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作者简介:

尚晓冬,男,1990年生。博士,副研究员,主要从事埃迪卡拉系古生物学与地层学研究。E-mail:shangxiaodong13@mails.ucas.ac.cn。

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目录contents

    摘要

    本文首次报道并描述了华南黄陵背斜西翼湖北宜昌兴山县乡儿湾剖面埃迪卡拉系陡山沱组二段中上部的燧石结核和条带中的疑源类微体化石,经系统鉴定共识别出8属8种,包括Appendisphaera grandis, Crassimembrana multitunica, Leiosphaeridia minutissima, Megasphaera inornata, Schizofusa zangwenlongii, Symphysosphaera basimembrana, Urasphaera nupta, Variomargosphaeridium gracile。该化石组合包含Liu Pengju and Moczydłowska (2019)所建立的第二生物组合带(Tanarium tuberosum-Schizofusa zangwenlongii组合带)的特征分子Schizofusa zangwenlongii,因此,可尝试性地与第二生物组合带相对比,但未见黄陵背斜东南翼陡山沱组二段的常见属Tianzhushania,据此推测黄陵背斜两翼在古环境上可能有所差异。

    Abstract

    The acritarch microfossils arediscovered from the chert nodules and bands in the Member Ⅱ of the Ediacaran Doushantuo Formation at Xiangerwan section in the western flank of Huangling anticline in the Xingshan County of Yichang, Hubei Province, South China. They are represented taxonomically by 8 species of 8 genera, including Appendisphaera grandis, Crassimembrana multitunica, Leiosphaeridia minutissima, Megasphaera inornata, Schizofusa zangwenlongii, Symphysosphaera basimembrana, Urasphaera nupta, Variomargosphaeridium gracile. Since the existence of Schizofusa zangwenlongii, it might be tentatively correlated with the second assemblage zone (Tanarium tuberosum-Schizofusa zangwenlongii) proposed by Liu and Moczydłowska (2019). However, it does not contain genus Tianzhushania which is common in the Member Ⅱ of the Doushantuo Formation in the southeastern flank of Huangling anticline, there might exist environmental variations between the two flanks of the Huangling anticline.

  • 近年来,华南埃迪卡拉纪早期地层陡山沱组陆续产出了大量的复杂具刺疑源类、多细胞藻类、丝状与球状蓝细菌、硫细菌、可能的海绵动物和后生动物胚胎、扇状海藻以及可能具有触手和类似肠道结构的后生动物等(Xiao Shuhai et al.,1998; Zhang Yun et al.,1998; Yin Leiming et al.,2007; Yuan Xunlai et al.,2011; Bailey et al.,2013; Xiao Shuhai et al.,2014; Yin Zongjun et al.,2015; Liu Pengju and Moczydłowska,2019; Shang Xiaodong and Liu Pengju,2022)。这些宏体与微体生物化石群为揭示地球早期多细胞真核生物的生命历史,阐明显生宙前夕的生物圈演化序幕提供了极其重要的证据。其中,具刺疑源类微体化石已经逐步应用于埃迪卡拉纪中下部地层的划分与对比(尹崇玉等,2009; McFadden et al.,2009; 刘鹏举等,2012; Liu Pengju et al.,20132014a2014b2022; Liu Pengju and Moczydłowska,2019; Zhou Chuanming et al.,2019; Xiao Shuhai and Narbonne,2020)。

  • 三峡地区的陡山沱组主要分布在黄陵背斜周缘,以往关于埃迪卡拉纪疑源类生物地层学的系统研究集中于黄陵背斜东翼和南翼,包括南晓峰河、北晓峰河、牛坪、王丰岗、陈家园子、田家园子、九龙湾、金瓜墩等剖面(图1)(尹崇玉等,2009; McFadden et al.,2009; Yin Chongyu et al.,2011; Liu Pengju et al.,20132014a2014b; 欧阳晴等,2015; Liu Pengju and Moczydłowska,2019)。虽然黄陵背斜西部地区青林口、九曲脑、五指山剖面也有相关化石的报道,但是目前的研究程度还很低(欧阳晴等,2015; Liu Pengju and Moczydłowska,2019; Ouyang Qing et al.,2021)。陡山沱组上部的地层序列在黄陵背斜的东部、西部地区出现明显变化,在碳同位素地层学方面也表现出一定的复杂性,而下部的地层在不同区域则能够进行良好的追踪对比(An Zhihui et al.,2015; Zhou Chuanming et al.,2017; Xiao Shuhai et al.,2017; Liu Pengju and Moczydłowska,2019; Zhou Chuanming et al.,2019; Ouyang Qing et al.,2021)。因此,有必要在黄陵背斜西部地区埃迪卡拉系开展深入的系统古生物学与生物地层学工作,以丰富早期微体古生物群的化石组合面貌,完善埃迪卡拉纪生物地层的区域与国际对比方案。

  • 图1 三峡地区地质简图,指示研究剖面及文中所提及的一些其他剖面的位置

  • Fig.1 Simplified geological map of the Yangtze Gorges area, indicating locations of the studied section and some other sections mentioned in the text

