 1262:Isozaki, Yukio; Servais, Thomas (8 December 2017). 988: 986: 220:Having historically been considered as part of the 189:and increased extinction rates near the end of the 4853: 4537: 4075: 3562: 3452: 2858:Clapham, M.E., Bottjer, D.J. and Shen, S. (2006). 2778: 2619: 2485: 2087: 1913: 1035: 1033: 4655: 4396:Palaeogeography, Palaeoclimatology, Palaeoecology 4332: 4261: 4213: 3917: 3808:Palaeogeography, Palaeoclimatology, Palaeoecology 3744: 3629:Palaeogeography, Palaeoclimatology, Palaeoecology 3612: 3466:Palaeogeography, Palaeoclimatology, Palaeoecology 3402: 3028:Palaeogeography, Palaeoclimatology, Palaeoecology 2627:Palaeogeography, Palaeoclimatology, Palaeoecology 2599:. The Geological Society of America. p. 38. 1890: 1888: 1733:Palaeogeography, Palaeoclimatology, Palaeoecology 1514:Palaeogeography, Palaeoclimatology, Palaeoecology 1309:Palaeogeography, Palaeoclimatology, Palaeoecology 1145:Palaeogeography, Palaeoclimatology, Palaeoecology 1047:Palaeogeography, Palaeoclimatology, Palaeoecology 724:excursion signifying a severe disturbance of the 353:in South Africa, including the extinction of the 288:The impact of the Capitanian extinction event on 4937: 4318:: CS1 maint: bot: original URL status unknown ( 3850: 2953: 2906:Shen, Shu-Zhong; Zhang, Yi-Chun (14 July 2015). 2724:: CS1 maint: bot: original URL status unknown ( 2706:. Archived from the original on 22 December 2022 1707: 1705: 1703: 1701: 983: 810: 341:exist for the Capitanian mass extinction. Among 4491: 4489: 4487: 4485: 4179: 3973: 3800: 3236: 3014: 3004:"Two Phases of the End-Permian Mass Extinction" 2588: 2586: 2271:Annals of the Geological Survey of South Africa 1961: 1829: 1827: 1825: 1823: 1779: 1777: 1153:https://dx.doi.org/10.1016/j.palaeo.2010.03.056 1030: 121: 26: 4806:Hallam, A.; Wignall, Paul B. (December 1999). 4543: 4388: 3458: 2737: 2735: 2667: 2533: 2380: 2083: 2081: 2079: 1885: 1261: 1096:Shen, Shu-Zhong; Shi, G. R. (September 2009). 457:may have occurred in the early Wuchiapingian. 171:Guadalupian-Lopingian boundary mass extinction 4805: 4300:. Archived from the original on 22 April 2023 2593:Keller, Gerta; Kerr, Andrew C., eds. (2014). 2174:Keller, Gerta; Kerr, Andrew C., eds. (2014). 1834:Keller, Gerta; Kerr, Andrew C., eds. 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" 2203: 2201: 2076: 1802: 1800: 1798: 1565: 1403: 1401: 1399: 1213: 4762: 2592: 2322: 2320: 2173: 1833: 1753: 1751: 1749: 1747: 845:Criticism of the volcanic cause hypothesis 616:that of the later end-Permian extinction. 337:Few published estimates for the impact on 4790: 3894: 3884: 3330: 3320: 3008:Pangea: Global Environments and Resources 2905: 2313:https://doi.org/10.1017/S001675680019675X 2294:https://doi.org/10.1017/S001675680012271X 2148: 2146: 2144: 2142: 1556: 1435: 1191: 332: 308:and clustering of extinctions in certain 4808:"Mass extinctions and sea-level changes" 3725: 3181:Clapham, Matthew E. (24 February 2015). 