Stepwise Oxygenation of Early Cambrian Ocean Drove Early Metazoan Diversification
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摘要: 伊迪卡拉-寒武纪过渡期是地球历史上海洋环境与生命演化变革的关键时期之一.有研究认为海洋氧化可能是驱动早寒武世动物快速多样化的主要原因，但有关海洋氧化及硫化水体演变过程和发生时间的认识仍有分歧.运用ICP-MS、FESEM、EDS和XRD技术对位于古陆架边缘-斜坡背景的湘西四都坪、黔东松江大桥等剖面牛蹄塘组黑色页岩进行氧化还原敏感元素、黄铁矿形态学、总有机碳及N、P等营养元素丰度变化进行高分辨研究.结果显示，研究区牛蹄塘沉积期底层海水条件经历了复杂的发展过程，表现为3个铁化与3个硫化时段的动态交互，在第3阶晚期转为亚氧化-氧化条件.不同相区的对比表明，海水氧化随时间由浅水向深水区扩展：台地相区发生在第2阶晚期，陆架边缘出现在第3阶晚期，而深水盆地以持续铁化为主，晚期出现硫化，表明海洋氧化界面逐步加深下移.沉积物Mo/TOC，U/TOC值自下而上持续增加，与Cr、Mn、P、N丰度变化趋势一致，表明海水氧化程度逐步增强.主要化石类群的时空分布与海水氧化过程对应良好，表明海洋阶段性氧化与硫化水体消失对后生动物多样化进程有明显的控制作用.Abstract: Ediacaran-Cambrian transition is a critical period in Earth history, during which both marine environment and life have experienced drastic changes. It is arguably suggested that pervasive oxygenation and associated geochemical changes in the ocean have potentially driven the rapid diversification of Early Cambrian metazoans, however, the timing and process of ocean oxygenation have not been well constrained. In this paper, an integrated study was conducted on the Lower Cambrian Niutitang (NTT) Fm. in Siduping area, West Hunan and Songjiang Bridge area, East Guizhou, which were paleogeographically situated at the shelf margin-upper slope setting of the Yangtze block. Using ICP-MS, FESEM, EDS and XRD techniques, the abundances and patterns of RSE (redox sensitive elements), pyrite morphology, TOC (total organic carbon), as well as N, P nutrients were investigated systematically. The results show that during NTT deposition, bottom seawaters in the study area experienced a complicated evolution, with a dynamic alternation of 3 ferruginous and 3 euxinic intervals, and suboxic-oxic conditions occurring only at the latest Cambrian Stage 3. Comparison with relevant data published previously from each facies belts within the Yangtze block shows that the seawater oxygenation was a progressive process expanding from shallow water area to deep-water area in time. Platform facies area oxygenated at the late Stage 2, shelf margin area in the late Stage 3, and the deep-water basin kept ferruginous until the latest Stage 3, when it became euxinic. Sediment Mo/TOC, U/TOC ratios increased from bottom to top consecutively in the section, coupled with elevated Cr, Mn, N and P secular trends, likely indicating an overall enhancement of seawater oxygenation. The spatial-temporal distribution and increased paleoecologic complexity of major fossil groups in the Yangtze block over time show a well coincidence with the seawater oxygenation process, likely implying that increased ocean oxygenation and declined euxinic seawaters have facilitated the rapid diversification of Early Cambrian metazoans.