  • 1 地质背景与剖面简介

  • 埃迪卡拉系陡山沱组广泛出露于扬子板块,以细碎屑岩与碳酸盐岩为主要岩性,沉积于被动大陆边缘(Jiang Ganqing et al.,2011; Li Zhengxiang et al.,2013)。在三峡地区,陡山沱组主要出露于黄陵背斜周边(图1),厚度多在170~230 m之间,常被分为四个岩性段,自下而上为:一段为厚约5 m的“盖帽”白云岩,含大量帐篷构造和板裂构造(Jiang Ganqing et al.,2006; Chen Xiaoshuai et al.,2023)二段主要为互层状出现的黑色碳质页岩、白云质泥页岩与白云岩,厚80~120 m,含大量毫米~厘米级燧石结核或条带,结核内含有特异保存的化石,另外在中部的白云岩中发育一侵蚀暴露面,部分地区在该段的中下部含胶磷矿层(Zhu Maoyan et al.,2013; Liu Pengju et al.,20132014a);三段主要为中厚层状白云岩,含化石的燧石条带或结核发育,上部为条带状灰岩,厚40~60 m;四段为约10 m厚的黑色碳质、硅质页岩,偶夹透镜状白云岩。其中,陡山沱组一段和二段在黄陵背斜周缘的多条剖面中均能够很好地对比追踪(Zhu Maoyan et al.,2013; Liu Pengju and Moczydłowska,2019; Zhou Chuanming et al.,2019)。

  • 研究剖面位于黄陵背斜西北部地区兴山县高岚镇乡儿湾附近,埃迪卡拉系陡山沱组发育良好且连续,属于近岸边缘的内陆棚盆地环境沉积(图1)(Jiang Ganqing et al.,2011; An Zhihui et al.,2015)。乡儿湾剖面的陡山沱组一段盖帽白云岩厚约5 m,由下至上依次由含砾白云岩、具有孔洞的白云岩、帐篷结构与交错层理发育、顶部含重晶石的白云岩组成。盖帽白云岩之下为成冰系南沱组冰碛砾岩,二者界线清晰(图2a、b)。其中,在盖帽白云岩底部可见较多冰坠石,其与南沱组冰碛砾岩表现为渐变过渡,呈整合接触关系。陡山沱组二段下部以黑色泥岩为主,偶夹灰色薄层状泥质白云岩,厚约26 m(图2c);中部开始逐渐出现中层状泥质白云岩,偶含厘米级大小的燧石结核,厚约12 m(图2d);上部为黑色页岩与中层状泥质白云岩呈现不等厚互层,向上表现为泥质白云岩开始增多,并含有大量的燧石结核与条带,厚约20 m(图2e)。其上为陡山沱组三段灰色中厚层状白云岩,二段向三段岩性过渡表现为突变面,二者界线截然清晰(图2f)。

  • 2 材料与方法

  • 本文报道的化石产自于乡儿湾剖面埃迪卡拉系陡山沱组二段黑色页岩与泥质白云岩中的燧石结核与条带,以陡山沱组二段与三段界线计为0 m标志,自上而下采集共计19层燧石结核或条带,14个层中发现疑源类微体化石(图3)。样品主要集中在陡山沱组二段的中上部,在下部由于较为缺乏燧石结核而导致样品稀少(图3)。样品采用平行层面方向切片,磨制厚约50 μm的岩石薄片共计约380张,在蔡司Axio Imager.A2透射光显微镜下观察。化石标本由显微镜附带的蔡司AxioCam MRc5数码相机拍照,度量在AxioVision Rel.4.8软件的辅助下完成。全部标本均保存于中国地质科学院地质研究所(标本号前缀为IGCAGS)。

  • 图2 湖北宜昌兴山地区乡儿湾剖面野外露头照片

  • Fig.2 Outcrop photos of the Xiangerwan section in the Xingshan area of Yichang City, Hubei Province, South China

  • (a)—成冰系南沱组冰碛砾岩;(b)—南沱组冰碛砾岩与陡山沱组一段盖帽白云岩呈整合接触关系;(c)—陡山沱组二段下部黑色页岩,偶夹灰色薄层状泥质白云岩;(d)—剖面位置~29.7 m处的陡山沱组二段含燧石结核泥质白云岩,箭头指示厘米级大小的燧石结核;(e)—陡山沱组二段上部含薄层燧石条带的泥质白云岩;(f)—陡山沱组二段含燧石条带泥质白云岩与三段灰色中厚层状白云岩界线

  • (a) —the Cryogenian Nantuo Formation diamictite; (b) —the Ediacaran Doushantuo Formation cap dolostone overlies conformably on the Cryogenian Nantuo Formation diamictite; (c) —black shales in the lower part of Member Ⅱ of the Doushantuo Formation, occasionally containing grey, thin-bedded argillaceous dolostone; (d) —argillaceous dolostone containing chert nodules at~29.7 m in the Member Ⅱ of the Doushantuo Formation, arrows denote the centimeter-sized chert nodules; (e) —argillaceous dolostone containing thin-bedded chert bands in the upper part of Member Ⅱ of the Doushantuo Formation; (f) —the boundary between the Member Ⅱ and Member Ⅲ of the Doushantuo Formation

  • 3 微体化石组成

  • 乡儿湾剖面陡山沱组二段虽然燧石结核与条带比较发育,但微体化石却较为稀少。经系统观察,仅发现微体化石共8属8种,即:Appendisphaera grandisCrassimembrana multitunicaLeiosphaeridia minutissimaMegasphaera inornataSchizofusa zangwenlongiiSymphysosphaera basimembranaUrasphaera nuptaVariomargosphaeridium gracile(图4~图7)。