2815: 2813: 2240: 2198: 1795: 1396: 1138: 1136: 1134: 1132: 639:that died when the seas closed, or were 376:International Commission on Stratigraphy 4185: 3180: 2819: 2317: 1744: 1095: 820:carbon dioxide in the oceans triggered 589:87% of brachiopod species found at the 329:or even entire ecosystems themselves). 4938: 4221:Geological Society of America Bulletin 2838: 2231:". GEOS, Moscow (original in Russian). 2158:Geological Society of America Bulletin 2139: 1969:Geological Society of America Bulletin 1770:https://doi.org/10.1098/rspb.2015.0834 1717:Geological Society of America Bulletin 1630:. Springer-Verlag, Berlin, pp. 35–51. 401:, which are interbedded with tropical 80: 73: 64: 59: 2810: 2363: 1163: 1129: 794: 715: 314:Cretaceous–Paleogene extinction event 108: 101: 94: 87: 4119:– via Elsevier Science Direct. 2475:– via Elsevier Science Direct. 1894: 930: 728:, was triggered by eruptions of the 460: 283: 215:Guadalupian-Lopingian boundary event 52: 44: 4663:Earth and Planetary Science Letters 4269:Earth and Planetary Science Letters 3925:Earth and Planetary Science Letters 3522:Earth and Planetary Science Letters 3362:Earth and Planetary Science Letters 2961:Earth and Planetary Science Letters 2453:Earth and Planetary Science Letters 903: 684: 647:. Evidence from marine deposits in 46:Marine extinction intensity during 13: 1763:Proceedings of the Royal Society B 1415:Proceedings of the Royal Society B 540: 117: 22: 14: 4967: 2182:The Geological Society of America 1842:The Geological Society of America 1812:Journal of the Geological Society 1408:Sahney, S.; Benton, M.J. 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Archived from 1383:10.1016/j.gr.2014.09.002 591:Kapp Starostin Formation 481:Superassemblage and the 477:Superzone and later the 4676:2014E&PSL.396..201J 4282:2023E&PSL.61018128S 3938:2023E&PSL.60217940W 3886:10.1073/pnas.2202039119 3535:2007E&PSL.255..306H 3375:2013E&PSL.375..156S 2974:2017E&PSL.460..180X 2913:Journal of Paleontology 2384:Paleontological Journal 2117:10.1126/science.1171956 1590:10.1126/science.1102127 1193:10.5194/bg-19-3369-2022 878:Abrahamskraal Formation 265:more severely than the 211:Late Guadalupian crisis 3981:The Journal of Geology 1683:Stanley, S. M., 2016. 1428:10.1098/rspb.2007.1370 917:biological competition 857:values from the tooth 733:large igneous province 339:terrestrial ecosystems 333:Terrestrial ecosystems 135: 40: 4813:Earth-Science Reviews 4496:Weidlich, O. (2002). 3010:(Memoir 17): 813–822. 