黄烈斌, 刘一龙, 纵瑞文, 高丹霞, 申岑. (2024). 新疆东准噶尔地区早泥盆世软舌螺的发现及其古生态和古地理意义* [J]. 古地理学报, 26(2): 387-400.
HUANG Liebin, LIU Yilong, ZONG Ruiwen, GAO Danxia, SHEN Cen. (2024). Discovery of the Early Devonian hyoliths in East Junggar,Xinjiang and their palaeoecological and palaeogeographic significances[J]. Journal Of Palaeogeography, 26(2): 387-400.   
Doi: 10.7605/gdlxb.2024.02.023
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新疆东准噶尔地区早泥盆世软舌螺的发现及其古生态和古地理意义*
黄烈斌1, 刘一龙2, 纵瑞文3, 高丹霞1, 申岑1
1 昆明理工大学国土资源工程学院,云南昆明 650031
2 中国地质大学(武汉)地球科学学院,湖北武汉 430074
3 中国地质大学(武汉)生物地质与环境地质国家重点实验室,湖北武汉 430074
通讯作者简介 申岑,女,1985年生,副教授,主要从事寒武纪磷酸盐化化石研究。E-mail: cshen@kmust.edu.cn。

第一作者简介 黄烈斌,男,1998年生,硕士研究生,主要从事泥盆纪古生物学与地层学研究。E-mail: hliebin@163.com。

收稿日期:2023-08-01 修回日期:2023-09-22
基金资助:*云南省高层次人才培养支持计划“青年拔尖人才”专项和昆明理工大学分析测试基金(编号: 2022M20212101024)联合资助
摘要

本文首次系统报道了来自中国新疆东准噶尔地区早泥盆世地层中的软舌螺化石,共计 3属 3种,分别为: 王氏富蕴螺Fuyunotheca wangi gen. et sp. nov.,施氏纵肋螺Costulatotheca schleigeri Earp, 2019和奥氏螺未定种Ottomarites sp.。其中,王氏富蕴螺作为首次发现的新属种,典型特征为横切面呈扁椭圆形至亚三角形,背部膨胀,具有 1条贯穿壳体的背中脊,腹部较平至微微凸起,壳体前 1/3处具有横向成组的肋,整个壳体装饰着横向比纵向明显的细线。古生态学分析表明,王氏富蕴螺是一类生活在具有一定水动力的、富氧的正常浅海环境中的软舌螺。另外,笔者总结了包括新属种在内的目前所有报道过的整个泥盆纪产出的软舌螺的有效属种(不包括未定种和未定属),共计来自 11个国家 12个地区的 14属 66种,依据其古地理分布特征,认为泥盆纪的软舌螺主要分布在南半球的劳俄大陆和冈瓦纳大陆附近,其他地区仅零星出现。此次新疆东准噶尔地区软舌螺化石的发现, 不仅扩大了早泥盆世软舌螺类的古地理分布范围,同时为洲际间的地层对比提供了更多依据。

关键词: 软舌螺; 早泥盆世; 古生态; 古地理; 东准噶尔
中图分类号:Q911.5 文献标志码:A 文章编号:1671-1505(2024)02-0387-14
Discovery of the Early Devonian hyoliths in East Junggar,Xinjiang and their palaeoecological and palaeogeographic significances
HUANG Liebin1, LIU Yilong2, ZONG Ruiwen3, GAO Danxia1, SHEN Cen1
1 Faculty of Land Resources Engineering,Kunming University of Science and Technology,Kunming 650031,China
2 School of Earth Sciences,China University of Geosciences(Wuhan),Wuhan 430074,China
3 State Key Laboratory of Biogeology and Environmental Geology,China University of Geosciences(Wuhan),Wuhan 430074,China
About the corresponding author SHEN Cen,born in 1985,is an associate professor. She is engaged in the research of Cambrian phosphatized fossils. E-mail: cshen@kmust.edu.cn.

About the first author HUANG Liebin,born in 1998,is a master’s degree candidate. He is engaged in the study of Devonian palaeontology and stratigraphy. E-mail: hliebin@163.com.

Fund:Co-fund by the top-notch young person under the “Ten thousand Talents Program” in Yunnan Province and Kunming University of Science and Technology Analysis Test Fund(No.2022M20212101024)
Abstract

The newly discovered hyoliths yielded from the Lower Devonian of East Junggar of Xinjiang,China,are documented herein for the first time,including three genera and three species: Fuyunotheca wangi gen. et sp. nov., Costulatotheca schleigeriEarp,2019,and Ottomarites sp. As a new genus and species, F.wangi gen. et sp. nov. is mainly distinguished by typically oblate oval to subtriangular cross-section,swollen dorsal side with a pronounced median ridge,ventral side weakly convex. Surface sculpture prominent transverse striations and weakly longitudinal streaks on both dorsum and venter,and several sets of remarkable ribs parallel to the aperture in the anterior third of the conch. Palaeoecological analysis shows that F.wangi gen. et sp. nov. is a kind of orthothecids,which could live in a normal,oxygenated shallow-marine environment with certain degree of hydrodynamics. Furthermore,based on the Devonian hyoliths reported so far from all over the world,it has been recognized 14 genera and 66 species as valid from 12 areas in 11 countries(excluding indeterminate genera and species). Palaeogeographic distribution analysis shows that the Devonian hyoliths were mainly distributed in the southern hemisphere,that were positioned near the Laurussia and Gondwana,with a few species sporadically distributed in other plates or blocks. Newly discovered specimens of these early Devonian hyoliths not only expands the palaeogeographic distribution range of hyoliths during the Devonian period,but also provides new evidence for inter-continental biostratigraphic correlation.

Phylum uncertain

Class HyolithaMarek, 1963

Order OrthothecidaMarek, 1966

Family INDET.

Fuyunotheca gen. nov.

Type species Fuyunotheca wangi gen. et sp. nov.

Etymology Fuyun(Gr.)is derived from the Chinese Pinyin of the word “Fuyun” of Fuyun County,Altay Prefecture,Xinjiang. The theca(Gr.)means the capsule,membrane,sheath,and shell of organisms.

Diagnosis Orthothecids having straight and middle-sized conch with oblate oval to subtriangular cross-section;swollen dorsal side with a pronounced median ridge;ventral side flat to weakly convex. Aperture straight or almost straight. Apical part of shell blunt. No apical septa were found. Surface sculpture prominent transverse striations and weakly longitudinal streaks on both dorsum and venter,and several sets of remarkable ribs parallel to the aperture in anterior third of the conch near the apertural side. The separation distances between each set of transverse ribs gradually widen from the aperture to the apical end of the shell. Operculum is unknown.

Remarks The main difference between Hyolithida and Orthothecida is that the former has protrusions(lips)on the ventral side of aperture(Malinky and Racheboeuf,2010). The aperture of the new genus Fuyunotheca gen. nov. is straight and without protrusions,so it can be attributed to the order OrthothecidaMarek,1966. According to the taxonomy of OrthothecidaeSysoev,1958 revised byMalinky(2009b),its venter is concave to flat,and sculpture of conch consists of longitudinal ridges or lines,without transverse ornamentation. The new genus has the similar conch with that of the family Orthothecidae,but differs from them in having a flat to slightly convex venter and sets of unique transverse ribs on the conch. The new genus is similar to BolithecaMarek and Isaacson,1992 and NeobactrothecaMarek and Isaacson,1992 from the Middle Devonian Icla Formation of Bolivia. However,specimens of Bolitheca exhibit pronounced longitudinal ridges on the dorsum,the semi-elliptical cross-section,and rounded lateral ridges(Malinky and Racheboeuf,2011). Specimens of Neobactrotheca have many longitudinal ridges and no median ridges on the dorsum. Besides,the conch of Neobactrotheca differs from that of this new species by having elliptical cross-section and smooth transitions to the lateral ridges(Marek and Isaacson,1992). It is for this reason that we establish a new genus,belonging to the indeterminate family.

