Gradual Collapse of Global Marine Ecosystem in the Late Permian and Its Link to the Anoxia
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摘要: 一般认为二叠纪末生物大灭绝持续的时间为3~6万年.然而，越来越多的研究显示在生物灭绝高峰期到来之前存在着环境危机预警信号，但相关研究仍然较少.本文聚焦于大灭绝全过程，包括灭绝高峰期到来之前、灭绝高峰期以及大灭绝之后残存期生物与环境的变化，揭示海洋生态系统坍塌的过程.通过对全球30个海相剖面的化石和古环境记录综合研究，结果表明：（1）深水环境（包括远洋环境、深水陆架、深水盆地和台地边缘斜坡）生态系统衰退发生较早，浅水环境（包括浅水碳酸盐台地、礁和浅水陆架）生态系统衰退发生较晚；（2）浮游生态系统的衰退早于底栖生态系统的衰退.全球海洋生态系统衰退的这种时空差异与最小含氧带（OMZ）的形成及扩展，并导致缺氧有关.Abstract: The duration for the End-Permian mass extinction has been estimated as about 30 to 60 kyr. However, an ever-expanding body of papers has revealed that the evolution of Late Permian ecosystems possibly involved some yet under-studied 'early warning signals' prior to the End-Permian mass extinction. The study on the pre-extinction 'warning signals' is still limited. In order to understand the process of marine ecosystem collapse specifically the 'early warning signals' pointing to the approaching of a global ecosystem regime shift (tipping point), 30 marine Permian-Triassic Boundary sections from different palaeogeographic settings were selected globally to investigate the spatiotemporal biodiversity changes of different taxa and the spatiotemporal redox conditions. The results reveal that: (1) The marine ecosystem collapsed first in deep waters and then in shallow waters (first in offshore pelagic settings, then in moderately deep waters and deep-water basins and shelves and finally in shallow water environments); (2) in the same environments (in deep or moderately deep waters), a similar differential temporal pattern is also apparent in that the planktonic ecosystems were devastated earlier than benthic ecosystems. To account for the spatiotemporal and ecological (taxonomic) differences in extinction timing, we propose that the formation and expansion of an OMZ (oxygen minimum zone) and the related anoxia (or oxygen depletion), were most likely responsible for the differential temporal patterns of marine ecosystem collapses between deep and shallow waters, and between planktonic and benthic communities during the Late Permian.
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Key words:
- collapse of marine ecosystem /
- oxygen minimum zone /
- anoxia /
- Late Permian /
- stratigraphy /
- environmental impact
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图 1 全球晚二叠世古地理与研究剖面的分布
全球古地理图根据