2675:Earth-Science Reviews 2213:Earth-Science Reviews 525:radiometric ages via 397:The volcanics of the 302:background extinction 153:source and image info 134: 61:Millions of years ago 39: 4082:Geoscience Frontiers 1738:(3-4), pp. 289–297. 1151:(1-2), pp. 282-294. 767:Southern Hemispheres 582:, a family of large 520:biostratigraphically 245:In the aftermath of 193:, also known as the 175:pre-Lopingian crisis 165:, also known as the 4826:1999ESRv...48..217H 4783:2021FrEaS...9...15D 4727:2018GondR..59....1R 4620:2021TeNov..33..455B 4564:2021GeolJ..56.6073C 4516:2002Geobi..35..287W 4461:2013JAESc..67...51S 4409:2019PPP...53108630Z 4353:2021GondR..89...31Z 4150:2009Geo....37..323G 4094:2021GeoFr..1201140Z 3877:2022PNAS..11902039J 3871:(31): e2202039119. 3821:2023PPP...61811518K 3765:2023ChGeo.61721243L 3690:2016GeoM..153..285G 3677:Geological Magazine 3642:2016PPP...441...65J 3583:2016Litho.264...17H 3479:2020PPP...55009743A 3423:2023ChGeo.61721243L 3313:2008PLoSO...3.3733F 3257:2019GPC...180....1W 3209:10.1017/pab.2014.15 3201:2015Pbio...41..266C 3097:2023Pbio...49..509A 3041:2018PPP...499...13K 2845:The Washington Post 2796:2006JAESc..26..353O 2758:2000Letha..33..285W 2688:2022ESRv..22804011M 2640:2008PPP...269...78L 2554:2019ChGeo.53019318W 2506:2014JAESc..82...47K 2397:2015PalJ...49.1346G 2343:2009JAESc..36..491L 2306:Geological Magazine 2287:Geological Magazine 2109:2009Sci...324.1179W 2103:(5931): 1179–1182. 2047:2012Palai..27...78W 1982:2022GSAB..134.2399L 1976:(9–10): 2399–2414. 1722:(11-12): 1398-1411. 1582:2004Sci...306..264V 1527:2022PPP...59911043M 1375:2015GondR..28...61M 1322:2013PPP...370..260M 1184:2022BGeo...19.3369K 1060:2019PPP...519....8H 1009:2012GPC....94...46D 961:10.1038/nature03339 953:2005Natur.434..208R 900:biodiversity loss. 867:specimens from the 822:ocean acidification 664:evolutionary faunas 633:epicontinental seas 527:uranium–lead dating 491:, Dinocephalian or 403:carbonate platforms 306:Signor–Lipps effect 300:when corrected for 294:marine invertebrate 4551:Geological Journal 3105:10.1017/pab.2023.1 2311:(5), pp. 203-208. 2184:. pp. 36–37. 2164:(9-10): 1411-1421. 1871:2019-06-24 at the 1695:(42), E6325–E6334. 1221:Historical Biology 882:Teekloof Formation 864:Diictodon feliceps 834:soil acidification 795:Anoxia and euxinia 716:Volcanic emissions 471:Superzone and the 423:In the absence of 257:replacing extinct 247:Olson's Extinction 136: 41: 4946:Extinction events 4714:Gondwana Research 4628:10.1111/ter.12530 4340:Gondwana Research 4158:10.1130/G25325A.1 4040:10.1111/sed.12117 2606:978-0-8137-2505-5 2391:(12): 1346–1352. 2191:978-0-8137-2505-5 1851:978-0-8137-2505-5 1576:(5694): 264–266. 1479:10.5252/g2015n3a1 1422:(1636): 759–765. 1362:Gondwana Research 1282:10.1111/let.12252 1178:(14): 3369–3380. 947:(7030): 209–210. 