The Evolution of Trace Fossils across the Permian⁃Triassic Mass Extinction in Deep Marine from the Western Qinling
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摘要: 二叠纪‒三叠纪之交大灭绝事件前后的海洋遗迹生态学研究程度很高，但大部分研究局限于浅海环境，大灭绝前后深海环境中的遗迹化石面貌及其变化特征还不清楚.本文系统描述了采自我国西秦岭地区3条深海剖面地层中的遗迹化石，共计18属.研究表明大灭绝前后深海环境中的遗迹化石属种多样性没有发生变化，但早三叠世出现了大量的雕画迹.进一步通过生态空间利用和生态系统工程定量分析方法发现大灭绝之后，深海环境中的造迹生物的生态空间利用能力和生态系统工程参数相比于灭绝前表现出显著的下降，深生态阶层更为显著.与硅质浅海临滨宜居带避难所不同，深海生态系统工程师对大灭绝之后生态系统复苏的反馈效应较弱.深海环境不利于浅海机会分子的殖居，可能并不是大灭绝之后的避难所.Abstract: The ichnoecological variations across the Permian⁃Triassic mass extinction have been well studied, but most studies focused on shallow marine environments. The ichnological characteristic and its evolution in deep marine remain unclear across the mass extinction. This paper investigates abundant ichnofossils in three deep marine sections from the West Qinling area of North China, and systematically describes 18 ichnogenera. The study shows that the diversity of ichnofossil in deep marine did not change across the mass extinction, but a large number of graphoglyptid trace fossils appeared in the Early Triassic. Further quantitative analysis by using ecospace utilization and ecosystem engineering methods reveals that ecospace utilization and ecosystem engineering parameters of trace-making organisms apparently declined after the mass extinction, especially in the deep tier. Unlike the siliciclastic nearshore 'habitable zone' refuges, the feedback effects of ecosystem engineers on the recovery of in deep marine ecosystem after the mass extinction is relatively weak. The deep marine environmental setting is unfavourable to the colonization of shallow-sea opportunistic organisms and may not have acted as a refuge after the mass extinction.
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Key words:
- trace fossils /
- deep marine /
- refuge /
- ecosystem engineers /
- West Qinling area /
- ecology /
- stratigraphy
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图 2 西秦岭地区三条研究剖面晚二叠世至早三叠世地层柱状图及各类沉积构造和遗迹化石的地层分布
a.麻隆沟剖面晚二叠世毛毛隆组；b.扎油沟剖面早三叠世山尕岭群；c.隆务峡剖面早三叠世果木沟组
Fig. 2. Columnar sections of the Upper Permian to Lower Triassic successions, showing stratigraphic distributions of trace fossils and sedimentary structures
图 3 西秦岭地区二叠系‒三叠系麻隆沟和扎油沟剖面地层和沉积特征
a.毛毛隆组下部厚层含砾砂岩；b.毛毛隆组薄层板岩；c.毛毛隆组薄层砂岩与板岩互层；d.山尕岭群中粉砂质板岩中含砾石；e.山尕岭群中的波痕；f.山尕岭群砂岩底面上保存的槽模.a~c来自麻隆沟剖面；d~f为扎油沟剖面
Fig. 3. Field photos of the Upper Permian to Lower Triassic successions from the Malonggou and Zhayougou sections
图 4 西秦岭地区下三叠统隆务峡剖面地层和沉积特征
a.下三叠统果木沟组中段地层；b.果木沟组砂岩被安山岩顺层侵入，红色虚线为界线，字母A代表安山岩；c.照片b中安山岩特写；d.果木沟组中段砂岩底面大量槽模；e.双脊波痕，表面含有丰富遗迹化石，包括雕画迹；f.双脊波痕及其表面的丰富遗迹化石
Fig. 4. Field photos of the Lower Triassic successions from the Longwuxia section
图 5 西秦岭地区研究剖面晚二叠世至早三叠世遗迹化石标本照片(1)
a~b.Acanthorhaphe delicatula；c.Capodistria vettersi；d.Chondrites targionii；e.Circulichnis ligusticus；f.Cruziana isp.；g~i.Dendrotichnium seilacheri.图d和f来自麻隆沟剖面毛毛隆组，其余均来自隆务峡剖面果木沟组，图f为内生迹，其余均保存为底迹凸起
Fig. 5. Ichnotaxa (1) from the Upper Permian to Lower Triassic successions from the studied sections of western Qinling
图 6 西秦岭地区研究剖面晚二叠世至早三叠世遗迹化石标本照片(2)