Fuyunotheca wangigen. et sp. nov.

Fig.2

Etymology The species is named in honor of Wang Hongzhen,the famous paleontologist who described hyoliths from China for the first time.

Holotypoe One well-preserved dorsum of conch,specimen number: XJLD-001(Fig.2-A).

Paratype One well-preserved venter of conch,specimen number: XJLD-003(Fig.2-C).

Other materials Specimen registration number: XJLD-002,004—046. Venter or dorsum of 44 conchs with incomplete aperture or initial part.

Diagnosis As for the genus.

Description The conch is straight and conical,expanding gradually and evenly from the apex to the aperture, and the aperture is thickened(Fig.2-E,ta). The growth angle of the conch is about 9.90°. The length of the shell is about 41 mm,and the diameter of the aperture is about 9 mm. The dorsum passes through the oblate lateral ridge to the venter,and its cross-section has oblate oval to subtriangular shape(Fig.2-B2). There is a median ridge running through the conch(Fig.2-A,B1,dmr)in the middle of dorsum. There are slightly inclined surfaces on both sides of the ridge,and a longitudinal groove on the right slope(Fig.2-A,gr). Sets of remarkable ribs(Fig.2-A,2-C,2-D,tr)parallel to the aperture are distributed in the anterior third of the conch. There are five groups of ridges(Fig.2-A,tr1-tr5)which are made up of 2-5 transverse ridges in each group. The separation distances between each set of transverse ribs gradually widen from the aperture to the apical end of the shell. Surface of shell from the middle to the initial part covered with slight growth lines(Fig.2-F,tl). In addition,the conch is also decorated with weakly longitudinal lines(Fig.2-G,ll),that are most prominent next to aperture but become fainter in direction of apex. But longitudinal lines become thicker near the lateral ridge(Fig.2-G,tll),which may be caused by pressure applied to the shell during diagenes. Ventral side almost flat to very slightly convex,and bears the same ornamentation as the dorsum without a median ridge(Fig.2-C,2-D,tr). The apex is damaged to some extent,but it still can be seen that it is bluntly rounded in form,and no septa are visible.

Locality and horizon The first member of the Tuoranggekuduke Formation,Emsian Age,Lower Devonian,Chawukar area,Fuyun County,Altay Prefecture,Xinjiang.

Key words: Hyoliths; Lower Devonian; Palaeoecology; Palaeogeography; East Junggar
1 概述

软舌螺是生活于寒武纪纽芬兰世幸运期中期至二叠纪末期的一类两侧对称的海生无脊椎动物(Wills, 1993; Malinky, 2009a; Valent et al., 2019), 其既是寒武纪最早出现的小壳生物群中的代表性生物, 同时也是寒武纪生命大爆发的见证者(李洛阳等, 2023)。寒武纪是软舌螺的鼎盛时期(Malinky and Racheboeuf, 2010), 或许是由于其他具有竞争力的底栖生物的崛起(如腹足类(Yochelson, 1984)和腕足类(Wang et al., 1987)), 造成该类群自奥陶纪之后便开始逐渐衰退, 直至二叠纪末全部灭绝(Malinky and Yochelson, 2007)。

自Eichwald(1840)首次报道软舌螺化石至今已有180余年, 但其亲缘关系一直存在争议。部分学者认为软舌螺是软体动物门下已经灭绝的一个纲(Marek and Yochelson, 1964, 1976; Malinky and Yochelson, 2007), 也有学者认为应该独立为软舌螺动物门(Runnegar, 1980; Pojeta, 1987), 而最近在加拿大不列颠哥伦比亚省布尔吉斯页岩(Burgess Shale)和美国犹他州斯潘塞页岩(Spence Shale)中发现的大量软舌螺动物的软体组织支持其应属于触手冠类(Moysiuk et al., 2017), 但也有学者认为它们应属于冠轮动物门的基础类群(Liu et al., 2020)。笔者遵循主流观点, 认为该类群属于已经灭绝的软舌螺纲, 门类未定。根据壳口是否有唇这一特征将软舌螺分为2个大类: 一类是生活于寒武纪纽芬兰世至泥盆纪的壳口平直且无唇的直管螺类(目), 另一类是生活于寒武纪第二世至二叠纪乐平世的壳口处有唇的软舌螺类(目)(Marek, 1963; Malinky, 1987, 2009b; Malinky and Racheboeuf, 2010)。研究表明, 直管螺类中的一支在寒武纪第二世早期演化出有唇的软舌螺类(Skovsted et al., 2016; Liu et al., 2022a, 2022b)。

软舌螺一般由始部封闭的锥形管壳(Conch)、内表面具有锁骨和主突起的口盖(Operculum)以及分布壳口两侧的1对附肢(Helens)组成, 其中附肢被认为是由口盖内表面的锁骨异速生长而来的(Skovsted et al., 2020)具有游泳功能的结构(Fisher, 1962; Marek, 1963)。但最初直管螺类口盖的内表面几乎没有复杂的结构(锁骨和主突起), 只有其雏形(Dzik, 1994), 到目前为止, 尚未发现附肢结构。除了在特异埋藏生物群中软舌螺的口盖和锥壳常可以组合保存外(甚至有时会保存比较好的软体组织)(e.g.Moysiuk et al., 2017; Liu et al., 2021), 一般很难发现口盖和锥壳组合保存的生物标本, 这一保存特征在寒武纪早期(纽芬兰世)的含磷岩系中最为突出(Li et al., 2018, 2020; Feng et al., 2022)。

通过统计地质历史时期软舌螺的数量分布, 刘璠(2021)认为地史上该类群主要存在9次程度不同的繁盛期, 其中寒武纪早中期的数量最多且研究最为详细透彻, 也是软舌螺繁盛和辐射的最大时期(e.g., Sun et al., 2015, 2018a, 2018b, 2018c; Moysiuk et al., 2017; Valent et al., 2019; Skovsted et al., 2020; Peel, 2021; Peel and Gubanov, 2022; Peel and Kouchinsky, 2022; Liu et al., 2022a, 2022b), 这可能与软舌螺的繁盛期和保存埋藏特征有关。除了寒武纪和奥陶纪外, 泥盆纪有关软舌螺类的研究则显得犹为薄弱, 且主要分布在劳俄大陆(Karpinsky, 1884; Williams, 1935; Sinclair, 1946; Ellison, 1965; Malinky, 1987; Malinky et al., 1987)、冈瓦纳大陆(Sinclair, 1946; Hiller, 1990; Marek and Isaacson, 1992; Malinky and Racheboeuf, 2011; Earp, 2019)和孤立的微小板块上, 如阿莫里凯板块(Armorica; Malinky and Racheboeuf, 2010)和波西米亚板块(Bohemia; Nová k, 1891; Barrande, 1867; Sinclair, 1946; Valent, 2006; Valent and Malinky, 2008)等。