  • 虽然化石属种较为单调,但部分化石属种不仅是华南扬子板块多个地区埃迪卡拉系中的常见分子,而且在全球不同的古大陆也具有广泛的分布。其中,Appendisphaera grandisMoczydłowska et al.(1993)在西伯利亚埃迪卡拉系发现,被认为具有实心的刺状突起,并作为模式种首次建立起来。伴随着显微成像系统技术的进步,Moczydłowska(2005)观察到该种的刺突为空心状态,并将其种征进行修订。随后,Appendisphaera grandis在全球范围内包括澳大利亚(Willman and Moczydłowska,2008)、印度(Prasad and Asher,2016)、蒙古北部(Anderson et al.,20172019)、华南扬子板块(陈寿铭等,2010; Xiao Shuhai et al.,2014; Ouyang Qing et al.,20172019; Liu Pengju and Moczydłowska,2019; Shang Xiaodong et al.,2019; 尚晓冬和刘鹏举,2020; 刘海等,2021)等多个板块或地区的埃迪卡拉纪地层中被广泛发现,显示出重要的生物地层学意义。

  • 图3 湖北宜昌兴山地区乡儿湾剖面埃迪卡拉系陡山沱组地层序列与化石产出层位

  • Fig.3 The Ediacaran stratigraphic succession and fossil horizons of the Xiangerwan section in the Xingshan area of Yichang City, Hubei Province, South China

  • Variomargosphaeridium属以往在黄陵背斜东部地区晓峰河剖面、王丰岗剖面陡山沱组三段的下部较为常见(Liu Pengju et al.,2014b)。随后,欧阳晴等(2015)在黄陵背斜西南地区青林口、南部地区金瓜墩和九龙湾剖面的陡山沱组二段中上部也发现了此类化石(图1)。类似地,以往常见于晓峰河剖面陡山沱组三段下部的Symphysosphaera basimembrana在本次研究的化石组合中也被发现。这一现象表明以上属种可能具有较长的地层延限,可下延至陡山沱组二段中。随着研究工作的深入开展,当前新发现的化石材料表明陡山沱组二段中的化石组成可能更加丰富,从而增加了三峡地区埃迪卡拉系陡山沱组下部化石物种的多样性。

  • 4 生物地层意义

  • 在华南三峡地区的陡山沱组中,已识别出来上部和下部两个具刺疑源类生物带或组合(Liu Pengju et al.,20132014a2014b)。下部生物带(Tianzhushania spinosa带)由它的代表性物种Tianzhushania spinosa作为此带的主要分子,伴有属Tianzhushania的其他类型大量出现,局限在陡山沱组二段;上部生物带(Tanarium conoideum-Hocosphaeridium scaberfacium-Hocosphaeridium anozos带)在陡山沱组三段中被识别出来,以大量出现HocosphaeridiumTanarium、而不含Tianzhushania为主要特征,物种丰度与分异度均比下部生物带更加丰富(Liu Pengju et al.,2014b)。然而,随着化石材料的不断积累,特征分子Hocosphaeridium anozosHocosphaeridium属在陡山沱组二段也有发现,推动着以往所建立的上、下生物带的修订工作(Xiao Shuhai et al.,2014; Hawkins et al.,2017; Liu Pengju and Moczydłowska,2019; 刘海等,2021)。基于三峡地区多条剖面的古生物学与地层学数据,Liu Pengju and Moczydłowska(2019)选取某些全球广布的属种建立了4个新的生物组合带,用于中下埃迪卡拉系的划分。这些生物带自下至上包括:①Appendisphaera grandis-Weissiella grandistella-Tianzhushania spinosa组合带;②Tanarium tuberosum-Schizofusa zangwenlongii组合带;③Tanarium conoideum-Cavaspina basiconica组合带;④Tanarium pycnacanthum-Ceratosphaeridium glaberosum组合带。

  • 本研究在黄陵背斜西部乡儿湾剖面陡山沱组二段的多层燧石结核与条带中均未发现属Tianzhushania、也并未发现属Hocosphaeridium,呈现出了与黄陵背斜东部地区不同的生物组合面貌,表明当前的生物组合尚不能与先前建立的Tianzhushania spinosa组合带相对比。然而,在距离研究剖面并不遥远的黄陵背斜东部地区多条剖面上,Tianzhushania spinosa以及Tianzhushania属内其他类型的化石广泛且大量地分布于陡山沱组二段中,是下部组合的典型常见分子(丰度通常大于60%)(McFadden et al.,2009; Liu Pengju et al.,2013)。与黄陵背斜东部地区相比,乡儿湾剖面陡山沱组二段产出化石的丰度和分异度较低,呈现不同的生物群面貌特征,可能是由于黄陵背斜东、西两翼地区在埃迪卡拉纪早期古环境的不同所导致。在岩石地层学方面,本次研究剖面的化石产自陡山沱组二段的中上部至顶部位置(图3)。化石组合产出层位较高,接近于陡山沱组二段与三段的界线位置,因此推断常见于东部地区的Tianzhushania属在当前研究剖面中可能已经绝灭消失。鉴于本次研究中的化石组合中包含Schizofusa zangwenlongii,该种为新建立的四个化石带中的第二生物组合带(Tanarium tuberosum-Schizofusa zangwenlongii组合带)的特征分子,暂将此次在陡山沱组二段中上部发现的微体化石组合对比于第二生物组合带。