645:Paleotethys Ocean 461:Terrestrial realm 413:foraminifera and 327:ecological niches 290:marine ecosystems 284:Marine ecosystems 69: 4963: 4930: 4929: 4891: 4885: 4884: 4882: 4880: 4851: 4845: 4844: 4842: 4840: 4803: 4797: 4796: 4794: 4760: 4754: 4753: 4751: 4749: 4704: 4695: 4694: 4692: 4690: 4653: 4647: 4646: 4644: 4642: 4597: 4591: 4590: 4588: 4586: 4558:(2): 6073–6087. 4541: 4535: 4534: 4532: 4530: 4493: 4480: 4479: 4477: 4475: 4455:. 67–68: 51–62. 4442: 4436: 4435: 4433: 4431: 4386: 4380: 4379: 4377: 4375: 4330: 4324: 4323: 4317: 4309: 4307: 4305: 4259: 4253: 4252: 4250: 4248: 4234:10.1130/B35281.1 4228:(5–6): 931–942. 4211: 4202: 4201: 4183: 4177: 4176: 4174: 4172: 4127: 4121: 4120: 4118: 4116: 4073: 4067: 4066: 4064: 4062: 4034:(6): 1812–1829. 4019: 4013: 4012: 4010: 4008: 3971: 3965: 3964: 3962: 3960: 3915: 3909: 3908: 3898: 3888: 3854: 3848: 3847: 3845: 3843: 3798: 3792: 3791: 3789: 3787: 3752:Chemical Geology 3742: 3736: 3735: 3723: 3717: 3716: 3714: 3712: 3667: 3661: 3660: 3658: 3656: 3619: 3610: 3609: 3607: 3605: 3560: 3554: 3553: 3551: 3549: 3529:(3–4): 306–323. 3512: 3506: 3505: 3503: 3501: 3456: 3450: 3449: 3447: 3445: 3410:Chemical Geology 3400: 3394: 3393: 3391: 3389: 3351: 3345: 3344: 3334: 3324: 3290: 3284: 3283: 3281: 3279: 3234: 3228: 3227: 3225: 3223: 3178: 3172: 3171: 3169: 3167: 3130: 3124: 3123: 3121: 3119: 3074: 3068: 3067: 3065: 3063: 3018: 3012: 3011: 2999: 2993: 2992: 2990: 2988: 2951: 2945: 2944: 2942: 2940: 2926:10.1666/07-118.1 2903: 2897: 2896: 2890: 2882: 2880: 2879: 2855: 2849: 2848: 2836: 2830: 2829: 2817: 2808: 2807: 2790:(3–4): 353–368. 2776: 2770: 2769: 2739: 2730: 2729: 2723: 2715: 2713: 2711: 2665: 2659: 2658: 2656: 2654: 2617: 2611: 2610: 2590: 2581: 2580: 2578: 2576: 2541:Chemical Geology 2531: 2525: 2524: 2522: 2520: 2483: 2477: 2476: 2474: 2472: 2443: 2437: 2423: 2417: 2416: 2378: 2372: 2371: 2361: 2355: 2354: 2324: 2315: 2302: 2296: 2292:(1), pp. 29-30. 2283: 2277: 2263: 2257: 2256: 2238: 2232: 2225: 2219: 2205: 2196: 2195: 2171: 2165: 2150: 2137: 2136: 2085: 2074: 2073: 2071: 2069: 2023: 2017: 2016: 2014: 2012: 1990:10.1130/B36227.1 1959: 1953: 1952: 1950: 1948: 1911: 1905: 1904: 1892: 1883: 1862: 1856: 1855: 1831: 1818: 1804: 1793: 1792: 1781: 1772: 1755: 1742: 1729: 1723: 1709: 1696: 1681: 1675: 1674: 1640: 1634: 1624: 1618: 1617: 1563: 1554: 1553: 1551: 1549: 1504: 1498: 1497: 1495: 1493: 1456: 1450: 1449: 1439: 1405: 1394: 1393: 1391: 1389: 1352: 1341: 1340: 1338: 1336: 1299: 1293: 1292: 1290: 1288: 1259: 1253: 1252: 1250: 1248: 1211: 1205: 1204: 1202: 1200: 1195: 1161: 1155: 1140: 1127: 1126: 1124: 1122: 1110:(2–3): 152–161. 1093: 1087: 1086: 1084: 1082: 1037: 1028: 1027: 1025: 1023: 1003:. 94–95: 46–61. 