a~b，e.Dictyodora zimmermanni；c.Diplichnites isp.；d.Halopoa imbricata；f.Helminthopsis abeli；g.Helminthoidichnites tenuis（黄色箭头）；h.Nereites missouriensis.图a~e来自麻隆沟剖面毛毛隆组，图f来自扎油沟剖面山尕岭群，图g-h来自隆务峡剖面果木沟组，均保存为内生迹
Fig. 6. Ichnotaxa (2) from the Upper Permian to Lower Triassic successions from the studied sections of western Qinling
图 7 西秦岭地区研究剖面晚二叠世至早三叠世遗迹化石标本照片(3)
a~b.Nereites missouriensis；c~d.Palaeophycus heberti；e.Protovirgularia longespicata；f，i.Planolites beverleyensis；g~h.Lorenzinia isp..图a~d和f来自扎油沟剖面山尕岭群，图g~h来自隆务峡剖面果木沟组，图i来自麻隆沟剖面毛毛隆组，均保存为底迹凸起
Fig. 7. Ichnotaxa (3) from the Upper Permian to Lower Triassic successions from the studied sections of western Qinling
图 8 西秦岭地区研究剖面晚二叠世至早三叠世遗迹化石标本照片（4）
a~b.Thalassinoides paradoxicus；c~e.？Ubinia isp..图d为照片c局部放大（红框）.图a~b来自麻隆沟剖面毛毛隆组，保存为内生迹；图c~e来自隆务峡剖面果木沟组，保存为底迹凸起
Fig. 8. Ichnotaxa (4) from the Upper Permian to Lower Triassic successions from the studied sections of western Qinling
图 9 二叠纪‒三叠纪大灭绝前后深海和硅质临滨环境中的造迹生物生态空间利用和生态系统工程三维立体图
a~b.西秦岭深海环境造迹生物生态空间利用（a）和生态系统工程（b）3D图.c~d.我国南祁连地区硅质临滨环境造迹生物生态空间利用（c）和生态系统工程（d）3D图，数据来自于Feng et al.（2022）
Fig. 9. Cubes of ecospace utilization and ecosystem engineering across the Permian-Triassic mass extinction in deep marine and siliciclastic environmental settings
表 1 西秦岭地区二叠纪‒三叠纪之交大灭绝前后深海遗迹类型及其歧异度
Table 1. Ichnotaxa and categories of architectural designs across the Permian-Triassic mass extinction in deep marine of the western Qinling
晚二叠世 早三叠世 遗迹属 歧异度类型 遗迹属 歧异度类型 Chondrites 轴状潜穴或者向下辐射延伸的束状潜穴 Acanthorhaphe 规则或不规则网络状雕画迹 Cruziana 双叶状的行迹和成对的沟槽 Capodistria 辐射状或莲花形潜穴 Dictyodora 具有复杂垂直方向蹼状构造的潜穴 Chondrites 轴状潜穴或者向下辐射延伸的束状潜穴 Diplichnites 行迹和抓痕 Circulichnis 简单水平移迹 Halopoa 具有简单垂直方向蹼状构造的水平潜穴 Dendrotichnium 单枝形蜿蜒雕画迹 Helminthopsis 简单水平移迹 Helminthopsis 简单水平移迹 Helminthoidichnites 简单水平移迹 Helminthoidichnites 简单水平移迹 Nereites 复杂的具有主动回填构造的水平潜穴 Lorenzinia 辐射状雕画迹 Palaeophycus 被动式充填的水平或倾斜潜穴 Nereites 复杂的具有主动回填构造的水平潜穴 Planolites 简单、主动充填的水平或倾斜潜穴 Palaeophycus 被动式充填的水平或倾斜潜穴 Phycodes 具有水平到垂直的分枝的水平的潜穴 Planolites 简单、主动充填的水平或倾斜潜穴 Protovirgularia V形移迹 Ubinia 单枝形蜿蜒雕画迹 Thalassinoides 迷宫式和箱式潜穴 
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表 2 西秦岭地区二叠纪‒三叠纪之交大灭绝前后深海遗迹生态空间占据和生态系统工程参数
Table 2. Parameters of ecospace utilization and ecosystem engineering across the Permian-Triassic mass extinction in deep marine of the western Qinling
晚二叠世 遗迹属 阶层 移动性 觅食方式 与沉积底质接触关系 改造沉积物方式 Chondrites 深 非自由 化学共生 压实 廊道扩散 Cruziana 半内栖 自由 非专业食沉积物 挖掘 再生型 Dictyodora 深 非自由 专业食沉积物 回填 传送型 Diplichnites 表层 自由 非专业食沉积物/捕食 压实 扩散 Halopoa 浅 自由 非专业食沉积物 压实 廊道扩散 Helminthopsis 半内栖 自由 非专业食沉积物 回填 传送型 Helminthoidichnites 半内栖 自由 非专业食沉积物 回填 传送型 Nereites 浅 自由 专业食沉积物 回填 传送型 Palaeophycus 浅 非自由 悬食/捕食 压实 廊道扩散 Planolites 浅 自由 非专业食沉积物 回填 传送型 Phycodes 半内栖 自由 专业食沉积物 回填 传送型 Protovirgularia 浅 自由 非专业食沉积物 侵入 扩散 Thalassinoides 深 自由 非专业食沉积物/捕食 挖掘 再生型 早三叠世 遗迹属 阶层 移动性 觅食方式 与沉积底质接触关系 改造沉积物方式 Acanthorhaphe 半内栖 自由 农艺 压实 廊道扩散 Capodistria 浅 自由 专业食沉积物 压实 廊道扩散 Chondrites 深 非自由 化学共生 压实 廊道扩散 Circulichnis 半内栖 自由 非专业食沉积物 回填 传送型 Dendrotichnium 半内栖 自由 农艺 压实 廊道扩散 Helminthopsis 半内栖 自由 非专业食沉积物 回填 传送型 Helminthoidichnites 半内栖 自由 非专业食沉积物 回填 传送型 Lorenzinia 半内栖 自由 农艺 压实 廊道扩散 Nereites 浅 自由 专业食沉积物 回填 传送型 Palaeophycus 浅 非自由 悬食/捕食 压实 廊道扩散 Planolites 浅 自由 非专业食沉积物 回填 传送型 Ubinia 半内栖 自由 农艺 压实 廊道扩散
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