  • 5 系统古生物学描述

  • 疑源类  Group Acritarcha Evitt,1963

  •  Genus AppendisphaeraMoczydłowska,Vidal and Rudavskaya,1993,emend. Moczydłowska,2005

  •  Species AppendisphaeragrandisMoczydłowska,Vidal and Rudavskaya,1993,emend. Moczydłowska,2005

  • (图4a、b)

  • non 2014 Appendisphaera grandis Moczydłowska et al., 1993, Shukla and Tiwari, p.215, fig.4D~E (=Appendisphaera tenuis) ;

  • non 2016 Appendisphaera grandis, Sharma et al., fig.4B (=Appendisphaera tenuis) ;

  • 2016 Appendisphaera grandis, Prasad and Asher, p.42, pl. II, figs.3~4;

  • 2017 Appendisphaera grandis, Anderson et al., fig.2B;

  • 2017 Appendisphaera fragilis, Ouyang et al., fig.8D~F;

  • non 2017 Appendisphaera? Hemisphaerica, Hawkins et al., fig.9C~D;

  • 2019 Appendisphaera grandis,Liu and Moczydłowska,pp.48~54,figs.21~23,及其文中同异名;

  • 2019 Appendisphaera grandis, Anderson et al., pp.507~509, fig.6A~D;

  • 2019 Appendisphaera grandis, Ouyang et al., fig.8I~K;

  • 2019 Appendisphaera grandis, Shang et al., p.7, fig.3;

  • 20 20 Appendisphaera grandis,尚晓冬和刘鹏举,156页,图4;

  • 20 21 Appendisphaera grandis,刘海等,图5.4;

  • 2021 Appendisphaera grandis, Moczydłowska and Liu, fig.2a~d;

  • 2021 Appendisphaera grandis, Ouyang et al., figs.10M~P;

  • 2022 Appendisphaera grandis, Shi et al., figs.7A~B;

  • 2022 Appendisphaera grandis, Ye et al., pp.20~21, fig.10;

  • 2022 Appendisphaera grandis, Xiao et al., pp.9~13, fig.7;

  • 2023 Appendisphaera grandis, Golubkova, pl.7, fig.1.

  • 材料 一枚保存良好的标本。

  • 描述 膜壳大,椭球形,表面具有丰富、稠密且均匀分布的刺状突起。刺突较长,形态简单,呈细的柱管状,具有相同的形态。刺突在其基部无明显的膨大构造,向尖端略微变细。刺管呈中空,刺的末端细且尖。部分刺管被埋藏保存过程中的有机质颗粒附着沉淀,导致其与膜壳内部空间的连通状态不清晰。在膜壳内部可见一些大小不等的球形体,其分布不均,或松散或聚集排列。

  • 度量 膜壳直径约390 μm;刺长约55 μm;刺长与膜壳直径比值约14.1%。

  • 讨论  Appendisphaera grandis以具有密集排列的、细长、柱状刺突为主要特征,刺突在基部略微加宽,向尖端逐渐变细(Moczydłowska,2005)。Sharma et al.(2016)Shukla and Tiwari(2014)中被鉴定作Appendisphaera grandis的化石为同一枚标本。该标本与模式种相比,其刺突较短且不够稠密,因而归入更为Appendisphaera tenuis合适。Liu Pengju and Moczydłowska(2019)与Shang Xiaodong et al.(2019)曾将Hawkins et al.(2017)在湖南四都坪地区鉴定为Appendisphaera? hemisphaerica的标本归入Appendisphaera grandis,由于其所附插图的标本具有稠密分布的长柱状刺突。随后,Xiao Shuhai et al.(2022)Hawkins et al.(2017)中的附图标本进行检查,通过显微镜下调整焦距的观察,确认了该标本具有基部相互分隔的双形刺突,其单个刺突由基部的膨大结构(直径约4 μm)和位于尖端的细长刺状突起(直径约1 μm)构成,因此仍将其鉴定为Appendisphaera? hemisphaericaXiao Shuhai et al.,2022)。虽然Appendisphaera grandis的一些刺突在基部会稍微膨大,但总体上较为窄小(约1~3 μm; Liu Pengju and Moczydłowska,2019; Shang Xiaodong et al.,2019),并不表现为明显的双形刺突。综上,Hawkins et al.(2017)在湖南四都坪地区发现的Appendisphaera? hemisphaerica标本符合A.? hemisphaerica的鉴定特征。

  • 图4 乡儿湾剖面埃迪卡拉系陡山沱组二段部分微体化石照片

  • Fig.4 Photomicrographs of microfossils from the Ediacaran Doushantuo Formation at Xiangerwan section

  • (a、b)—Appendisphaera grandis Moczydłowska,Vidal and Rudavskaya,1993,emend. Moczydłowska,2005,标本号IGCAGS-XE-8.1-0201,薄片号XE-8.1-02;(b)—图4a中箭头所指部分的放大,示其稠密分布的柱管状刺突;(c)—Leiosphaeridia minutissima Naumova,1949,emend. Jankauskas,Mikhailova and Hermann,1989,标本号IGCAGS-XE-4.5-0301,薄片号XE-4.5-03;(d、e)—Crassimembrana multitunica Ouyang,Zhou,Xiao,Guan,Chen,Yuan and Sun,2021,标本号IGCAGS-XE-5.3-0301,薄片号XE-5.3-03;(e)—图4d中箭头所指部分的放大,示其多层状胶质膜