990: 981: 980: 934: 904:Other hypotheses 869:Karoo Supergroup 776: 685:Terrestrial life 641:dominant species 609:rhynchonelliform 524: 523:well-constrained 425:radiometric ages 279: 187:species richness 183:extinction event 113: 106: 99: 92: 85: 78: 71: 67: 62: 57: 56: 50: 4971: 4970: 4966: 4965: 4964: 4962: 4961: 4960: 4936: 4935: 4934: 4933: 4892: 4888: 4878: 4876: 4852: 4848: 4838: 4836: 4804: 4800: 4761: 4757: 4747: 4745: 4705: 4698: 4688: 4686: 4654: 4650: 4640: 4638: 4598: 4594: 4584: 4582: 4572:10.1002/gj.4151 4542: 4538: 4528: 4526: 4494: 4483: 4473: 4471: 4443: 4439: 4429: 4427: 4387: 4383: 4373: 4371: 4331: 4327: 4311: 4310: 4303: 4301: 4260: 4256: 4246: 4244: 4212: 4205: 4198: 4184: 4180: 4170: 4168: 4128: 4124: 4114: 4112: 4074: 4070: 4060: 4058: 4020: 4016: 4006: 4004: 3972: 3968: 3958: 3956: 3916: 3912: 3855: 3851: 3841: 3839: 3799: 3795: 3785: 3783: 3743: 3739: 3724: 3720: 3710: 3708: 3668: 3664: 3654: 3652: 3620: 3613: 3603: 3601: 3561: 3557: 3547: 3545: 3513: 3509: 3499: 3497: 3457: 3453: 3443: 3441: 3401: 3397: 3387: 3385: 3352: 3348: 3291: 3287: 3277: 3275: 3235: 3231: 3221: 3219: 3179: 3175: 3165: 3163: 3131: 3127: 3117: 3115: 3075: 3071: 3061: 3059: 3019: 3015: 3000: 2996: 2986: 2984: 2952: 2948: 2938: 2936: 2904: 2900: 2884: 2883: 2877: 2875: 2856: 2852: 2837: 2833: 2818: 2811: 2777: 2773: 2740: 2733: 2717: 2716: 2709: 2707: 2666: 2662: 2652: 2650: 2618: 2614: 2607: 2591: 2584: 2574: 2572: 2532: 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4481: 4437: 4381: 4325: 4254: 4203: 4197:978-0691142098 4196: 4178: 4144:(4): 323–326. 4122: 4068: 4048:2027.42/108696 4014: 3994:10.1086/524120 3966: 3910: 3849: 3793: 3737: 3732:Earth Magazine 3718: 3684:(2): 285–297. 3662: 3611: 3555: 3507: 3451: 3395: 3346: 3285: 3229: 3195:(2): 266–279. 3173: 3147:(4): 465–483. 3125: 3091:(3): 509–526. 3069: 3013: 2994: 2946: 2920:(5): 924–937. 2898: 2850: 2831: 2809: 2771: 2752:(4): 285–294. 2731: 2660: 2634:(1–2): 78–93. 2612: 2605: 2582: 2526: 2478: 2438: 2418: 2373: 2356: 2337:(6): 491–502. 2316: 2297: 2278: 2258: 2251: 2233: 2220: 2197: 2190: 2166: 2138: 2075: 2018: 1954: 1906: 1884: 1857: 1850: 1844:. p. 37. 1819: 1817:, pp. 607–619. 1794: 1773: 1743: 1724: 1697: 1676: 1657:(4): 522–542. 1635: 1619: 1555: 1499: 1473:(3): 283–313. 1451: 1395: 1342: 1294: 1276:(2): 173–186. 1254: 1228:(5): 716–722. 1206: 1171:Biogeosciences 1156: 1128: 1088: 1029: 982: 928: 927: 925: 922: 905: 902: 894:sea level fall 889: 888:Sea level fall 886: 846: 843: 812: 809: 796: 793: 751:carbon dioxide 730:Emeishan Traps 717: 714: 712: 711:Emeishan Traps 709: 707: 704: 686: 683: 674:Diplosphaerina 560:photosymbiotic 556:photosynthesis 547: 544: 542: 539: 535:Tapinocephalus 494:Tapinocephalus 462: 459: 438: 435: 399:Emeishan Traps 366: 363: 355:dinocephalians 334: 331: 285: 282: 242: 239: 191:Middle Permian 