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图 1 扬子地区晚伊迪卡拉纪-早寒武世沉积古地理背景及研究区地层序列
a.沉积古地理简图，据Jiang et al.(2007)、Steiner et al.(2007)、Xu et al.(2012)、Och et al.(2013)等修改补充；b.NW-SE向古地势示意图；c.研究区晚伊迪卡拉-早寒武世地层序列.图a中红色五星表示前人研究的剖面，蓝色三角表示本文研究的剖面
Fig. 1. Simplified paleogeography of South China during Ediacaran-Cambrian transition and typical stratigraphic succession in the study area
图 2 扬子地台不同相区晚伊迪卡拉纪至寒武纪早期地层划分与对比
据Steiner et al.(2007)、Zhao et al.(2012)、Jin et al.(2016)和Xiang et al.(2017)等资料综合修改.SSF.小壳类化石组合(small shelly fossils)；FAD.化石首现点(first appearance datum)；灰色方块表示黑色页岩为主的地层单位；竖线条表示地层缺失
Fig. 2. Stratigraphic subdivisions and correlation of the Late Ediacaran-Early Cambrian transition on the Yangtze platform
图 3 研究区下寒武统牛蹄塘组(NTT)沉积类型与微相特征
a.松桃松江大桥牛蹄塘组下部黑色页岩与留茶坡组分界；b.江口平引村牛蹄塘组底部黑色泥岩中的磷结核；c.湘西四都坪牛蹄塘组下部硅化-硅质页岩；d.松江大桥牛蹄塘组一段下部黑色页岩；e.松江大桥牛蹄塘组下部黑色泥页岩中的黄铁矿条带；f.四都坪牛蹄塘组下部硅质页岩中的硅化球粒和石英粉砂；g.四都坪牛蹄塘组一段上部黄铁矿条带；h.四都坪牛蹄塘组一段自形黄铁矿；i.四都坪牛蹄塘组二段上部草莓状黄铁矿；j.四都坪牛蹄塘组二段上部草莓状黄铁矿.a~e.野外照片；f~i.显微照片；j.SEM照片.照片中，Euhd.自形黄铁矿；Fr.草莓状黄铁矿；Qz.硅化微球粒
Fig. 3. Sedimentary types and micro-facies features of the Niutitang (NTT) Fm. in study area
图 4 四都坪剖面牛蹄塘组草莓状黄铁矿分布及其特征
a.草莓状黄铁矿地层分布；b.草莓状黄铁矿平均粒径与标准偏差散点分布图；c.草莓状黄铁矿平均粒径与偏斜散点分布图.图a数据点中的红线表示平均粒径值，方框表示95%置信区间，黑色横线表示粒径分布范围；LCP.留茶坡组，QXD.清虚洞组
Fig. 4. Distribution and characteristics of framboidal pyrites in the NTT Fm., Siduping, West Hunan
图 5 四都坪牛蹄塘组RSE、TOC、N、P丰度及其相关比值的地层分布特征
LCP.留茶坡组；QXD.清虚洞组
Fig. 5. Stratigraphic distribution of RSE, TOC, N, P abundances and their related ratios in the NTT Fm., Siduping, West Hunan
图 6 研究区牛蹄塘组Mo-TOC浓度与现代硫化盆地局限度比较(a)及MoEF/UEF相关性限定的水体化学条件(b)
Fig. 6. Comparison of the Mo vs. TOC concentrations from the NTT Fm. with those in some recent anoxic-euxinic basins (a), and the possible seawater redox conditions constrained by MoEF vs. UEF plot in the NTT Fm. (b)
图 7 寒武纪大爆发过程中主要后生动物类群出现与海水氧化的时间对比
生物资料据Shu(2008)、朱茂炎(2010)、Erwin(2011)和Shu et al.(2014)等资料综合；海水条件变化据本文研究
Fig. 7. Time correlation of major metazoan group occurrence with ocean oxygenation during the Cambrian explosion
表 1 四都坪牛蹄塘组(NTT)黑色页岩主量元素分析数据
Table 1. Major element data of the black shale from the NTT Fm., Siduping, West Hunan
单元平均值 SiO2
(%)TiO2
(%)Al2O3
(%)Fe2O3T
(%)MgO
(%)CaO
(%)Na2O
(%)K2O
(%)MnO
(μg/g)P2O5
(μg/g)LOI
(%)总量
(%)第1段(0~13 m；n=14) 下部 73.7 0.4 9.7 0.5 0.9 0.2 0.7 2.9 30.8 165.3 10.4 99.5 上部 77.8 0.4 8.9 0.5 0.7 0.1 0.7 2.6 26.4 148.9 7.8 99.5 全段 76.0 0.4 9.3 0.5 0.8 0.2 0.7 2.7 28.3 155.9 8.9 99.5 第2段(13~55 m；n=69) 下部 75.8 0.4 9.8 1.1 0.9 0.4 1.0 2.4 32.4 318.3 7.5 99.5 中部 74.7 0.5 11.0 1.7 1.2 0.2 1.1 2.6 46.5 391.8 6.5 99.5 上部 72.7 0.6 13.3 1.2 1.3 0.2 1.1 3.4 51.3 288.3 5.8 99.6 全段 73.8 0.5 12.0 1.4 1.2 0.2 1.1 3.0 47.0 325.7 6.3 99.5 第3段(55~74 m；n=28) 下部 69.9 0.7 16.7 1.4 1.2 0.1 0.7 4.6 57.7 338.3 4.2 99.7 上部 71.5 0.6 13.0 2.8 0.2 0.1 0.9 3.3 126.9 625.9 5.4 98.0 顶部 73.1 0.3 6.7 2.8 0.0 0.1 0.7 1.6 76.2 1815.3 12.8 98.3 全段 70.8 0.7 14.5 2.1 0.7 0.1 0.8 3.9 93.0 534.9 5.1 98.8 牛蹄塘组 全组 73.3 0.6 12.3 1.5 1.0 0.2 1.0 3.2 56.2 357.0 6.3 99.4 注：由于测试样品较多，本表仅给出各层段的统计平均值，n表示该段测试样品数；以下各表同. 