  • (a, b) —Appendisphaera grandis Moczydłowska, Vidal and Rudavskaya, 1993, emend. Moczydłowska, 2005, No.IGCAGS-XE-8.1-0201, thin section XE-8.1-02; (b) —magnified fragment shown by the arrow in Fig.4a, showing the densely distributed, cylindrical processes; (c) —Leiosphaeridia minutissima Naumova, 1949, emend. Jankauskas, Mikhailova and Hermann, 1989, No.IGCAGS-XE-4.5-0301, thin section XE-4.5-03; (d, e) —Crassimembrana multitunica Ouyang, Zhou, Xiao, Guan, Chen, Yuan and Sun, 2021, No.IGCAGS-XE-5.3-0301, thin section XE-5.3-03; (e) —magnified fragment shown by the arrow in Fig.4d, showing the multi-laminate vesicle wall

  •  Genus Crassimembrana Ouyang,Zhou,Xiao,Guan,Chen,Yuan and Sun,2021

  •  Species Crassimembranamultitunica Ouyang,Zhou,Xiao,Guan,Chen,Yuan and Sun,2021

  • (图4d、e)

  • 2017 possible Tianzhushania spinosa, Hawkins et al., fig.7A~B;

  • non 2019 Laminasphaera capillata Liu and Moczydłowska, p.124, fig.67A~H;

  • 2021 Crassimembrana multitunica gen. et sp. nov. Ouyang et al., p.25, fig.13J~O.

  • 材料 一枚被压扁变形的标本。

  • 描述 膜壳巨大,遭受压扁变形,原始呈球形,膜壳壁内层薄而坚实,外层围绕膜壳内壁包裹着一圈厚的、多层状的、呈胶质的膜。膜壳不具有刺突或其他装饰。由于膜壳的形变与生物有机质的降解,位于外层的胶质膜发生起皱,造成外膜部分分层不明显。

  • 度量 膜壳直径约1800 μm;膜壳厚度约40 μm。

  • 讨论  Crassimembrana multitunica主要特征为膜壳巨大且外围具有一圈厚的、多层状的、呈胶质的膜。本次研究中的标本虽遭受较为强烈的压扁变形,化石周边也保存了呈黑色的有机质颗粒或残余生物膜碎片,但化石标本的膜结构仍被保存下来,在高倍镜下可见其外圈膜壳呈胶质多层状的膜(图4d、e)。Hawkins et al.(2017)所附插图中被鉴定为可能的Tianzhushania spinosa的膜壳壁较厚,由多层状微弯曲的薄层构成,且缺乏刺突,因此插图标本很可能为Crassimembrana multitunica。以往出现在埃迪卡拉纪地层中的部分微体化石膜壳也具有与此相似的多层膜结构,如Laminasphaera capillata Liu and Moczydłowska2019。然而,L. capillata因其具有细长且柔软、纤维状的刺突而显著区别于Crassimembrana multitunicaOuyang Qing et al.(2021)将Liu Pengju and Moczydłowska(2019)中的两枚Laminasphaera capillata标本归入Crassimembrana multitunica,认为其不具有刺突且具有相似于C. multitunica的多层膜结构。经过对以上两枚标本的镜下观察与重新检查,发现化石膜壳的多层状结构是截然的且顺层平直延展、分层界线清晰。相比之下,Crassimembrana multitunica膜壳多层状薄层的分层界线通常稍微波状,相邻的薄层大多相互平行,有时候弯曲并相互接触、发生黏连。这一现象曾被Ouyang Qing et al.(2021)认为是由化石埋藏原因导致,而在很多保存完好的C. multitunica标本上均具有相似的膜壳结构。因此,多层状的薄层膜发生弯曲并相互接触或粘连的现象应视为该类化石的生物学特征,其不同于分层界线截然且延展清晰的Laminasphaera capillata多层膜结构。综上,Liu Pengju and Moczydłowska(2019)中的两枚Laminasphaera capillata标本鉴定合理,不应被归入Crassimembrana multitunica

  •  Genus LeiosphaeridiaEisenack,1958,emend. Turner,1984

  •  Species Leiosphaeridiaminutissima Naumova,1949,emend. Jankauskas,Mikhailova and Hermann,1989

  • (图4c)

  • 1949 Leiosphaeridia minutissima sp. nov. Naumova, pl.1, fig.1;

  • 1989 Leiosphaeridia minutissima Naumova, 1949, emend. Jankauskas et al., pp.79~80, pl.9, figs.1~4, 11;

  • 2005 Leiosphaeridia minutissima Naumova, 1949, Grey, p.184, fig.63D;

  • 2009 Leiosphaeridia minutissima Naumova, 1949, emend. Yankauskas et al., 1989, Vorobeva et al., p.185, fig.14.9 (Yankauskas=Jankauskas) ;

  • non 2012 Leiosphaeridia minutissima Naumova, 1949, emend. Yankauskas et al., 1989, Sergeev et al., pp.332~333, pl.26.7 (Yankauskas=Jankauskas) ;

  • 2015 Leiosphaeridia minutissima Naumova, 1949, emend. Yankauskas et al., 1989, Schopf et al., p.724, fig.13.10 (Yankauskas=Jankauskas) ;

  • 2019 Leiosphaeridia minutissima Naumova, 1949, emend. Jankauskas et al., 1989; Shang et al., p.24, fig.21A.