中国拥有幅员辽阔的国土面积和相对较好的软舌螺类研究基础, 但软舌螺的相关研究主要集中在寒武纪地层中(钱逸, 1977, 1978; 毛家仁等, 1992; 钱逸等, 2000, 2003; 陈笑源等, 2003; Sun et al., 2018b; Skovsted et al., 2020; Liu et al., 2021, 2022a; 李洛阳等, 2023), 奥陶纪地层中虽然曾有学者提到过, 但一直未有详细的研究和报道(钱逸和肖立功, 1995), 后续的地质历史时期中关于软舌螺的报道几乎没有。文中报道的来自新疆东准噶尔地区早泥盆世的软舌螺化石是中国泥盆纪软舌螺类的首次报道, 此次发现不仅丰富了中国的软舌螺类研究材料, 同时也为泥盆纪软舌螺的形态学、古地理学和古生态学研究提供了新证据。

2 材料与方法

此次研究的软舌螺化石材料均采自新疆维吾尔自治区阿勒泰地区富蕴县西南部约50 km处的恰乌卡尔剖面(图1-A, E88° 57'37.50″, N46° 43'12.28″)。该剖面出露下泥盆统托让格库都克组, 在该地区该组被分为4个段: 第1段为灰色、深灰色钙质/泥质细砂岩、硅质粉砂状碎屑灰岩与晶屑岩屑凝灰岩、凝灰砂砾岩不均匀互层; 第2段为灰绿色、灰紫色安山玢岩、辉石安山玢岩, 夹凝灰砂岩、砂砾岩及钙质砂岩; 第3段主要为玄武玢岩、安山玢岩以及火山碎屑岩夹少量凝灰角砾岩; 第4段为安山玢岩、英安斑岩及凝灰砂岩夹凝灰岩(新疆地质局区域地质调查大队, 1979)。该组以上覆地层北塔山组的暗灰色凝灰圆砾岩为界, 两者呈不整合接触(新疆维吾尔自治区地质矿产调查局, 1999)。本次获得的化石产自该组第1段的第7层, 其岩性为灰黑色至深灰色钙质细砂岩(图 1-B), 风化面为灰黄色, 厚度约188.30 m。同一层位除了软舌螺化石外, 还发现有三叶虫(Odontochile tenuis, O. (kasazhstamia) intermedia, Odontochile sp., “ Phacops” sp., Acanthopyge sp.)、珊瑚、腕足类、双壳类、苔藓虫、腹足类和头足类等化石。本次获得含软舌螺化石共计145块, 锥壳205枚(编号为XJLD-001至XJLD-205), 除少部分化石保存完整外, 大部分化石均存在不同程度的残缺。使用气动笔对软舌螺化石标本进行清修, 莱卡显微镜(M205-C)进行图像采集, Adobe Photoshop CC 2019和CorelDraw X8软件进行排版和制图。所有标本均保存于昆明理工大学地学博物馆。

图 1 新疆阿勒泰地区富蕴县恰乌卡尔剖面地理位置(A)和下泥盆统托让格库都克组岩性柱状图(B)Fig.1 Location(A)and stratigraphic occurrence of the Lower Devonian Tuoranggekuduke Formation(B) of Qiawukaer section in Fuyun County, Altay Prefecture, Xinjiang

3 系统古生物学

门未定 Phylum uncertain

软舌螺纲 Class Hyolitha Marek, 1963

直管螺目 Order Orthothecida Marek, 1966

科未定 Family INDET.

富蕴螺属 Fuyunotheca gen. nov.

词源 Fuyun(Gr.)取产地新疆阿勒泰地区富蕴县的“ 富蕴” 二字汉语拼音, theca(Gr.)生物的囊、膜、鞘和壳。

模式种 王氏富蕴螺 Fuyunotheca wangi gen. et sp. nov.

属征 壳体直, 中等大小; 横切面呈扁椭圆形至亚三角形; 壳体背部膨胀凸起, 中间具1条贯穿壳体的中间脊, 腹部平至微微凸起; 壳口截切平直; 始部钝圆, 无隔板; 腹部与背部具有相同纹饰且整个壳体具横向纹饰和纵向纹饰, 其中横向纹饰比纵向纹饰清楚; 壳体前1/3部分处具成组横肋, 每组横肋之间的间隔由壳口向始部方向逐步变宽; 口盖未知。

讨论 软舌螺目与直管螺目的主要区别是前者壳口腹侧有突出物(唇)使得壳口不在一个平面上(Malinky and Racheboeuf, 2010), 而新属壳口平直且无突出物, 可归属于直管螺目。根据Malinky(2009b)修订的Orthothecidae Sysoev, 1958科征, 其腹部凹至平, 壳体上纹饰只有纵向的脊或线没有横向纹饰, 而新属腹部平至微凸, 且壳体表面装饰非常特殊的横向纹饰, 其他的特征与该科科征相似。新属与玻利维亚中泥盆统Icla组中的BolithecaMarek and Isaacson, 1992和NeobactrothecaMarek and Isaacson, 1992较为相似, 但又存在明显区别。Fuyunotheca gen. nov.与Bolitheca的区别在于后者背部具有明显的、较多的纵向脊, 横切面为半椭圆形, 侧脊圆等(Marek and Isaacson, 1992); 而Neobactrotheca的背部没有中脊, 代替的是许多纵向脊, 横切面呈椭圆形, 侧脊圆滑过渡(Marek and Isaacson, 1992)。综上所述, 笔者建立一新属, 但科未定。

王氏富蕴螺 Fuyunotheca wangi gen. et sp. nov.

图 2-A至2-G

图 2 新疆阿勒泰富蕴县恰乌卡尔地区下泥盆统托让格库都克组第1段中的Fuyunotheca wangi gen. et sp. nov.
A— 较完整的软舌螺壳体背部, 正模, 标本编号: XJLD-001; B— 壳口和始部缺失的背部, 标本编号: XJLD-002; C— 较完整的软舌螺壳体腹部, 副模, 标本编号: XJLD-003; D— 较完整的软舌螺壳体腹部, 标本编号: XJLD-004; E— 壳口纹饰特写; F-G— 正模局部放大。缩写解释: tr, transverse rib, 横向肋; dmr, dorsal median ridge, 背中脊; gr, groove, 凹槽; ll, longitudinal lines, 纵向线; tl, transverse lines, 横向线; tll, thicken longitudinal lines, 加粗纵向线; ta, thicken aperture, 壳口加粗; cs, cross-section, 横切面Fig.2 Fuyunotheca wangi gen. et sp. nov. from the First Member of Lower Devonian Tuoranggekuduke Formation in Qiawukaer area, Fuyun County, Altay Prefecture, Xinjiang

词源 种名为纪念中国第1个发现软舌螺化石的著名古生物学家王鸿祯院士。

正模 1个保存较好的壳体背部, 标本编号: XJLD-001(图 2-A)。

副模 1个保存较好的壳体腹部, 标本编号: XJLD-003(图 2-C)。

其他材料 44个壳口或始部残缺的壳体背部或腹部, 标本编号为: XJLD-002, 004— 046。

种征 见属征。

描述 壳体中等大小, 笔直呈锥形, 由壳顶向壳口处缓慢均匀扩大, 壳口处无唇, 壳口圈加粗(图 2-E, ta)。壳体生长角约9.90° , 壳体长约41 mm, 壳口直径约9 mm。壳体背部经扁圆的侧脊过渡至腹部, 其横切面为扁椭圆形至亚三角形(图 2-B2)。壳体背部中间有1条贯穿壳体的中间脊(图 2-A, B1, dmr), 脊两边存在稍微倾斜的面, 右侧斜面上可见1条纵向凹槽(图 2-A, gr), 凹槽自始部出现到中部截止。壳体背部前端约1/3部分内可见极其明显的横向肋(图 2-A, 2-C, 2-D, tr), 肋成组分布, 共有5组(图 2-A, tr1-tr5), 每组具2~5条横肋; 每组的间距向后逐渐变宽, 该区域内发育众多较清楚的横向线, 壳体中部至始部也存在比较淡的横向线(图 2-F, tl)。此外还存在许多极细的纵向线(图 2-G, ll), 纵向线在壳体近壳口处较清晰, 向壳顶逐渐变淡, 靠近侧脊处加粗(图 2-G, tll), 这也有可能是后期挤压造成的。壳体腹部平至微微突起, 没有中间脊, 其纹饰与背部相同(图 2-C, 2-D, tr)。壳顶部分有一定程度的损坏, 但仍见其呈钝圆形, 未见隔板。