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表 2 四都坪牛蹄塘组黑色页岩各层段微量及其他关注元素的分析结果
Table 2. Trace element data and concerned ratios for subdivided intervals of the NTT Fm., Siduping, West Hunan
单元平均值 TOC
(%)TN
(%)TS
(%)Mo
(μg/g)U
(μg/g)V
(μg/g)MoEF UEF VEF Mo/TOC
((μg/g)/%)U/TOC
((μg/g)/%)δEu P
(μg/g)Cr
(μg/g)Mn
(μg/g)Cu
(μg/g)Zn
(μg/g)(N/C)mol FePy/
FeTFe/
Al下部 7.9 0.2 0.3 105 25.1 1327.1 206.4 14.8 18.2 14.1 3.4 1.2 72.2 99.8 23.9 8.6 14.5 0.03 0.72 0.07 上部 7.2 0.2 0.3 57.6 10.5 322.4 126.8 6.1 4.1 8.4 1.5 1.4 65.0 65.1 20.4 7.3 11.7 0.03 0.71 0.07 全段 7.5 0.2 0.3 78.2 16.8 753.0 160.9 9.8 10.1 10.9 2.3 1.4 68.1 80.0 21.9 7.9 12.9 0.03 0.72 0.07 下部 6.4 0.2 0.3 54.7 12.7 186.2 105.0 6.9 1.6 8.6 2.0 1.0 139.0 59.7 25.1 18.1 11.8 0.02 0.41 0.15 中部 5.3 0.2 0.7 43.4 10.3 151.6 74.6 4.8 0.9 8.2 2.0 1.0 171.1 66.6 36.0 23.3 24.6 0.03 0.48 0.20 上部 3.4 0.2 0.4 31.8 8.8 217.1 44.9 3.1 1.2 10.4 2.8 1.1 125.9 75.5 39.7 8.5 17.6 0.06 0.27 0.13 全段 4.4 0.2 0.5 38.8 9.9 191.7 63.1 4.2 1.2 9.5 2.4 1.0 142.2 70.4 36.4 14.6 19.0 0.04 0.36 0.16 下部 1.9 0.2 0.1 30.5 9.6 325.6 33.9 2.5 1.8 22.0 6.9 1.1 147.7 89.4 44.7 6.1 17.4 0.14 0.07 0.11 上部 1.8 0.3 0.7 38.9 8.7 292.0 97.3 4.8 3.9 21.4 5.1 1.2 273.3 93.1 98.3 19.8 6.3 0.13 0.31 0.42 顶部 7.5 3.1 3.5 135 42.8 959.2 381.9 38.0 18.3 18.0 5.7 1.4 792.6 75.6 59.1 68.2 3.3 0.35 1.48 0.56 全段 2.0 0.4 0.5 38.5 10.3 331.4 78.0 4.9 3.4 21.6 6.0 1.1 233.5 90.8 72.0 15.2 11.3 0.14 0.24 0.28 全组 4.2 0.2 0.5 43.7 10.8 297.8 79.2 5.1 2.9 12.7 3.3 1.1 155.9 76.7 43.6 13.9 16.3 0.07 0.37 0.18 注：FePy据TS(总硫)含量的95%经化学式计算；(N/C)mol据TOC和TN(总氮)含量的摩尔值计算.