  • 材料  11枚保存完好的标本。

  • 描述 小的球形膜壳,具有单层、表面光滑的壁。化石个体通常聚集,以丛状出现。

  • 度量 膜壳直径25~35 μm,壁厚度约1 μm。

  • 讨论 基于膜壳大小与膜壳壁厚度,Jankauskas et al.(1989)将元古代光面球疑源类Leiosphaeridia划分为4个种:L. minutissima(壁薄,膜壳直径小于70 μm);L. tenuissima(壁薄,膜壳直径70~200 μm);L. crassa(壁厚,膜壳直径小于70 μm);L. jacutica(壁厚,膜壳直径70~800 μm)。乡儿湾剖面埃迪卡拉系产出的光面球膜壳直径为25~35 μm,因而归入Leiosphaeridia minutissimaSergeev et al.(2012)中被鉴定为Leiosphaeridia minutissima的标本具有不光滑的膜壳,表面看起来可能还具有很短的刺突,因此将其排除出去。

  •  Genus Megasphaera Chen and Liu,1986,emend. Xiao,Zhou,Liu,Wang and Yuan,2014

  •  Species Megasphaerainornata Chen and Liu,1986,emend. Xiao,Zhou,Liu,Wang and Yuan,2014

  • (图5a~f)

  • 2020 Megasphaerainornata Chen and Liu,1986,emend. Xiao et al.,2014,尚晓冬和刘鹏举,158页,图5C,及其文中同异名;

  • 2020 Acritarcha gen. et sp. indet. 尚晓冬和刘鹏举,159页,图6D~E;

  • 2020 Megasphaera inornata Chen et Liu,1986,emend. Xiao et al.,2014,阳乐等,9~10页,图3;

  • 2021 Megasphaera inornata Chen and Liu, 1986, emend. Xiao et al., 2014; Ouyang et al., fig.16A~D;

  • 2022 Megasphaera inornata Chen and Liu, 1986, emend. Xiao et al., 2014, Ye et al., pp.47~49, fig.29.

  • 材料  23枚保存完好的标本。

  • 描述 膜壳巨大,呈球形或椭球形,膜壳壁薄,表面未见任何刺饰,通常光滑。

  • 度量 膜壳直径1200~2300 μm。

  • 讨论  Megasphaera inornata以大型的光滑球形膜壳为显著特征,膜壳内部可具有一个或多个子细胞(Xiao Shuhai et al.,2014)。根据以往报道标本的测量数据,M. inornata的膜壳直径表现出很大的范围,最大可达2300 μm(Ye Qin et al.,2022)。本次研究中发现的标本膜壳巨大,内部通常空洞,可能保存有暗色、不成形的有机残留物,膜壳内部一般不具有球状内含物。膜壳可能由双层膜构成,分为内膜和外膜,均很薄且光滑。由于膜壳的外围缺乏一圈厚的、多层状呈胶质的膜,因此该类标本不同于Crassimembrana multitunica,但部分标本很有可能是由C. multitunica膜壳的外膜受埋藏因素脱落并未保存下来而导致,二者的膜壳均巨大且其直径范围接近。

  •  Genus Schizofusa Yan,1982

  •  Species Schizofusazangwenlongii Grey,2005

  • (图5g~i)

  • 1992 Schizofusa sp. Zang, Zang and Walter, p.96, fig.71;

  • 2005 Schizofusa zangwenlongii sp. nov. Grey, p.191, figs.71~72;

  • 2011 Schizofusa zangwenlongii Grey, 2005, Sergeev et al., p.1007, fig.9.8;

  • 2013 Schizofusa zangwenlongii, Liu et al., fig.11M;

  • 2014a Schizofusa zangwenlongii, Liu et al., fig.9D;

  • 2014b Schizofusa zangwenlongii, Liu et al., p.132, fig.108;

  • 2015 Schizofusa zangwenlongii; 王楚楚等,figs.3~4;

  • 2019 Schizofusa zangwenlongii, Shang et al., p.26, fig.21B.

  • 材料  4枚保存完好的标本。

  • 描述 巨大的、椭球形或纺锤形膜壳,表面光滑,无刺饰。沿纵轴具有一道开裂缝,顺着开裂边平滑或者微微褶皱。开裂缝的开口宽阔,开裂的两瓣膜壳在末端相连接,不同标本的膜壳开裂程度不同。

