华南寒武纪碳酸盐岩中最早腕足动物的辐射、发育与分布

doi:10.13745/j.esf.sf.2020.6.4

地学前缘 ›› 2020, Vol. 27 ›› Issue (6): 79-103.DOI: 10.13745/j.esf.sf.2020.6.4

华南寒武纪碳酸盐岩中最早腕足动物的辐射、发育与分布

张志亮¹^,²(), 陈飞扬¹^,², 张志飞¹^,^*()

1.西北大学地质学系大陆动力学国家重点实验室与早期生命与环境陕西省重点实验室, 陕西西安 710069
2.麦考瑞大学生命科学系, 澳大利亚悉尼 NSW 2109

收稿日期:2020-03-23 修回日期:2020-05-27 出版日期:2020-11-02 发布日期:2020-11-02
通讯作者: 张志飞
作者简介:张志亮(1988—),男,博士,主要从事寒武纪早期壳体化石形态、结构与发育研究。E-mail:zhiliang. zhang@mq.edu.au
基金资助:
国家自然科学基金项目(41425008);国家自然科学基金项目(41720104002);国家自然科学基金项目(41621003);国家自然科学基金项目(41890844);中国科学院战略性先导科技专项(B类)项目(XDB26000000);高等学校学科创新引智计划111项目(D17013);国家留学基金委公派研究生项目(CSC 201806970026);澳大利亚麦考瑞大学研究奖学金项目(Macquarie University Research Fellowships)(2019 MQRF)

The earliest phosphatic-shelled brachiopods from the carbonates of South China: their diversification, ontogeny and distribution

ZHANG Zhiliang¹^,²(), CHEN Feiyang¹^,², ZHANG Zhifei¹^,^*()

1. State Key Laboratory of Continental Dynamics and Shaanxi Key Laboratory of Early Life and Environments, Department of Geology, Northwest University, Xi’an 710069, China
2. Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia

Received:2020-03-23 Revised:2020-05-27 Online:2020-11-02 Published:2020-11-02
Contact: ZHANG Zhifei

摘要/Abstract

摘要：

腕足动物是寒武纪冠轮动物分支的重要类群,在寒武纪大爆发期间海洋底栖动物群落结构构建中发挥着重要作用。研究表明华南最早的腕足动物出现在寒武系第二统。通过大量酸蚀处理和系统分析,发现华南寒武纪第二世碳酸盐岩相中赋存的腕足动物可分为8属12种和3个未定属种,都属于磷酸钙质舌形贝型,揭示了寒武纪早期磷酸钙质壳腕足动物的多样性及其形态差异。通过全球对比,力求探索寒武纪最早腕足动物的起源、多样性、分布与辐射。通过个体发育研究,揭示了异时发育在乳孔贝形态多样性中发挥的重要作用,同时表明寒武纪舌形贝型腕足动物发育中普遍存在滤食型浮游幼虫和变态发育的过程。因此变态发育的浮游幼虫可能是腕足动物的祖先特征。这与现代舌形贝明显不同,现代舌形贝的浮游幼虫为次级幼虫,演变为直接发育。此外,对比全球不同大陆腕足动物的首现,认为舌形贝型亚门腕足动物可能在寒武纪第二世初起源于东冈瓦纳与华南板块,随后开始向全球扩散。对寒武纪早期腕足动物多样性、个体发育与生物地层的深入研究,有助于增进对关键动物门类的早期起源与系统演化的认识,同时也将推动全球和区域寒武纪生物地层的划分与对比。

关键词: 腕足动物, 磷酸钙质壳, 个体发育, 多样性, 寒武纪第二世, 华南

Abstract:

Brachiopoda, as a key phylum of lophotrochozoan animals, plays an important role in studying the biodiversification of benthic communities during the Cambrian evolutionary radiation. Eight genera and twelve species of brachiopods, with the exception of three indeterminate taxa, are revealed and recovered from the Cambrian Series 2 carbonate facies near the Ediacaran-Cambrian boundary in the Yangtze Platform. They represent one of the earliest assemblages or components of brachiopods from South China. Based on the brachiopod data, the shell morphology and textural structures of the earliest variety of brachiopod groups and their occurrences and dispersals are discussed here to explore the origin and biodiversification of phosphatic-shelled brachiopods. We then summarised the recent research advances on the development of Cambrian brachiopods known from the Xihaoping Member of the Dengying, Shuijingtuo, and Yu’anshan formations of the Yangtze Platform. Lines of studies demonstrated that the linguliform brachiopods achieved adaptive diversity in the ornamentation of metamorphic and mature shells and shell structures, and in the configuration of pedicle foramen during the Cambrian explosion. In addition, three ontogenetic stages, including the pedicle foramen formation stage, pedicle foramen enclosure stage, and intertrough increasing stage, are observed in Cambrian Epoch 2 acrotretides. Compared with the ontogenetic developmental processes in different genera and species of acrotretides, the post-metamorphic allometry of shell valves was recognized as it played an increasingly significant role in the morphological diversification of different acrotretide clades. Furthermore, our recent studies on the earliest ontogenetic characteristics, preserved in the metamorphic shells in the oldest known lingulides and acrotretides, demonstrated that metamorphosis of planktotrophic larva is plesiomorphic for all brachiopods while the living lingulids are modified to obtain secondary larva with direct development. Interestingly, separate and diachronous occurrences of the oldest known brachiopods in the Cambrian palaeocontinents implied that linguliform brachiopods probably originated in East Gondwana and South China during the early Cambrian Epoch 2 and achieved a cosmopolitan dispersal subsequently. More studies on the diversity, ontogeny and biostratigraphy of the earliest phosphatic-shelled brachiopods will help us improve our understanding of their origination and the evolutionary changes of different lineages to uncover the correlation of Cambrian strata in detail.