139: 116: 43: 21: 20: 19: 18: 15: 9: 6: 4: 3: 2: 4968: 4957: 4954: 4952: 4949: 4947: 4944: 4943: 4941: 4927: 4923: 4919: 4915: 4911: 4907: 4903: 4899: 4898: 4890: 4875: 4871: 4867: 4863: 4862: 4857: 4850: 4835: 4831: 4827: 4823: 4819: 4815: 4814: 4809: 4802: 4793: 4788: 4784: 4780: 4776: 4772: 4771: 4766: 4759: 4744: 4740: 4736: 4732: 4728: 4724: 4720: 4716: 4715: 4710: 4703: 4701: 4685: 4681: 4677: 4673: 4669: 4665: 4664: 4659: 4652: 4637: 4633: 4629: 4625: 4621: 4617: 4613: 4609: 4608: 4603: 4596: 4581: 4577: 4573: 4569: 4565: 4561: 4557: 4553: 4552: 4547: 4540: 4525: 4521: 4517: 4513: 4509: 4505: 4504: 4499: 4492: 4490: 4488: 4486: 4470: 4466: 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1995: 1991: 1987: 1983: 1979: 1975: 1971: 1970: 1965: 1958: 1943: 1939: 1935: 1931: 1927: 1923: 1922: 1917: 1910: 1902: 1898: 1891: 1889: 1881: 1878: 1874: 1870: 1867: 1861: 1853: 1847: 1843: 1839: 1838: 1830: 1828: 1826: 1824: 1816: 1813: 1809: 1803: 1801: 1799: 1790: 1786: 1780: 1778: 1771: 1767: 1764: 1760: 1754: 1752: 1750: 1748: 1741: 1737: 1734: 1728: 1721: 1718: 1714: 1708: 1706: 1704: 1702: 1694: 1690: 1686: 1680: 1672: 1668: 1664: 1660: 1656: 1652: 1651: 1646: 1639: 1633: 1629: 1623: 1615: 1611: 1607: 1603: 1599: 1595: 1591: 1587: 1583: 1579: 1575: 1571: 1570: 1562: 1560: 1544: 1540: 1536: 1532: 1528: 1524: 1520: 1516: 1515: 1510: 1503: 1488: 1484: 1480: 1476: 1472: 1468: 1467: 1466:Geodiversitas 1462: 1455: 1447: 1443: 1438: 1433: 1429: 1425: 1421: 1417: 1416: 1411: 1404: 1402: 1400: 1384: 1380: 1376: 1372: 1368: 1364: 1363: 1358: 1351: 1349: 1347: 1331: 1327: 1323: 1319: 1315: 1311: 1310: 1305: 1298: 1283: 1279: 1275: 1271: 1270: 1265: 1258: 1243: 1239: 1235: 1231: 1227: 1223: 1222: 1217: 1210: 1194: 1189: 1185: 1181: 1177: 1173: 1172: 1167: 1160: 1154: 1150: 1146: 1139: 1137: 1135: 1133: 1117: 1113: 1109: 1105: 1104: 1099: 1092: 1077: 1073: 1069: 1065: 1061: 1057: 1053: 1049: 1048: 1043: 1036: 1034: 1018: 1014: 1010: 1006: 1002: 1001: 996: 989: 987: 978: 974: 970: 966: 962: 958: 954: 950: 946: 942: 941: 933: 929: 921: 918: 914: 910: 909:Global drying 901: 899: 898:Palaeotethyan 895: 885: 883: 879: 875: 870: 866: 865: 860: 856: 852: 842: 840: 835: 831: 827: 823: 818: 808: 806: 802: 792: 789: 785: 781: 772: 768: 764: 760: 756: 752: 747: 743: 738: 734: 731: 727: 723: 703: 701: 696: 692: 682: 680: 675: 671: 670: 665: 660: 656: 654: 650: 646: 642: 638: 634: 630: 625: 622: 617: 614: 610: 605: 604:fossil record 600: 596: 592: 587: 585: 581: 580:Verbeekinidae 577: 573: 569: 564: 561: 557: 553: 538: 536: 532: 528: 521: 516: 510: 508: 506: 501: 497: 495: 490: 489: 484: 483:Theriodontian 480: 479:Dinocephalian 476: 475: 470: 469: 468:Titanophoneus 458: 456: 452: 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