比较 新种与其他早— 中泥盆世软舌螺均存在不同, 如背中脊特征, 具有背中脊的泥盆纪软舌螺物种有很多, 但是各不相同。Bolitheca crasquinaeMalinky and Racheboeuf, 2011的背中脊窄尖, 而新种较宽缓(图 2-A, dmr); Brittanella triliratus(Hall, 1879)的背中脊较狭窄, 且脊上有1条纵沟, 而新种比较单一。新种的纹饰最为独特, 成组的横肋仅在壳体前1/3处存在, 且整个壳体发育横向和纵向的细线, 这一特殊纹饰目前还未在早泥盆世的其他软舌螺属种中有所发现。综上所述, 新种以独特的背中脊、壳口处横向纹饰分2级(横肋和横向生长线)成组、始部无隔板等特征而成立。

产地与层位 新疆维吾尔自治区阿勒泰地区富蕴县恰乌卡尔地区, 托让格库都克组第1段, 埃姆斯阶, 下泥盆统。

纵肋螺属 Genus CostulatothecaEarp, 2019

模式种Costulatotheca schleigeriEarp, 2019

属征 锥壳中等大小, 横截面圆形, 在背部和腹部均有狭窄的纵脊; 除了不明显的适应性凸起的生长线外, 没有横向纹饰; 口盖内表面有2个大的主突起(Earp, 2019)。

施氏纵肋螺 Costulatotheca schleigeriEarp, 2019

图 3-A至3-D

图 3 新疆阿勒泰地区富蕴县恰乌卡尔地区下泥盆统托让格库都克组第1段中的软舌螺和共生化石
A-D— 为Costulatotheca schleigeriEarp, 2019; E— 为F壳体横向生长纹特写; F-G— 不完整的奥氏螺未定种Ottomarites sp., 登记号: XJLD-157(F), XJLD-158(G); H— 三叶虫 Odontochile sp.; I— 腕足类 Schellwienella sp.。缩写解释: to, transverse ornamentation, 横向纹饰; lr, longitudinal ribs, 纵肋; hcs, half cross-section, 半横截面Fig.3 Hyolithids and other fossils from the First Member of Lower Devonian Tuoranggekuduke Formation in Qiawukaer area, Fuyun County, Altay Prefecture, Xinjiang

材料 110个完整程度不一的锥壳, 标本编号为: XJLD-047至XJLD-156。

描述 壳体锥形, 中等大小, 壳口处平直, 横切面圆形, 但有些被压成透镜状, 标本大多数被剥蚀, 因而有些标本仅保存了壳体的一半, 可见其半横切面(图 3-B, 3-C, 3-D, hcs)。最大壳口直径为11 mm, 壳体长度3~40 mm, 生长角为10° ~40° 。壳体表面可见纵向的肋, 肋在壳顶处收缩, 壳体大的其纵肋比较宽(图 3-A, lr), 每5 mm约有7条, 壳体小的其纵肋小而密(图 3-C, lr), 每5 mm约有14条。口盖未知。

比较 可以与Bolitheca steinmanni和Neobactrotheca pharetra比较, 它们都有纵向的肋, 但是Bolitheca steinmanni和Neobactrotheca pharetra的肋比较尖锐, 而且肋之间的间隔比较宽, Costulatotheca schleigeri反之。Neobactrotheca pharetra的横切面为半圆形, 而Costulatotheca schleigeri则为圆形。与Earp(2019)所发现的标本相比较主要不同在于肋的密度, 本次发现的标本密度比较稀疏, 前者的比较密(12~13/mm), 其他特征相似。

产地与层位 新疆维吾尔自治区阿勒泰地区富蕴县恰乌卡尔地区, 托让格库都克组第1段, 埃姆斯阶, 下泥盆统。

软舌螺目 Order Hyolithida Sysoev, 1957

泡克西螺科 Family Pauxillitidae Marek, 1967

奥氏螺属 Genus Ottomarites Marek & Galle, 1976

奥氏螺未定种 Ottomarites sp.

图 3-E至3-G

材料 9个比较完整的印痕锥壳, 标本编号为: XJLD-157— XJLD-165。

描述 壳体长约18 mm, 壳口宽约6 mm, 生长角约15.85° 。壳体较直, 呈锥形, 壳口处具有向前突起的唇, 壳体自壳顶向壳口方向均匀扩大, 且壳体表面具有突起的细密横向纹饰(图 3-E, to), 每5 mm约有36条, 始部较尖, 口盖未知。XJLD-157(图 3-F)标本壳顶未完全出露, 壳体表面横向生长纹清楚。XJLD-158(图 3-G)标本受到后期挤压在壳体中间形成1条纵向的凸起脊, 具有横向纹饰且比XJLD-157(图 3-F)稍微模糊。

比较 由于标本保存不完整, 未保存口盖, 锥壳也有部分残缺。但突出的唇和具有清楚的、向壳口处弯的横向纹饰的壳体等特征符合OttomaritesMarek & Galle, 1976属征(Nová k, 1886; Valent and Malinky, 2008), 但由于没有更多的细节特征, 因此将其定义为未定种。该未定种壳体与Ottomarites discors(Barrande, 1867)的区别较小, 主要在于本文的未定种横向纹饰粗细统一、间隔均匀且壳体比较小, 而Ottomarites discors(Barrande, 1867)的壳体横向纹饰粗细不统一、间隔也不规律且壳体比较大, 因此无法归属于该种。

产地与层位 新疆维吾尔自治区阿勒泰地区富蕴县恰乌卡尔地区, 托让格库都克组第1段, 埃姆斯阶, 下泥盆统。

4 古生态与生物古地理
4.1 古生态

除了高盐度、白云质的水体和生物礁相的沉积环境外, 几乎在所有类型的海相沉积环境中软舌螺类都可以生存(Fisher, 1962), 它们经常与腹足类、海绵类、腕足类以及三叶虫等共同生活于正常盐度的浅海环境中(Hiller, 1990)。同时苔藓虫、珊瑚和海百合也可能会寄生在软舌螺壳体的背部, 利用流经壳体背部的水流来获得更多的营养物质(Marek and Galle, 1976; Galle and Parsley, 2005)。在浅海且富氧的环境中, 软舌螺的丰度和多样性均相对较高, 其个体也相对比较大, 如捷克的巴朗德地区(Barrande, 1867; Nová k, 1891; Sinclair, 1946, Malinky, 2009b)。反之在深水、缺氧的暗色泥页岩或硅质岩中则很少有软舌螺被发现, 如德国中西部地区(Malinky and Lichtenberger, 2005)。

