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维基百科，自由的百科全书
寒武紀奧陶紀志留紀泥盆紀石炭紀二疊紀三疊紀侏羅紀白堊紀古近紀新近紀
單位：百萬年
寒武紀奧陶紀志留紀泥盆紀石炭紀二疊紀三疊紀侏羅紀白堊紀古近紀新近紀
本圖僅呈現地質年代各期的海洋生物滅絕比例，包含規模最大的五次滅絕事件。藍色部份代表大致的滅絕屬比例，以該時期與下一時期的化石紀錄比較計算。

瓜达洛普世末期灭绝事件（英語：end-Guadalupian extinction event），也称卡匹敦期灭绝事件（Capitanian extinction event）[1]、瓜达洛普世—乐平世边界大灭绝（Guadalupian-Lopingian boundary mass extinction）[2]、前乐平世危机（pre-Lopingian crisis）[3]或二叠纪中期灭绝事件（Middle Permian extinction），是古生代二叠纪后期瓜达洛普世和乐平世之交的一次生物集群灭绝事件，比著名的二叠纪末大灭绝早约720万。此次灭绝事件的背景是二叠纪中后期从2.62亿年前就开始出现物种多样性下降和灭绝率上升——即所谓的“晚瓜达洛普世危机”（Late Guadalupian crisis）[4]，其中地层学研究发现灭绝高峰出现在约2.59亿年前的卡匹敦期后半叶[5]。

瓜达洛普世末期灭绝事件历史上曾被视为二叠纪末大灭绝的一部分，直到近年才被视作是独立的灭绝事件[6]，从物种灭绝数量方面看其实是整个显生宙第三大灭绝事件（仅次于二叠纪末大灭绝和奥陶纪末大灭绝）[7]和第五严重的生态破坏事件[8]。因为一些分析发现此次事件只影响北半球低纬度的类群，所以许多古生物学家质疑其是否可以被视作全球性事件[9]。

程度

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全球的物种多样性在奥尔森灭绝事件后逐渐恢复，很可能是因为幸存的先锋物种得益于新空出的生态位并演替了之前优势的灭绝物种（如各种盘龙目）。瓜达洛普世末期的灭绝事件严重减少了同资源种团的数量（共有八个种团丧失），其在各群落内的多样性损失成都超过了有“大死亡”之称的二叠纪末大灭绝[10]。虽然动物群随后就开始恢复[11][12]，生态系统营养级结构和种团复杂性得到了一定重建和补充[10]，物种多样性和差异性仍整体持续下跌并持续到了二叠纪—三叠纪边界[13]。.

海洋生态

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瓜达洛普世末期灭绝事件对海洋生态系统造成的冲击在古生物学界仍有争议。瓜达洛普世末期灭绝事件中海洋无脊椎动物的损失程度可与白垩纪末大灭绝媲美[14]，早期估测显示有35％至47％的属损失[15][16]，而2016年发表的一份估测认为考虑背景灭绝率、西格诺尔-利普斯效应和类群间灭绝的不对称性后海洋生物属的损失应该是33～35％[17]。一些研究认为从海洋无脊椎动物的角度，瓜达洛普世末期灭绝事件是有史以来第三或第四大灭绝事件；另一项研究发现从属的损失率来看，瓜达洛普世末期灭绝事件只是有史以来的第九大，但从生态冲击程度上则是第五大[18]。

陆地生态

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瓜达洛普世末期灭绝事件对陆地生态系统影响的研究相对缺乏。关于南非卡鲁盆地脊椎动物化石的研究认为有74～80％的四足动物属因此损失[19]，包括当时处于优势的恐头兽亚目完全灭绝[20]。而对中国华北石盒子组的陆地植物研究发现有56％的植物种类灭绝[21]，而华南有24％的植物灭绝[22]。

时间段

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虽然学界确定瓜达洛普世末期灭绝事件发生在奥尔森灭绝事件和二叠纪末大灭绝之间，[10]，确切的时间段仍有争议，部分因为卡匹敦期与吴家坪期间边界本身仍未确定——目前的估计是约2.591年前[19][20][23]，但国际地层委员会可以随时对二叠纪的地层学定义进行调整。除此之外，当时在现今中国发生的集群灭绝是否和在北欧斯匹次卑尔根岛的灭绝发生在同一时段仍有争议[24]。有一项研究认为瓜达洛普世末期灭绝事件不是一个单一事件，而是始于沃德期末期的持续性多样性降低[25]。另一项审查斯瓦尔巴化石岩相的研究并未发现突发性大规模灭绝的证据，因而认为当地生物相在卡匹敦期的变化是因为许多类群开始向南迁徙穿过史前的蔡希斯坦海[26]。匈牙利和伊兹拉岛的碳酸盐岩地台没有发现卡匹敦期末的灭绝事件，而是发现了卡匹敦期中期有灭绝事件[27]。