http://www.scotese.com ；华南古地理图根据He et al.（2023）；黑五角星标注为代表性剖面；黄五角星标注为参考剖面.a. 华南以外的研究剖面及其古地理位置：①挪威Spitsbergen；②意大利Western Dolomites；③巴基斯坦Salt Range；④中国西藏色龙；⑤克什米尔地区Guryul Ravine；⑥中国西藏土隆；⑦阿曼；⑧西伯利亚Kuznetsk Basin；⑨加拿大北极West Blind Fiord；⑩澳大利亚悉尼盆地；11新西兰Island Bay of Waiheke Island；12日本.b. 华南研究剖面及其古地理分布：1. 浅海碳酸盐台地；2. 陆相；3. 近岸浅海碎屑岩陆架；4. 孤立碳酸盐台地；5. 深水硅质岩盆地（半深海及以下）；6. 浅海硅质岩局限盆地；7. 古陆；8. 水下隆起；9. 古特提斯洋俯冲；10. 裂谷盆地；11. 礁和微生物岩相；12. 古地理方向.剖面代号含义：SS. Shangsi，四川广元上寺；YDZ. Yudongzi，四川江油鱼洞子；LLD. Laolongdong，重庆老龙洞；LFY. Liangfengya，重庆凉风垭；RCP. Rencunping，湖南桑植仁村坪；DXK. Daxiakou，湖北兴山大峡口；CL. Cili，湖南慈利；CB. Chibi，湖北赤壁；MJS. Majiashan，安徽巢湖马家山；MS. Meishan，浙江长兴煤山；HZS. Huangzhishan，浙江湖州黄芝山；YG. Yangou，江西沿沟；CH. Chahe，贵州威宁岔河；ZZ. Zhongzhai，贵州六枝中寨；KJ. Kejiao，贵州惠水克脚；DJ. Dajiang，贵州罗甸打讲；ZD. Zuodeng，广西田东作登；DP. Dongpan，广西扶绥东攀Fig. 1. Late Permian global palaeogeography and localities of studied sections
图 3 全球不同古地理环境中代表性剖面生物灭绝的层位及灭绝率（基于10个代表性剖面）
剖面原始资料来源: 凉风垭.腕足类参考沈树忠和何锡麟(1991)，有孔虫参考Liu et al.2020).慈利.有孔虫和蜓参考杨利蓉等(2013).黄芝山.腕足类参考Heet al.(2015), 有孔虫参考Chen etal. (2009).煤山.腕足类参考He et al.(2015, 2023): 有孔虫参考Song et al.(2009): 放射虫参考Heet al.(2005).上寺.有孔虫参考Liu et al.(2020): 放射虫在聂小妹等(2012)基础上补充(根据课题组最近的工作).仁村坪.腕足类参考He et al.(2015, 2019): 放射虫参考Xiao et al.(2024).克脚.腕足类和放射虫根据Wang el al.(2023).东攀.腕足类根据He et al.(2019): 放射虫参考Feng et al.，2007), He et al.(2024).日本Nf1212.放射虫参考Sano et al.(2010).北极加拿大.硅质海绵参考Algeo el al.(2012).本图各剖面生物灭绝层位及灭绝率分析过程见He el al.(2025)的附件材料.岩性符号含义: (1).泥岩: (2).钙质泥岩: (3)碳质泥岩: (4).钙质碳质泥岩: (5).硅质泥岩: (6).碳质硅质泥岩: (7).碳质锰质泥岩: (8).钙质硅质泥岩: (9).含凝灰质泥岩: 10).含凝灰质钙质泥岩: (1).含凝灰质硅质泥岩; (12).含凝灰质粉砂质泥岩; (13).碳质页岩; (14).页岩; (15).粉砂岩; (16).粉砂质泥岩; (17).泥质粉砂岩; (18).含粉砂质钙质泥岩; (19).燧石条带; (20).燧石; （(21).碳质硅质岩; (22).微生物岩；(23).灰岩: (24).泥质灰岩: (25).含燧石团块泥质灰岩: (26).含燧石团块灰岩: (27).生物碎屑灰岩；(28).含燧石团块生物碎屑灰岩; (29).鲕状灰岩; (30).蠕虫状灰岩; (31).硅质灰岩; (32).碳质灰岩; (33).藻灰岩; (34).硅质锰质灰岩; (35).含硅的泥质灰岩; (36).含凝灰质泥质灰岩; (37).黄铁矿; (38).火山粘土; (39).铁质泥岩
Fig. 3. Columnar Permian-Triassic sections showing extinction horizons/intervals and/or extinction rates of different taxa in varied water depths from pelagic setting, deep-water basin, moderately deep-water slope to shallow-water facies (based on 10 representative sections)
图 4 全球不同古地理环境中浮游生物‒底栖生物灭绝时间的对比
华南研究剖面的地层对比根据图 2；其他地区的地层对比根据He et al. (2025)的附件材料.生物灭绝时间的数据来源：中国华南凉风垭、慈利、黄芝山、煤山、上寺、仁村坪、克脚以及东攀剖面，日本和加拿大北极剖面根据图 3；中国中寨根据Zhang et al.（2015）、He et al.（2023）；阿曼根据Clarkson et al.（2016）；挪威根据Nabbefeld et al.（2010）、Zuchuat et al.（2020）、Rodríguez-Tovar et al.（2021）；意大利根据Posenato（2009, 2019）；巴基斯坦、中国西藏色龙根据Shen et al.（2006）；克什米尔地区、中国西藏土隆根据Shen et al.（2006）
Fig. 4. Comparison of extinction timing between planktons and benthos across shallow to deep waters
图 5 全球不同古水深环境浮游生物‒底栖生物灭绝时间的对比和总结