大多数的直管螺类腹部均存在1个纵向的沟, Malinky(2009b)曾认为软舌螺腹部纵沟的深度与其生活的基底软硬程度有密切关系, 基底越硬腹部纵沟越浅, 反之腹部纵沟越深, 这可能是软舌螺为适应生活环境而做出的身体结构上的变化。因此生活在海洋基底为砂质或含砂质的软舌螺, 其腹部呈现出较平至轻微凸起的变化, 以来自日本的Joachimilites fukujiensis Kase, 1987为典型代表(Kase et al., 1987); 海洋基底为黏土的或泥质的软舌螺, 腹部呈现出平至凹的变化, 以南非的Hyolithes? subaequalis (Salter, 1845)为典型代表(Hiller, 1990)。王氏富蕴螺(Fuyunotheca wangi gen. et sp. nov.)的壳体较大, 壳体背部有一突起的中间脊, 腹部较平至微凸, 因此可以判断该属可能生活于较硬的沉积物表面。同时依据软舌螺常与分散的三叶虫背壳、腕足类和苔藓虫化石碎片共同保存, 软舌螺化石均未发现口盖和岩性以灰黄色中厚层钙质细砂岩等特征, 推测王氏富蕴螺生活在具有一定水动力的浅海环境中, 这样的条件能为它们带来更多的营养物质或者是借助水动力进行缓慢的移动, 这或许与它的滤食性生活有关, 而不仅仅底栖掘泥生活。Costulatotheca schleigeri与F.wangi gen. et sp. nov. 生活在相同环境之中, 其个体最大的达到了40 mm以上, 可能是由于浅海中的水压力小, 氧气丰富, 营养物质充足, 为它们的发育提供了客观条件。Earp(2019)所发现的标本是产出于复理石中, 其埋藏环境为缺氧条件的低能海底, 根据保存于同层位的腕足类与双壳类推测软舌螺是经历了一定的漂流, 因此并不能代表Costulatotheca schleigeri的原始古生态环境。综上所述, 本次发现的软舌螺类可能与三叶虫和腕足类共同生活于正常浅海、富氧、营养物质丰富且具有一定水动力的环境中。

4.2 生物古地理分布

依据目前收集到的资料, 笔者共统计了来自11个国家12个地区所报道的泥盆纪软舌螺化石, 共计14个属66个种(附表1), 并对部分国家代表性种的特征进行了总结对比(表 1)。古地理投图显示泥盆纪软舌螺主要分布在中亚造山带(中国新疆=1, 本文)、劳俄大陆(俄罗斯乌拉尔地区=2, 加拿大魁北克省=3, 美国纽约州=4, 美国宾夕法尼亚州=5, Karpinsky, 1884; Williams, 1935; Sinclair, 1946; Ellison, 1965; Malinky, 1987; Malinky et al., 1987)、冈瓦纳大陆(南非南部=6, 玻利维亚西部=7, 澳大利亚维多利亚州=12, Sinclair, 1946; Hiller, 1990; Marek and Isaacson, 1992; Malinky and Racheboeuf, 2011; Earp, 2019)、阿莫里凯板块(Armorica, 法国西北部=8, Malinky and Racheboeuf, 2010)、莱茵地块(德国西南部=9, Sinclair, 1946; Houbrick et al., 1988)、波西米亚板块(Bohemia, 捷克=10, Nová k, 1891; Barrande, 1867; Sinclair, 1946; Valent, 2006; Valent and Malinky, 2008)、华南板块(日本中部=11, Kase et al., 1987)(图 4)。

表 1 部分国家泥盆纪软舌螺代表种的特征 Table1 Typical species characteristics of the Devonian hyoliths in some countries

图 4 泥盆纪软舌螺古地理分布及各国属种统计(资料来源于Scotese, 2016; Hiller, 1990; Marek and Isaacson, 1992; Sinclair, 1946; Malinky, 1987; Malinky et al., 1987; Houbrick et al., 1988; Valent, 2006; Valent and Malinky, 2008; Malinky, 2009b; Malinky and Racheboeuf, 2010, 2011; 有修改)Fig.4 Palaeogeographic distribution of the Devonian hyoliths and genera and species statistics of every country(histogram)(modified from Scotese, 2016; Hiller, 1990; Marek and Isaacson, 1992; Sinclair, 1946; Malinky, 1987; Malinky et al., 1987; Houbrick et al., 1988; Valent, 2006; Valent and Malinky, 2008; Malinky, 2009b; Malinky and Racheboeuf, 2010, 2011)

中国钱逸和肖立功(1995)在出版的《软舌螺》专著中统计了中国的软舌螺, 曾提到“ 早寒武世有36属79种, 中寒武世22属41种, 晚寒武世4属6种, 奥陶纪2属2种” , 其中并没有泥盆纪的软舌螺记载, 后续的几十年研究中也未见到相关中国泥盆纪软舌螺的报道。本文报道的来自中国新疆东准噶尔下泥盆统埃姆斯阶的软舌螺化石是中国泥盆纪软舌螺的首次正式报道, 共描述3属3种, 包括1新属新种和1未定种。

俄罗斯 俄罗斯记录的软舌螺Hyolithes uralicusKarpinsky, 1884由俄罗斯著名的地质学家Karpinsky(1884)报道, 该物种也是迄今为止俄罗斯唯一报道的泥盆纪软舌螺化石。

加拿大 1892年, Whitevaves在研究泥盆系地层产出的化石组成时曾经记录过软舌螺化石, 但Hyolithes alatus Whiteaves, 1892目前已被归属于腹足纲(Rocque, 1949), 因而不进行统计。确切的加拿大软舌螺物种仅有2属2种, 且均来自魁北克省(Sinclair, 1946)。

美国 美国目前所报道的泥盆纪软舌螺共有5属12种, 大部分分布于美国北部地区。在此之前, 虽然已有零星的软舌螺化石记录(Hall, 1876, 1879, 1888; Williams, 1935; Ellison, 1965), 但是直到很多年之后, 其中的泥盆纪软舌螺化石才被Malinky(1987)和Malinky等(1987)整理和纠正。

南非Hiller(1990)报道了来自南非南部的泥盆系Voorstehoek组中的软舌螺Hyolithes capensis Reed, 1925和 H. subaequalis(Salter), 1845, 共计1属2种。

玻利维亚Ulrich(1892)首次报道了玻利维亚的泥盆纪软舌螺化石, 随后Knod(1908)描述了一新种Orthotheca steinmanniKnod, 1908, 后来Marek和Isaacson(1992)认为该物种有着不同于Orthotheca属的特点而建立了一新属Neobactrotheca。因此, 目前玻利维亚共报道泥盆纪软舌螺化石4属5种, 主要分布在4个层位: 第1处是玻利维亚中部Icla地区Icla组的中段上部, 主要产出Bolitheca steinmanni (Knod, 1908)和Bolithes crasquinaeMalinky and Racheboeuf, 2011; 第2处是玻利维亚西北地区Belé n组的上部, 主要产出Neobactrotheca pharetraMarek & Isaacson, 1992; 第3处是玻利维亚南部Gamoneda组的底部; 第4处是玻利维亚Presto地区Huamamoampa组的下部产出了一些软舌螺未定属种(Marek and Isaacson, 1992)。

法国Malinky和Racheboeuf(2010)首次报道了法国西北部中泥盆统Kersadiou组中的软舌螺化石, 并将Hyolithes triliratusHall, 1879修订为Brittanella triliratus (Hall, 1879)。这一属也曾在美国东部发现过(Malinky et al., 1987)。