当时处于热带的峨眉山暗色岩火山（现位于四川峨眉山茅口组的碳酸盐岩地台中）是整个灭绝事件的起因，同时也保留了最多的化石和地质证据[22]。峨眉山准岩浆型火山喷发造成了䗴类有孔虫和钙化藻类的灭绝[28]。

因为尚且没有直接测量海洋生态群的放射测年法，近年大部分研究都不愿设定瓜达洛普世末期灭绝事件的具体数字年份，但根据二叠纪的时间段外推可以估计出约在2.6～2.62亿年前[19][29]——这与陆地生态系统的放射年期相符，特别是植物相损失与海洋大灭绝大致在同一时期或稍晚一些[22]。

海洋

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中国西南地区的䗴类有孔虫灭绝期初被定为在瓜达洛普世末期，但2009年和2010年发表的研究将灭绝定为卡匹敦期中期[30]，比如腕足动物和珊瑚的损失[31]。菊石遭受的灭绝则发生在吴家坪期早期[31]。

陆地

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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 Titanophoneus Superzone and the Scutosaurus Superzone[32] and later the Dinocephalian Superassemblage and the Theriodontian Superassemblage, respectively. In South Africa, this corresponded to the boundary between the variously named Pareiasaurus, Dinocephalian or Tapinocephalus Assemblage Zone and the overlying assemblages.[33][34][35][36] In both Russia and South Africa, this transition was associated with the extinction of the previously dominant group of therapsid amniotes, the dinocephalians, which led to its later designation as the dinocephalian extinction.[37] Post-extinction origination rates remained low through the Pristerognathus Assemblage Zone for at least 1 million years, which suggests that there was a delayed recovery of Karoo Basin ecosystems.[38]

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.[14] Though it was subsequently suggested that because the Russian Ischeevo fauna, which was considered the youngest dinocephalian fauna in that region, was constrained to below the Illawarra magnetic reversal and therefore had to have occurred in the Wordian stage, well before the end of the Guadalupian,[37] this constraint applied to the type locality only. The recognition of a younger dinocephalian fauna in Russia (the Sundyr Tetrapod Assemblage)[39] and the retrieval of biostratigraphically well-constrained radiometric ages via uranium–lead dating of a tuff from the Tapinocephalus Assemblage Zone of the Karoo Basin[19][40] demonstrated that the dinocephalian extinction did occur in the late Capitanian, around 260 million years ago.

对生命的影响

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海洋生物

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In the oceans, the Capitanian extinction event led to high extinction rates among ammonoids, corals and calcareous algal reef-building organisms, foraminifera, bryozoans, and brachiopods. It was more severe in restricted marine basins than in the open oceans.[41] It appears to have been particularly selective against shallow-water taxa that relied on photosynthesis or a photosymbiotic relationship;[42] many species with poorly buffered respiratory physiologies also became extinct.[43][44] The extinction event led to a collapse of the reef carbonate factory in the shallow seas surrounding South China.[45][46]

The ammonoids, which had been in a long-term decline for a 30 million year period since the Roadian, suffered a selective extinction pulse at the end of the Capitanian.[13] 75.6% of coral families, 77.8% of coral genera and 82.2% of coral species that were in Permian China were lost during the Capitanian mass extinction.[47] The Verbeekinidae, a family of large fusuline foraminifera, went extinct.[48]

87% of brachiopod species found at the Kapp Starostin Formation on Spitsbergen disappeared over a period of tens of thousands of years; though new brachiopod and bivalve species emerged after the extinction, the dominant position of the brachiopods was taken over by the bivalves.[49] Approximately 70% of other species found at the Kapp Starostin Formation also vanished.[50] The fossil record 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.[29] Whereas rhynchonelliform 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, molluscs made up 61.2% of the individuals found in similar environments after the extinction.[51] 87% of brachiopod species and 82% of fusulinacean foraminifer species in South China were lost.[29] Although severe for brachiopods, the Capitanian extinction's impact on their diversity was nowhere near as strong as that of the later end-Permian extinction.[52]

Biomarker evidence indicates red algae 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.[53]

Most of the marine victims of the extinction were either endemic species of epicontinental seas around Pangaea that died when the seas closed, or were dominant species of the Paleotethys Ocean.[54] Evidence from marine deposits in Japan and Primorye suggests that mid-latitude marine life became affected earlier by the extinction event than marine organisms of the tropics.[55]

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,[9] others suggest that there was little latitudinal variation in extinction patterns.[56] 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.[57]