部分图例解释参考图 4]
Fig. 5. Summary and comparison of extinction timings across shallow water, moderately deep water to deep water settings
图 6 最小含氧带OMZ的形成与扩展对生态系统坍塌的影响
a. Stage Ⅰ，OMZ形成导致深水浮游生物灭绝；b. Stage Ⅱ，OMZ向下扩展导致深水底栖生态系统坍塌；c. Stages Ⅲ和Ⅳ，OMZ继续向下和向上扩展导致深水和浅水生态系统倒塌
Fig. 6. Formation/expansion of the OMZ and its influence on the marine ecosystem collapse
表 1 研究剖面古水深相关信息及文献来源
Table 1. Summary of studied sections, including details of palaeogeographic setting (palaeo-water depths) and source of published information for each chosen section
序号 剖面名称 地区 代表性剖面重点研究内容 古纬度/
古经度二叠系‒三叠系界线附近地层古水深证据 古水深 作为研究剖面的原因 参考文献 1 中寨 中国
华南14°20′45″S
105°23′47″E粉砂质泥岩、泥岩；
含大量腕足类(近岸)浅水 < 50 m 腕足类研究详细(但界线附近有疑问); 有关键层位的地层划分与对比 Shen et al., 2011; Zhang et al., 2015; He et al., 2023 2* 慈利 生物灭绝, 氧化‒还原环境 10°22′07″S
109°54′13″E生物碎屑灰岩;
微生物岩(礁)化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比王钦贤等, 2009;
杨利蓉等, 2013;
廖卫, 20203* 鱼洞子 氧化‒还原环境 8°18′S
104°32′E生物碎屑灰岩;
微生物岩(礁)氧化‒还原环境（Ce异常）研究详细; 有关键层位的地层划分与对比 方宇恒, 2021 4* 老龙洞 氧化‒还原环境 8°47′56″S
114°39′59″E生物碎屑灰岩;
微生物岩(礁)化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Huang et al., 2022 5* 打讲 氧化‒还原环境 14°45′54″S
106°44′15″E生物碎屑灰岩;
微生物岩(礁)化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Song et al., 2009; Jiang et al., 2014; Liao et al., 2017 6* 作登 氧化‒还原环境 16°49′03″S
107°21′39″E生物碎屑灰岩;
微生物岩(礁)化石和氧化‒还原环境（Ce异常）研究详细;
有关键层位的地层划分与对比Wan et al., 2019; He et al., 2023;
廖卫, 20207* 黄芝山 生物灭绝, 氧化‒还原环境 7°04′36″S
117°17′15″E生物碎屑灰岩、泥质灰岩(台地) 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比陈军等，2008；Chen et al., 2009; He et al., 2015 8* 沿沟 氧化‒还原环境 9°20′55″S
115°31′43″E生物碎屑灰岩(台地) 化石和氧化‒还原环境（Ce异常）研究详细;
有关键层位的地层划分与对比田力等, 2014;
Li et al., 2017;
Qiu et al., 20199* 凉风垭 生物灭绝, 氧化‒还原环境 10°53′39″S
105°57′44″E生物碎屑灰岩、泥质灰岩(台地) 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比沈树忠和何锡麟, 1991; 李国山, 2016; 袁东勋和沈树忠, 2011;
Liu et al., 202010* 煤山 生物灭绝, 氧化‒还原环境 6°59′29″S
117°00′30″E生物碎屑灰岩, 硅质岩; 丘状交错层理; 球形放射虫(斜坡) 中等深水50~200 m 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Song et al., 2009; Shen et al., 2011; Yuan et al., 2014; Li et al., 2016; He et al., 2023 11* 上寺 生物灭绝, 氧化‒还原环境 8°12′15″S
104°46′55″E生物碎屑灰岩, 硅质岩; 鲍玛序列; 深水放射虫（La. and Al.）(斜坡) 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Yuan et al., 2019; Zhang et al., 2020; Liu et al., 2020; He et al., 2025 12* 赤壁 氧化‒还原环境 9°34′28″S
112°25′13″E生物碎屑灰岩, 硅质岩; 钙质角砾; 球形放射虫(斜坡) 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比李国山, 2016; Yang et al., 2022; He et al., 2025 13* 大峡口 氧化‒还原环境 8°41′53″S