德国 目前在德国所报道的泥盆纪软舌螺共计2属13种, 最早被发现的是Hyolithes anceps (Mü nster), 1840(Sinclair, 1946)。随后在德国陆续报道了其他物种(Ludwing, 1864; Mauz, 1933; Sinclair, 1946), 最新记录的是Houbrick等(1988)曾经在研究德国南部下泥盆统中产出的软体动物群时发现的一软舌螺化石, 但由于标本保存原因仅进行了图版展示, 并未对其进行详细鉴定和描述。

捷克 捷克是目前报道泥盆纪软舌螺最多的国家, 共计5属32种(Barrande, 1867; Nová k, 1891; Sinclair, 1946; Valent, 2006; Valent and Malinky, 2008)。Barrande(1867)曾统计了12个物种, Nová k(1891)共记录10个物种, 笔者从其他文献中又整理了10个物种, 这些属种大多数都来自捷克巴兰德和波西米亚地区的下— 中泥盆统。

日本 仅报道了泥盆纪软舌螺化石1属1种— — Joachimilites fukujiensis Kase, 1987。该种来自日本中部地区下或中泥盆统Fukuji组中, 其岩性为黑色钙质含细砂泥页岩。

澳大利亚Earp(2019)首次报道了澳大利亚维多利亚州的泥盆纪直管螺类化石, 并建立了一新属种Costulatotheca schleigeriEarp, 2019。该种在本次研究中也有发现。

5 结论

1)文中报道了来自新疆东准噶尔早泥盆世埃姆斯期的软舌螺化石3属3种, 包括1新属新种— — 王氏富蕴螺(Fuyunotheca wangi gen. et sp. nov.), 其典型特征是横切面呈扁椭圆形至亚三角形, 背部膨胀具有1条中脊, 壳体前1/3部分具有横向成组的脊, 整个壳体具有横向和纵向的细线, 腹部较平至微微凸起且纹饰与背面相同。

2)统计了到目前为止所报道过的全球泥盆纪软舌螺化石属种(不包括未定种和未定属), 并分析了泥盆纪时期软舌螺的古地理位置, 发现它们大多数都存在于南半球的劳俄大陆和冈瓦纳大陆附近, 其他地区仅零星分布。

3)古生态学研究表明王氏富蕴螺可能与三叶虫和腕足类共同生活于正常浅海、富氧、营养物质丰富且具有一定水动力的环境中, 推测其为营滤食生活。

4)此次研究是中国泥盆纪地层中软舌螺化石的首次报道, 不仅丰富了中国新疆泥盆纪地层中古生物的多样性, 同时为泥盆纪时期各板块之间的地层对比和生物交流提供了更多的依据。

致谢 感谢中国科学院南京地质古生物研究所宋俊俊副研究员和硕士生孙语聪提供部分论文资料, 感谢中国科学院南京地质古生物研究所孙海静副研究员和西北大学地质系刘璠博士解答部分专业问题, 感谢中国地质大学(武汉)地球科学学院副教授范若颖给予英文修改, 博士生郭超和尹家一及硕士生杜晓琦、马娟和刘士博协助野外工作。感谢两位匿名审稿人给予本文诸多建设性意见, 使本文得到质的提升。

(责任编辑 张西娟)