  • 度量 膜壳长度1300~2300 μm,膜壳宽度400~800 μm。

  • 讨论  Schizofusa属是阎玉忠于1982年依据中元古界串岭沟组的标本所建,其形态特征为:膜壳梭形至椭圆形,单层壁,厚薄不一,大小各异,表面光滑,颗粒状的少见;沿纵轴具有开裂缝,一般一侧开裂,顺开裂边有或无边褶(阎玉忠,1982)。此类化石大量出现于华北燕山地区长城系常州沟组、串岭沟组(阎玉忠,19821991; 罗其玲等,1985; Miao Lanyun et al.,2019),在瑞典拉伸系Visingso群的上部地层中也有发现(Loron and Moczydłowska,2018)。近年来随着研究的深入,在华南三峡地区(Liu Pengju et al.,20132014a2014b; 王楚楚等,2015; 当前研究)和贵州松林地区(Shang Xiaodong et al.,2019)、澳大利亚(Zang Wenlong and Walter,1992; Grey,2005)、西伯利亚(Sergeev et al.,2011)埃迪卡拉纪地层中也逐渐发现了大量的Schizofusa zangwenlongii化石材料。对于S. zangwenlongii膜壳大小,从以往所发表的埃迪卡拉纪数据发现其膜壳长度为200~400 μm,宽度约50~200 μm。然而,本次研究发现的化石标本具有巨大的膜壳,比以往标本均大得多。基于当前标本的膜瓣长度和宽度,其大小范围非常接近于本研究同时发现的Megasphaera inornata。而具有巨大膜壳和裂缝状开口的Schizofusa zangwenlongii是否是由Megasphaera inornata开裂发展而来,因当前化石材料不够充足而尚不清楚。

  • 图5 乡儿湾剖面埃迪卡拉系陡山沱组二段部分微体化石照片

  • Fig.5 Photomicrographs of microfossils from the Ediacaran Doushantuo Formation at Xiangerwan section

  • (a~f)—Megasphaera inornata Chen and Liu,1986,emend. Xiao,Zhou,Liu,Wang and Yuan,2014;(a)—标本号IGCAGS-XE-13.8-0301,薄片号XE-13.8-03;(b)—标本号IGCAGS-XE-5-0601,薄片号XE-5-06;(c)—标本号IGCAGS-XE-15.5-0601,薄片号XE-15.5-06;(d)—标本号IGCAGS-XE-14.8-0101,薄片号XE-14.8-01;(e)—标本号IGCAGS-XE-4.5-0802,薄片号XE-4.5-08;(f)—标本号IGCAGS-XE-10.5-0101,薄片号XE-10.5-01;(g~i)—Schizofusa zangwenlongii Grey,2005; 箭头示其裂缝状的开口;(g)—标本号IGCAGS-XE-15.5-0101,薄片号XE-15.5-01;(h)—标本号IGCAGS-XE-13.8-0601,薄片号XE-13.8-06;(i)—标本号IGCAGS-XE-15.5-0502,薄片号XE-15.5-05

  • (a~f) —Megasphaera inornata Chen and Liu, 1986, emend. Xiao, Zhou, Liu, Wang and Yuan, 2014; (a) —No.IGCAGS-XE-13.8-0301, thin section XE-13.8-03; (b) —No.IGCAGS-XE-5-0601, thin section XE-5-06; (c) —No.IGCAGS-XE-15.5-0601, thin section XE-15.5-06; (d) —No.IGCAGS-XE-14.8-0101, thin section XE-14.8-01; (e) —No.IGCAGS-XE-4.5-0802, thin section XE-4.5-08; (f) —No.IGCAGS-XE-10.5-0101, thin section XE-10.5-01; (g~i) — Schizofusa zangwenlongii Grey, 2005; arrows denote the slit-like apertures; (g) —No.IGCAGS-XE-15.5-0101, thin section XE-15.5-01; (h) —No.IGCAGS-XE-13.8-0601, thin section XE-13.8-06; (i) —No.IGCAGS-XE-15.5-0502, thin section XE-15.5-05

  • 图6 乡儿湾剖面埃迪卡拉系陡山沱组二段部分微体化石照片

  • Fig.6 Photomicrographs of microfossils from the Ediacaran Doushantuo Formation at Xiangerwan section

  • (a~c)—Symphysosphaera basimembrana Liu,Xiao,Yin,Chen,Zhou and Li,2014,标本号IGCAGS-XE-5.3-0201,薄片号XE-5.3-02;(b、c)分别为图6a中红色与蓝色箭头所指部分的放大,示其围绕膜壳的一层小的、球状的细胞;(d~f)—Urasphaera nupta Liu,Xiao,Yin,Chen,Zhou and Li,2014,标本号IGCAGS-XE-10.7-0101,薄片号XE-10.7-01;(e、f)—图6d中刺状突起在不同焦距下的放大,示其较长的、似裙状基部膨大

  • (a~c) —Symphysosphaera basimembrana Liu, Xiao, Yin, Chen, Zhou and Li, 2014, No.IGCAGS-XE-5.3-0201, thin section XE-5.3-02; (b, c) —magnified fragments shown by red and blue arrows in Fig.6a, respectively, showing the layer of small, spheroidal cells surrounding the vesicle; (d~f) —Urasphaera nupta Liu, Xiao, Yin, Chen, Zhou and Li, 2014, No.IGCAGS-XE-10.7-0101, thin section XE-10.7-01; (e, f) —magnified fragments of processes in Fig.6d at different focal levels, showing the long, skirt-like basal expansions

  •  Genus Symphysosphaera Yin,1992,emend. Liu,Xiao,Yin,Chen,Zhou and Li,2014

  •  Species Symphysosphaerabasimembrana Liu,Xiao,Yin,Chen,Zhou and Li,2014

  • (图6a~c)

  • 2014b Symphysosphaera basimembrana sp. nov. Liu et al., p.134, fig.111.