109°30′45″E碳质灰岩, 碳质泥岩; 小有孔虫(盆地) 深水 > 200 m 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Zhao et al., 2013; Shen et al., 2016; Zhang and Gu, 2015 14* 马家山 氧化‒还原环境 4°39′42″S
114°10′16″E硅质泥岩, 碳质泥岩; 深水放射虫（La.）(盆地) 化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比方宇恒, 2021;
He et al., 202315* 克脚 生物灭绝, 氧化‒还原环境 14°31′18″S
106°35′26″E硅质泥岩, 碳质泥岩; 深水放射虫
（La. and Al.）(盆地)化石和氧化‒还原环境（草莓状黄铁矿）研究详细;
有关键层位的地层划分与对比Wang et al., 2023 16* 仁村坪 生物灭绝 10°19′32″S
109°10′60″E硅质泥岩, 硅质灰岩, 钙质泥岩;
深水放射虫
（La. and Al.）(盆地)化石研究详细;
有关键层位的地层划分与对比He et al., 2015, 2019, 2023;
Xiao et al., 202417* 东攀 生物灭绝, 氧化‒还原环境 17°53′28″S
108°12′41″E硅质泥岩, 燧石;
深水放射虫
（La. and Al.）(盆地)化石和氧化‒还原环境（Ce异常）研究详细;
有关键层位的地层划分与对比Feng et al., 2007; Shen et al., 2012; Baresel et al., 2017; He et al., 2023 18 Western Dolomites (②) 意大利 12°45′N
36°15′E生物碎屑灰岩、鲕状灰岩(浅水陆架) 浅水 < 50 m 化石和氧化‒还原环境初步研究；
有地层时代对比Posenato, 2009, 2019 19* Nf1212 日本⑫ 生物灭绝 2°06′N
131°56′E燧石; 深水放射虫（La. and Al.）(远洋) 深水 > 500 m 放射虫研究详细;
有关键层位的地层划分与对比Sano et al., 2010, 2012 20 Sasayama- Kinkazan-Tenjinmaru
复合剖面1°48′N
130°48′E燧石; 深水放射虫（La. and Al.）(远洋) 氧化‒还原环境和地层时代有初步研究 Kajiwara et al., 1993a, 1993b; Kato et al., 2002 21* Akkamori 氧化‒还原环境 4°23′N
137°04′E燧石; 深水放射虫（La. and Al.）(远洋) 氧化‒还原环境和关键层位的地层时代研究较详细 Takahashi et al., 2021 22 Salt Range (③) 巴基斯坦 35°S
55°E灰岩, 粉砂质泥岩;
大个体腕足类(浅水陆架)浅水 < 50 m 腕足类详细研究;
有关键层位的地层时代对比Shen et al., 2006 23 Selong (④) 中国西藏 47°01′S
53°38′E灰岩, 白云岩; 大个体腕足类(浅水陆架) 腕足类详细研究;
有关键层位的地层时代对比Shen et al., 2006 24 Site 1 (⑦1) 阿曼 53°06′58″N
22°17′37″E灰岩, 白云岩;
珊瑚, 有孔虫(台地)有初步遗迹化石和地层时代研究；
氧化‒还原环境研究详细Clarkson et al., 2016 25 Guryul Ravine(⑤) 克什米尔地区 36°S
56°E灰岩, 页岩;
腕足类(深水陆架)深水 > 200 m 腕足类和氧化‒还原环境（Ce异常）研究详细; 有地层时代对比 Shen et al., 2006; Algeo et al., 2007 26 Tulong (⑥) 中国西藏 47°33′S
53°29′E泥质灰岩, 页岩, 砂岩; 腕足类(深水陆架) 腕足类详细研究; 地层时代有初步研究 Shen et al., 2006 27 Site 6 (⑦2) 阿曼 52°52′34″N
21°47′37″E硅质泥岩, 页岩;
水平层理(盆地)氧化‒还原环境研究
详细；地层时代有初步研究Clarkson et al., 2016 28 Spitsbergen (①) 挪威 54°44′06″N
22°58′06″E海绿石砂岩, 泥岩(浅水陆架) 浅水 < 50 m 氧化‒还原环境研究
详细；有关键层位的地层时代对比Nabbefeld et al., 2010; Zuchuat et al., 2020; Rodríguez-Tovar et al., 2021 29* Island Bay of Waiheke Island (⑪) 新西兰 氧化‒还原环境 86°21′26″S
163°09′34″E燧石, 硅质泥岩; 放射虫; 水平层理(远洋) 深水 > 500 m 氧化‒还原环境（草莓状黄铁矿）研究详细；有关键层位的地层时代研究 Grasby et al., 2021 30* West Blind Fiord (⑨WBF) 加拿大北极 生物灭绝 63°N
42°E燧石, 硅质泥岩; 硅质海绵(远洋) 化石和关键层位的地层时代研究较详细 Algeo et al., 2012 注：*代表性剖面（之所以被选为代表性剖面的理由见本表第4栏“代表性剖面重点研究内容”，即在生物灭绝或者氧化‒还原环境研究方面基础较好）; ①为剖面编号，与图 1a一致; La. Latentifistularia十字多囊虫目, Al. Albaillellaria阿尔拜虫目（这两个目为深水分子，一般阿尔拜虫生活于200 m以下的深水环境，十字多囊虫生活于100~150 m水深以下，见 He et al., 2024 ）.
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