参考文献
[1] 陈笑媛, 赵元龙, 王萍丽. 2003. 贵州凯里生物群中软舌螺的初步研究. 微体古生物学报, 20(3): 296-302.
[Chen X Y, Zhao Y L, Wang P L. 2003. Preliminary research on hyolithids from the Kaili biota, Guizhou. Acta Micropalaeontologica Sinica, 20(3): 296-302] [文内引用:1]
[2] 李洛阳, 潘兵, 贠浩. 2023. 华北寒武系第二统辛集组软舌螺动物化石群. 古生物学报, 62(1): 30-65.
[Li L Y, Pan B, Yun H. 2023. Hyolith fauna from the Xinji Formation(Cambrian Series 2)in North China. Acta Palaeontologica Sinica, 62(1): 30-65] [文内引用:2]
[3] 刘璠. 2021. 滇东地区寒武系第二统软舌螺动物研究. 西北大学博士学位论文: 134-143.
[Liu F. 2021. Palaeobiological study of hyoliths from the Cambrian Series 2 Lagerstätten in eastern Yunnan, China. Doctoral dissertation of Northwest University: 134-143] [文内引用:3]
[4] 毛家仁, 赵元龙, 余平, 钱逸. 1992. 贵州台江中寒武世的一些软舌螺化石. 微体古生物学报, 9(3): 257-265.
[Mao J R, Zhao Y L, Yu P, Qian Y. 1992. Some Middle Cambrian hyolithids from Taijiang, Guizhou. Acta Micropalaeontologica Sinica, 9(3): 257-265] [文内引用:1]
[5] 钱逸. 1977. 华中西南区早寒武世梅树村阶的软舌螺纲及其他化石. 古生物学报, 16(2): 255-281.
[Qian Y. 1977. Hyolitha and some problematica from the Lower Cambrian Meishucun Stage in central and S. W. China. Acta Palaeontolgica Sinica, 16(2): 255-281] [文内引用:1]
[6] 钱逸. 1978. 华中西南区早寒武世软舌螺化石的研究及其地层意义. 中国科学院南京地质古生物研究所集刊, 11: 1-50.
[Qian Y. 1978. The Early Cambrian hyolithids in central and Southwest China and their stratigraphical significance. Journal of Nanjing Institute of Geology and Paleontology, Chinese Academy of Sciences, 11: 1-50] [文内引用:1]
[7] 钱逸, 肖立功. 1995. 软舌螺. 北京: 科学出版社, 65-79.
[Qian Y, Xiao L G. 1995. Hyolitha. Beijing: Science Press, 65-79] [文内引用:2]
[8] 钱逸, 尹恭正, 肖兵. 2000. 新疆下寒武统玉尔吐斯组软舌螺口盖和口盖状化石. 微体古生物学报, 17(4): 404-415.
[Qian Y, Yin G Z, Xiao B. 2000. Opercula of hyoliths and operculum-like fossils from the Lower Cambrian Yuertus Formation, Xinjiang. Acta Micropalaeontologica Sinica, 17(4): 404-415] [文内引用:1]
[9] 钱逸, 李国祥, 朱茂炎, 尹恭正. 2003. 新疆乌什下寒武统肖尔布拉克组软舌螺化石. 微体古生物学报, 20(4): 342-349.
[Qian Y, Li G X, Zhu M Y, Yin G Z. 2003. Hyoliths from the Lower Cambrian Xiaoerbulakr Formation of Wushi, Xinjiang. Acta Micropalaeontologica Sinica, 20(4): 342-349] [文内引用:1]
[10] 新疆地质局区域地质调查大队. 1978. 区域地质调查报告, 1︰200000 地质图, 锡泊渡幅(L-45-XI).
[Regional Geological Survey Brigade of Xinjiang Geological Bureau. 1978. Regional geological survey report, Geological Map 1︰200000, Xibodufu, Sheet(L-45-XI)] [文内引用:1]
[11] 新疆维吾尔自治区地质矿产调查局. 1999. 新疆维吾尔自治区岩石地层. 武汉: 中国地质大学出版社, 1-430.
[XBGMR(Xinjiang Bureau of Geology and Mineral Resources). 1999. The Xinjiang Region Rock Strata. Wuhan: China University of Geosciences Press, 1-430] [文内引用:1]
[12] Dzik J. 1994. Evolution of ‘Small Shelly Fossils’ assemblages of the Early Paleozoic. Acta Palaeontologica Polonica, 39(3): 247-313. [文内引用:1]
[13] Earp C. 2019. Costulatotheca schleigeri(Hyolitha: Orthothecida)from the Walhalla Group(Early Devonian)at Mount Pleasant, central Victoria, Australia. Alcheringa: An Australasian Journal of Palaeontology, 43(2): 220-227. [文内引用:10]
[14] Ellison R L. 1965. Stratigraphy and paleontology of the Mahantango Formation in south-central Pennsylvania. Pennsylvania Geological Survey Bulletin, 48: 298. [文内引用:3]
[15] Feng Q, Pan B, Yang A H, Miao L, Li G X. 2022. Biostratigraphy of the Small Shelly Fossils from the Upper Maidiping Formation(Terreneuvian)at the Fand ian Section, Sichuan Province, South China. Frontiers Earth Science: Section Paleontology, 10: 922439. [文内引用:1]
[16] Fisher D W. 1962. Small conoidal shells of uncertain affinities. In: Moore R C(eds). Treatise on Invertebrate Paleontology, part W. Lawrence, Kansas: Geological Society of America and University of Kansan Press, 98-143. [文内引用:2]
[17] Galle A, Parsley R L. 2005. Epibiont relationships on hyolithids demonstrated by Ordovician trepostomes(Bryozoa)and Devonian tabulates(Anthozoa). Bulletin of Geosciences, 80(2): 125-138. [文内引用:1]
[18] Hall J. 1876. Illustrations of Devonian fossils: Gasteropoda, Pteropoda, Cephalopoda, Crustacea and Corals of the Upper Helderberg, Hamilton and Chemung Groups. Albany: Weed, Parsons and Company: 1-39. [文内引用:1]
[19] Hall J. 1879. Palaeontology of New York: Gasteropoda, Pteropoda and Cephalopoda of the Upper Helderberg, Hamilton, Portage and Chemung Groups. Geological Survey of the State of New York, 5(2): 1-492. [文内引用:4]
[20] Hall J. 1888. Palaeontology of New York: supplement containing descriptions and illustrations of Pteropoda, Cephalopoda and Annelida. Geological Survey of the State of New York, 7: 1-42. [文内引用:1]
[21] Hiller N. 1990. Devonian hyoliths in South Africa, and their palaeoenvironmental significance. Palaeontologica Africana, 27: 5-8. [文内引用:5]
[22] Houbrick R S, Stürmer W, Yochelson E L. 1988. Rare Mollusca from the Lower Devonian Hunsriick Slate of southern Germany. Lethaia, 21(4): 395-402. [文内引用:2]
[23] Kase T, Hamada T, Niko S. 1987. First record of a hyolith(Paleozoic Mollusca)from Japan. Bulletin of the National Museum of Tokyo, 13(1): 29-34. [文内引用:2]
[24] Li L Y, Zhang X L, Skovsted C B, Yuan H, Pan B, Li G X. 2019. Homologous shell microstructures in Cambrian hyoliths and molluscs. Palaeontology, 62(4): 515-532. [文内引用:1]
[25] Li L Y, Skovsted C B, Yun H, Betts M J, Zhang X L. 2020. New insight into the soft anatomy and shell microstructures of early Cambrian orthothecids(Hyolitha). Proceedings of the Royal Society B: Biological Sciences, 287: 20201467. [文内引用:1]
[26] Liu F, Skovsted C B, Topper T P, Zhang Z F, Shu D G. 2020. Are hyoliths Palaeozoic lophophorates? National Science Review, 7(2): 453-469. [文内引用:1]
[27] Liu F, Skovsted C B, Topper T P, Zhang Z F. 2021. Soft part preservation in hyolithids from the lower Cambrian(Stage 4)Guanshan Biota of South China and its implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 562: 110079. [文内引用:1]
[28] Liu F, Skovsted C B, Topper T P, Zhang Z F. 2022a. Hyolithid-like hyoliths without helens from the early Cambrian of South China, and their implications for the evolution of hyoliths. BMC Ecology and Evolution, 22: 64. [文内引用:3]
[29] Liu F, Skovsted C B, Topper T P, Zhang Z F. 2022b. A fresh look at the Hyolithid Doliutheca from the Early Cambrian(Stage 4)Shipai Formation of the Three Gorges Area, Hubei, South China. Biology, 11(6): 875. [文内引用:2]
[30] Malinky J M. 1987. Taxonomic revision of Lower and Middle Paleozoic Orthothecida(Hyolitha)from North America and China. Journal of Paleontology, 61(5): 942-959. [文内引用:8]
[31] Malinky J M. 2009a. Permian Hyolithida from Australia: the last of the Hyoliths?Journal of Paleontology, 83(1): 147-152. [文内引用:1]
[32] Malinky J M. 2009b. First occurrence of orthotheca Novák, 1886(Hyolitha, Early Devonian)in North America. Journal of Paleontology, 83(4): 588-596. [文内引用:4]
[33] Malinky J M, Linsley R M, Yochelson E L. 1987. Taxonomic revision of hyolitha from the Middle Paleozoic of North America. Journal of Paleontology, 61(6): 1173-1186. [文内引用:1]
[34] Malinky J M, Lichtenberger M. 2005. Hyolitha and problematica from the Rhine Slate Mountains(Lower-Middle Devonian), and their palaeobiological significance. Neues Jahrbuch fur Geologie und Palaontologie-Abhand lungen, 238(1): 79-106. [文内引用:1]
[35] Malinky J M, Yochelson E L. 2007. On the systematic position of the Hyolitha(Kingdom Animalia). Memoir of the Association of Australasian Palaeontologists, 34: 521-536. [文内引用:2]