  • 材料 一枚保存良好的标本。

  • 描述 球状的结构,具有一层小的、球状的细胞围绕着膜壳。球状小细胞层位于基部的膜上,不具有外膜。

  • 度量 总体直径约90 μm;外围球状小细胞直径4~9 μm。

  •  Genus UrasphaeraNagovitsin and Moczydłowska in Moczydłowska and Nagovitsin,2012

  •  Species Urasphaeranupta Liu,Xiao,Yin,Chen,Zhou and Li,2014

  • (图6d~f)

  • 2014b Urasphaera nupta sp. nov. Liu et al., pp.119~128, figs.89.2, 90.

  • 材料 一枚保存状态良好的标本。

  • 描述 膜壳大,原始呈球状,形变至椭球状,具有许多基部分隔的、锥形刺状突起。刺突具有膨大的末端,刺突基部加宽呈锥形,向刺突的颈部逐渐变窄,在靠近末端膨大部位收缩。刺突呈中空状态,与膜壳内部自由连通。

  • 度量 膜壳直径166~283 μm;刺长22~69 μm;刺长与膜壳直径比值13.3%~24.4%;刺突基部宽度13~18 μm;刺突颈部收缩宽度3~5 μm;刺突膨大的末端宽度10~12 μm。

  • 讨论  Urasphaera属包括3个种,即模式种Urasphaera capitalisU. fungiformisU. nupta,其中模式种U. capitalis的刺突在颈部之下表现为柱状,而U. fungiformisU. nupta的刺突在基部部位显著加宽,向颈部逐渐变细。U. nupta能够区分于U. fungiformis因其刺突具有一个较长的、轮廓类似新娘礼服的裙状基部。当前标本的刺突在基部和尖端部位均膨大加宽,腰部更接近尖端,基部较长且似裙状,同Urasphaera nupta特征相符,故归入该种。

  •  Genus Variomargosphaeridium Zang in Zang and Walter,1992,emend. Xiao,Zhou,Liu,Wang and Yuan,2014

  •  Species Variomargosphaeridiumgracile Xiao,Zhou,Liu,Wang and Yuan,2014

  • (图7)

  • 2007? Meghystrichosphaeridium multispinatum,尹崇玉等,图版15,图1~4;

  • 2014 Variomargosphaeridium gracile, Xiao et al., pp.57, 58, fig.36.1, 36.2;

  • non 2014 Variomargosphaeridium gracile; Xiao et al., pp.57, 58, fig.36.3~36.6 (=Knollisphaeridium bifurcatum) .

  • 材料 两枚标本,保存较好。

  • 描述 膜壳小至中等大小,呈球状,具有均匀分布的刺状突起。刺突基部加宽,向末端逐渐变细。刺突末端连续多次分叉,发育很细的小分枝。刺突多被微小有机质颗粒充填,不清楚是否与膜壳内部自由连通。

  • 度量 膜壳直径60~242 μm;刺长9~14 μm;刺长与膜壳直径比值6%~23%;刺突基部宽度3~7 μm。

  • 讨论 被Xiao Shuhai et al.(2014)鉴定作Variomargosphaeridium gracile的部分标本(其文中插图36.3~36.6)刺突末端并非多次分叉,而是规则的二歧式分枝,故将其归入Knollisphaeridium bifurcatum更为合适。

  • 6 结论

  • (1)本文通过对湖北宜昌兴山地区黄陵背斜西翼乡儿湾剖面埃迪卡拉系陡山沱组二段中上部产出的燧石结核与条带进行切片研究,经系统古生物学鉴定与描述共识别出8属8种,包括Appendisphaera grandisCrassimembrana multitunica Leiosphaeridia minutissimaMegasphaera inornataSchizofusa zangwenlongiiSymphysosphaera basimembranaUrasphaera nuptaVariomargosphaeridium gracile

  • (2)新发现的微体化石增加了三峡地区埃迪卡拉系陡山沱组下部化石物种的多样性。部分化石属种不仅是华南扬子板块多个地区埃迪卡拉系中的常见分子,而且在全球不同的古大陆也具有广泛的分布,显示出重要的生物地层学意义。

  • (3)本次研究中的化石组合中包含Schizofusa zangwenlongii,该种为Liu Pengju and Moczydłowska(2019)所建立的第二生物组合带(Tanarium tuberosum-Schizofusa zangwenlongii组合带)的特征分子,暂将此次在陡山沱组二段中上部发现的微体化石组合对比于第二生物组合带。与黄陵背斜东部地区相比,该化石组合的属种丰度和分异度较低,呈现不同的生物群面貌特征,可能是由于黄陵背斜东、西两翼地区在埃迪卡拉纪早期古环境的不同所导致。

  • 图7 Variomargosphaeridium gracile Xiao,Zhou,Liu,Wang and Yuan,2014

  • Fig.7 Variomargosphaeridium gracile Xiao, Zhou, Liu, Wang and Yuan, 2014

  • (a)—标本号IGCAGS-XE-10.2-0101,薄片号XE-10.2-01;(b~d)—标本号IGCAGS-XE-12.7-0101,薄片号XE-12.7-01;(c、d)分别为图7b中红色与蓝色箭头所指部分的放大,示其连续二次或多次分叉的、很细的刺状突起

  • (a) —No.IGCAGS-XE-10.2-0101, thin section XE-10.2-01; (b~d) —No.IGCAGS-XE-12.7-0101, thin section XE-12.7-01; (c, d) —magnified fragments shown by red and blue arrows in Fig.7b, respectively, showing the thin processes that successively bifurcate (or multifurcate)

  • 致谢:衷心感谢匿名审稿专家对本文提出的建设性修改意见。

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