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表 3 四都坪牛蹄塘组黑色页岩中草莓状黄铁矿形态统计分析数据
Table 3. Morphological statistics of pyrite from the NTT Fm., Siduping, West Hunan
样品 高度(m) 数量(个) Dave(μm) Dmax(μm) Dmin(μm) 标准偏差(μm) 偏斜 密度(个/mm2) 第1段 SDP-0700 9.0 82 4.28 7.35 1.22 1.77 -0.13 182 SDP-0950 11.5 77 3.73 6.98 1.21 1.07 0.63 171 第2段 SDP-1900 21.0 64 3.54 10.23 1.82 2.24 1.40 142 SDP-1950 21.5 30 3.61 8.03 2.97 2.53 0.47 67 SDP-2050 22.5 51 4.32 9.12 2.46 2.15 0.34 113 SDP-2200 24.0 182 4.36 8.97 1.49 1.47 0.47 404 SDP-2250 24.5 55 4.08 7.17 2.28 1.87 0.35 122 SDP-2350 25.5 178 3.17 12.00 1.09 1.38 1.90 396 SDP-2400 26.0 165 2.71 6.65 0.91 1.12 0.80 367 SDP-2560 27.6 37 3.62 7.45 2.14 2.35 0.10 82 SDP-2900 31.0 85 3.99 9.34 1.40 1.75 0.92 189 SDP-2950 31.5 21 2.01 6.96 1.48 2.05 0.84 47 SDP-3000 32.0 49 4.13 10.95 1.77 2.02 1.25 109 SDP-3050 32.5 59 4.37 9.04 2.30 2.17 0.05 131 SDP-3300 35.0 145 3.19 7.05 1.26 1.13 0.82 322 SDP-3550 37.5 124 4.41 8.60 1.51 1.67 0.50 276 SDP-4050 42.5 231 3.90 8.06 0.65 1.33 0.60 513 第3段 13SSTW-14 62.0 79 3.43 7.56 1.25 1.51 0.71 176 13SSTW-13 62.7 159 4.21 8.88 1.28 1.38 0.86 353 13SSTW-11 64.1 54 2.46 9.65 1.27 1.77 2.38 120 13SSTW-10 64.8 92 3.85 7.29 1.49 1.34 1.14 204 13SSTW-9 65.5 200 3.94 12.96 1.22 1.55 1.65 444 13SSTW-8 66.2 172 3.76 12.69 1.51 1.57 2.00 382 13SSTW-5 68.3 57 4.17 6.90 1.67 1.30 0.05 127 13SSTW-2 70.4 50 3.14 12.78 1.51 2.05 3.00 111 注：Dave.平均粒径；Dmin.最小粒径；Dmax.最大粒径；高度指该样品距剖面底部距离，数量指测量的黄铁矿莓球个数；密度指单位面积内黄铁矿莓球个数.在剖面0~8.5 m，12.0~20.5 m，43.0~61.5 m和70.7~74.0 m的4个地层间隔段内样品中未见草莓状黄铁矿，故未列出.
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表 4 四都坪牛蹄塘组中硫化与非硫化时段微量元素数据比较
Table 4. Comparison of trace element data for recognized euxinic and non-euxinic intervals in the NTT Fm., Siduping, West Huanan
单元平均值 高度
(m)Mo
(μg/g)U
(μg/g)V
(μg/g)MoEF UEF VEF Mo/TOC
((μg/g)/%)U/TOC
((μg/g)/%)δEu P
(μg/g)Cr
(μg/g)Mn
(μg/g)(N/C)mol FePy/
FeT非硫化 非硫化A 2~8 105.7 25.1 1 327.1 206.4 14.8 18.2 14.1 3.4 1.2 72.2 99.8 23.9 0.03 0.72 非硫化B 12~21 46.9 11.6 180.7 90.9 6.2 1.5 7.2 1.8 1.0 122.5 59.6 24.0 0.02 0.46 非硫化C 43~61 31.4 8.9 285.3 40.7 2.7 1.7 16.7 4.8 1.1 116.3 82.3 39.3 0.10 0.11 硫化 硫化1 8~12 79.7 12.2 418.4 176.6 7.3 5.6 11.9 1.8 1.6 63.6 68.4 20.4 0.03 0.79 硫化2 21~43 38.0 9.8 164.8 60.5 4.1 0.9 8.2 2.2 1.0 164.3 70.0 39.6 0.04 0.44 硫化3 61~71 38.7 8.2 291.5 101.8 4.9 4.2 20.3 4.5 1.2 275.3 93.3 102.5 0.13 0.33 亚氧化-氧化 亚氧化-氧化 71~74 135.6 42.8 959.2 381.9 38.0 18.3 18.0 5.7 1.4 792.6 75.6 59.1 0.35 1.48
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