[36] Malinky J M, Racheboeuf P R. 2010. Devonian(Givetian)Hyolitha from the Kersadiou Formation, Massif Amoricain, France. Journal of Paleontology, 84(4): 739-745. [文内引用:6]
[37] Malinky J M, Racheboeuf P R. 2011. New Hyolitha from the Devonian of Bolivia. Journal of Paleontology, 85(6): 1077-1088. [文内引用:4]
[38] Marek L. 1963. New knowledge on the morphology of Hyolithes. Sborník Geologických Věd, Paleontologie, 1: 53-73. [文内引用:3]
[39] Marek L. 1966. New hyolithid genera from the Ordovician of Bohemia. Casopis Národního Muzea, 135(2): 89-92. [文内引用:1]
[40] Marek L. 1967. The class Hyolitha in the Caradoc of Bohemia. Sbornik Geologickych Ved, Rada Paleontologie, 9: 51-113. [文内引用:1]
[41] Marek L, Yochelson E L. 1964. Paleozoic Mollusk: Hyolithes. Science, 146(3652): 1674-1675. [文内引用:1]
[42] Marek L, Galle A. 1976. The tabulate coral Hyostragulum, an epizoan with bearing on hyolithid ecology and systematics. Lethaia, 9(1): 51-64. [文内引用:1]
[43] Marek L, Yochelson E L. 1976. Aspects of the biology of Hyolitha(Mollusca). Lethaia, 9: 65-82. [文内引用:3]
[44] Marek L, Isaacson P E. 1992. Devonian Hyoliths from Bolivia. Journal of Paleontology, 66(1): 24-28. [文内引用:9]
[45] Moysiuk J, Smith M R, Caron J B. 2017. Hyoliths are Paleozoic lophophorates. Nature, 541: 394-397. [文内引用:3]
[46] Peel J S. 2021. In-place operculum demonstrates that the Middle Cambrian Protowenella is a hyolith and not a mollusc. Alcheringa: An Australasian Journal of Palaeontology, 45(4): 385-394. [文内引用:1]
[47] Peel J S, Gubanov A P. 2022. Hyoliths from the Bystraya Formation(Cambrian Series 2)of eastern Transbaikalia(Zabaykalsky Krai), Siberia. Alcheringa: An Australasian Journal of Palaeontology, 46(2): 129-146. [文内引用:1]
[48] Peel J S, Kouchinsky A. 2022. Middle Cambrian(Miaolingian Series, Wuliuan Stage)molluscs and mollusc-like microfossils from North Greenland (Laurentia). Bulletin of the Geological Society of Denmark, 70: 69-104. [文内引用:1]
[49] Pojeta J J. 1987. Class Pelecypoda. In: Boardman R S, Cheetham A H, Rowell A J(eds). Fossil Invertebrates. Palo Alto: Blackwell Scientific Publications, 386-435. [文内引用:1]
[50] Rocque A L. 1949. New Uncoiled Gastropods from the Middle Devonian of Michigan and Manitoba. University of Michigan, 7(7): 113-122. [文内引用:1]
[51] Runnegar B. 1980. Hyolitha: status of the phylum. Lethaia, 13(1): 21-25. [文内引用:1]
[52] Scotese C R. 2016. PALEOMAP paleoAtlas for gplates. Https://www.earthbyte.org/paleomap-paleoatlas-for-gplates. [文内引用:1]
[53] Sinclair G W. 1946. Notes on the Nomenclature of Hyolithes. Journal of Paleontology, 20(1): 72-85. [文内引用:12]
[54] Skovsted C B, Pan B, Topper T P, Betts M J, Li G X, Brock G A. 2016. The operculum and mode of life of the lower Cambrian hyolith Cupitheca from South Australia and North China. Palaeogeography, Palaeoclimatology, Palaeoecology, 443: 123-130. [文内引用:1]
[55] Skovsted C B, Mus M M. Zhang Z L, Pan B, Li L Y, Liu F, Li G X, Zhang Z F. 2020. On the origin of hyolith helens. Palaeogeography, Palaeoclimatology, Palaeoecology, 555: 109848. [文内引用:3]
[56] Sun H J, Babcock L E, Peng J, Zhao Y L. 2015. Hyolithids and associated trace fossils from the Balang Formation(Cambrian Stage 4), Guizhou, China. Palaeoworld, 24(1-2): 55-60. [文内引用:1]
[57] Sun H J, Malinky J M, Zhu M Y, Huang D Y. 2018a. Palaeobiology of orthothecide hyoliths from the Cambrian Manto Formation of Hebei Province, North China. Acta Palaeontologica Polonica, 63(1): 87-101. [文内引用:1]
[58] Sun H J, Smith M R, Zeng H, Zhao F C, Li G X, Zhu M Y. 2018b. Hyoliths with pedicles illuminate the origin of the brachiopod body plan. Proceedings of the Royal Society B: Biological Sciences, 285(1887): 20181780. [文内引用:2]
[59] Sun H J, Zhao F C, Wen R Q, Zeng H, Peng J. 2018c. Feeding strategy and locomotion of Cambrian hyolithides. Palaeoworld, 27(3): 334-342. [文内引用:1]
[60] Sysoev V A. 1957. To the morphology, systematics and systematic position of the Hyolithoidea. Doklady Akademiya Nauk SSSR, 116(2): 304-307. [文内引用:1]
[61] Sysoev V A. 1958. The superorder Hyolithoidea. In: Luppov N P, Drushchits V V(eds). Principles of Palaeontology, Mollusca-Cephalopoda. Russia: Akademii Nauk SSSR, 184-190. [文内引用:1]
[62] Valent M. 2006. Stratigraphic distribution of the class Hyolitha(Mollusca)in the Barrand ian area(Czech Republic). Acta Universitatis Carolinae-Geologica, 47(1-4): 183-188. [文内引用:3]
[63] Valent M, Malinky J M. 2008. Early Devonian(Emsian)Hyolith Ottomarites discors(Barrand e, 1867)with colour pattern. Bulletin of Geosciences, 83(4): 503-506. [文内引用:4]
[64] Valent M, Fatka O, Marek L. 2019. Alfaites romeo gen. et sp. nov. ,a new Hyolitha from the Cambrian of Skryje-Týřovice Basin(Czech Republic). European Journal of Taxonomy, 491: 1-10. [文内引用:2]
[65] Wang Y, Boucot A J, Rong J Y, Yang X C. 1987. Community Paleoecology as a geologic tool: the Chinese Ashgillian-Eifelian(latest Ordovician through early Middle Devonian)as an example. Geological Society of America, Special Paper, 211: 1-101. [文内引用:1]
[66] Whiteaves J F. 1892. The fossils of the Devonian rocks of the island s, shores or immediate vicinity of Lakes Manitoba and Winnipegosis. Contributions to Canadian Palaeontology, 1(4): 255-359. [文内引用:1]
[67] Williams J S. 1935. Paleontology. In: Cooper G A, Williams J S(eds). Tully Formation of New York. Colorado: Bulletin of the Geological Society of America, 830-859. [文内引用:3]
[68] Wills M A. 1993. ?Mollusca Incertae Sedis. In: Benton M J. The Fossil Record 2. Cambridge: The University Press, 265-270. [文内引用:1]
[69] Yochelson E L. 1984. Speculative functional morphology and morphology that could not function: the example of Hyolithes and Biconulites. Malacologia, 25: 255-264. [文内引用:1]
[70] Barrand e J. 1867. Systême Silurien du Centre de la Bohême. Ordre des Ptéropodes. Privately publish, Prague & Paris, 3: 1-179. [文内引用:7]
[71] Eichwald K E V. 1840. Über das silurische Schichtensystem in Esthland . Zeitschrift fur Natur-und Heilkunde der Königlichen Medicinisch-chirurgischen Akademie St. Petersburg, 1/2: 1-210. [文内引用:1]
[72] Karpinsky A. 1884. Die fossilen Pteropoden an Ostabhange des Urals. Academy Imperial Science Saint Petersburg, Memoire, 7(32): 1-21. [文内引用:4]
[73] Knod R. 1908. Devonische Faunen Boliviens. In: Steinmann G(eds). Beiträge zur Geologie und Paläontologie von Südamerika. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 25: 493-600. [文内引用:3]
[74] Ludwing R. 1864. Pteropoden aus dem Devon in Hessen und Nassau sowie aus dem Tertiaer-Thon des Mainzer beckens. Paleontographica, 11: 311-322. [文内引用:1]
[75] Mauz J. 1933. Zur Fauna der Unterkoblenz Stufe. Senckenbergiana, 15(3/4): 274-294. [文内引用:1]
[76] Novák O. 1886. Zur Kenntniss der Fauna der Etage F-fl in der palaeozischen Schichtengruppe Böhmens. Aus den Sitzungsberichten der königlichen böhmens Gesellschaft der Wissenschaften, 1-27. [文内引用:1]
[77] Novák O. 1891. Revision der palaeozoischen Hyolithiden Böhmens. Abhand lungen der mathematisch-naturwischenschaftlichen Classe der böhmischen Gesellschaft der Wissenschaften, 7(4): 1-48. [文内引用:5]
[78] Ulrich A. 1892. Palaeozoische Versteinerungen aus Bolivien. In: Steinmann G(ed). Beitrage zur Geologie und Palaontologie von Sudamerika. Germany: Neues Jahrbuch für Geologie und Paläontologie, 1-116. [文内引用:1]