This mass extinction marked the beginning of the transition between the Palaeozoic and Modern evolutionary faunas.[1] 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.[58] After the Capitanian mass extinction, disaster taxa such as Earlandia and Diplosphaerina became abundant in what is now South China.[5] 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 Tubiphytes-dominated reefs, which in turn was followed by a return of metazoan, sponge-dominated reefs.[59] Overall, reef recovery took approximately 2.5 million years.[2]

陆地生物

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Among terrestrial vertebrates, the main victims were dinocephalian therapsids, which were one of the most common elements of tetrapod fauna of the Guadalupian; only one dinocephalian genus survived the Capitanian extinction event.[14] The diversity of the anomodonts that lived during the late Guadalupian was cut in half by the Capitanian mass extinction.[60] Terrestrial survivors of the Capitanian extinction event were generally 20公斤（44磅） to 50公斤（110磅） and commonly found in burrows.[14]

起因

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峨眉山暗色岩

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火山喷发

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It is believed that the extinction, which coincided with the beginning of a major negative δ13C excursion signifying a severe disturbance of the carbon cycle,[61][22] was triggered by eruptions of the Emeishan Traps large igneous province,[62][63][64] basalt piles from which currently cover an area of 250,000 to 500,000 km2, although the original volume of the basalts may have been anywhere from 500,000 km3 to over 1,000,000 km3.[44] The age of the extinction event and the deposition of the Emeishan basalts are in good alignment.[65][66] Reefs and other 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.[67] These eruptions would have released high doses of toxic mercury;[68][69] increased mercury concentrations are coincident with the negative carbon isotope excursion, indicating a common volcanic cause.[70] Coronene 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 anomalies coeval with mercury and coronene anomalies.[71] A large amount of carbon dioxide and sulphur dioxide is believed to have been discharged into the stratosphere of the Northern and Southern Hemispheres due to the equatorial location of the Emeishan Traps, leading to sudden global cooling and long-term global warming.[28] 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 the most precipitous in the entire geological history of the Earth.[72] 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.[73] Significant quantities of methane released by dikes and sills intruding into coal-rich deposits has been implicated as an additional driver of warming,[74] 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.[75][76][77] The emissions of the 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.[78]

缺氧硫化

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Global warming resulting from the large igneous province's activity has been implicated as a cause of marine anoxia.[79][80] Two anoxic events, the middle Capitanian OAE-C1 and the end-Capitanian OAE-C2, occurred thanks to Emeishan volcanic activity.[81] Volcanic greenhouse gas release and global warming increased continental weathering and mineral erosion, which in turn has been propounded as a factor enhancing oceanic euxinia.[82] Euxinia may have been exacerbated even further by the increasing sluggishness of ocean circulation resulting from volcanically driven warming.[83] 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.[67] 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.[84]

高二氧化碳和酸化

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Because the ocean acts as a carbon sink 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.[85] The dissolution of volcanically emitted carbon dioxide in the oceans triggered ocean acidification,[29][24][49] which probably contributed to the demise of various calcareous marine organisms, particularly giant alatoconchid bivalves.[86] By virtue of the greater solubility of carbon dioxide in colder waters, ocean acidification was especially lethal in high latitude waters.[80] Furthermore, acid rain would have arisen as yet another biocidal consequence of the intense sulphur emissions produced by Emeishan Traps volcanism.[28] This resulted in soil acidification and a decline of terrestrial infaunal invertebrates.[87] 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 pH.[88]

对火山起因说的批判

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Not all studies, however, have supported the volcanic warming hypothesis; analysis of δ13C and δ18O values from the tooth apatite of Diictodon feliceps specimens from the Karoo Supergroup 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.[89] Analysis of vertebrate extinction rates in the Karoo Basin, specifically the upper Abrahamskraal Formation and lower Teekloof Formation, 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.[90] 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.[9]

海平面下降

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The Capitanian mass extinction has been attributed to sea level fall,[91] with the widespread demise of reefs in particular being linked to this marine regression.[85] The Guadalupian-Lopingian boundary coincided with one of the most prominent first-order marine regressions of the Phanerozoic.[5] 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.[85] Additionally, a tremendous unconformity is associated with the Guadalupian-Lopingian boundary in many strata across the world.[92] The closure of the Sino-Mongolian Seaway at the end of the Capitanian has been invoked as a potential driver of Palaeotethyan biodiversity loss.[93]

其它假说

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因为盘古大陆整合造成的全球干旱、板块构造和种间竞争也可能部分造成了灭绝事件发生[5][21][47][89]。其它可能造成灭绝（特别是生物礁）的因素还包括海水盐度的波动和微大陆碰撞[85]。

另见

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参考

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