Key words: brachiopods, phosphatic-shelled, ontogeny, diversity, Cambrian Epoch 2, South China

中图分类号:

P52
P534.4

张志亮, 陈飞扬, 张志飞. 华南寒武纪碳酸盐岩中最早腕足动物的辐射、发育与分布[J]. 地学前缘, 2020, 27(6): 79-103.

ZHANG Zhiliang, CHEN Feiyang, ZHANG Zhifei. The earliest phosphatic-shelled brachiopods from the carbonates of South China: their diversification, ontogeny and distribution[J]. Earth Science Frontiers, 2020, 27(6): 79-103.

图/表 11

图1 寒武纪生命大爆发期间基础动物、原口动物和后口动物依次成型(据文献[5])

Fig.1 Successive formation of basal metazoans, protostomes and deuterostomes during the Cambrian explosion.Adapted from [5].

图2 华南寒武纪第二世古地理及化石产地 a—华南寒武纪第二世古地理简图[10],标注化石产地;b—湖北宜昌艾家河剖面岩家河组(YJH)-水井沱组(SJT)界线,箭头标注水井沱组底部钙质结核;c—陕南镇巴小洋坝剖面灯影组(DY)-西蒿坪段(XHP)-水井沱组(SJT)界线,箭头标注腕足动物Eoobolus incipiens的最低层位。

Fig.2 Palaeogeography and fossil localities of the Cambrian Series 2 in South China

表1 华南寒武系第二统磷酸钙质壳腕足动物属种分类(据文献[17]略做修改)

Table 1 Phosphatic-shelled brachiopod taxa from Cambrian Series 2 in South China. Modified after [17].

腕足动物	目	超科	科	属种Genus/species	文献
寒武纪第二世浅水碳酸盐岩 Cambrian Epoch 2 Carbonate sediments	Lingulida	Linguloidea	Obolidae	Palaeobolus? liantuoensis	[31]
				Kyrshabaktella? sp.	[31]
				Spinobolus popovi	[10]
			Eoobolidae	Eoobolus incipiens	[29]
				Eoobolus? shaanxiensis	[31]
				Eoobolussp.	[17]
				Eoobolidae gen. et sp. indet.	[17]
			Lingulellotretidae	Lingulellotreta yuanshanensis	[32]
			Lingulellotretidae	Lingulellotreta ergalievi	[17]
		Acrotheloidea	Botsfordiidae	Botsfordia minuta	[31]
				Botsfordiidae gen. et sp. indet. A	[10]
				Botsfordiidae gen. et sp. indet. B	[17]
	Acrotretida	Acrotretoidea	Acrotretidae	Eohadrotreta zhenbaensis	[31]
				Palaeotreta zhujiahensis	[30]
				Palaeotreta shannanensis	[30]

图3 华南寒武纪第二世磷酸盐化壳体化石 a-p—峡东水井沱组;a—三叶虫Tsunyidiscus actus(感谢代韬提供照片);b-e—多节类三叶虫ptychopariids,AJH S05 AO-03,AJH S05 AO-06,AJH S05 Q-03,AJH S05 AO-05;f,g—微网虫Microdictyon sp.,AJH S05 BE04;h,i—高肌虫Houlongdongella sp.,AJH 8-2-3 B-13;j—棘皮动物骨板,AJH S05 G-20;k-n—软舌螺Paramicrocornus zhenbaensis,AJH 8-2-3 CM-07;o,p—似软舌螺Hyolithellus,AJH S05 BH-12;q—岩家河组原牙形刺Protohertzina,YJH S01C V-42;r-u—峡东水井沱组;r—似软舌螺Torellella,AJH S05 Q-25;s—海绵骨针,AJH S05 P-41;t,u—古杯化石Robustocyathellus sp.,AJH 8-2-3 BJ-12;v,w—陕南西蒿坪段寒武钉Cambroclavus fangxianensis,XYB 5 AN-05;x—陕南水井沱组软舌螺口盖Paramicrocornus zhenbaensis;y,z,ab—陕南水井沱组三叶虫Hupeidiscus orientalis,XYB S4-3 AN-15,XYB S4-3 AN-13。

Fig.3 Phosphatised shelly fossils from Cambrian Series 2 in South China

图4 寒武纪第二世澄江化石库软躯体保存腕足动物化石 a,b—Diandongia pista,ELI-BD 0082,ELI-BD 0056;c,d—Heliomedusa orienta,ELI-BH 0011,ELI-BH 0036;e,f—Lingulellotreta yuanshanensis ELI-BL409b,ELI-425;g,h—Eoglossa chengjiangensis ELI-BC 0118,ELI-BC 035;i,j—Xianshanella haikouensis ELI-BX 002A,ELI-BX003A;k,l—Kuangshanotreta malungensis ELI-B CLP K 015,ELI-CLP K 023A。

Fig.4 Soft bodied preserved brachiopods from the Cambrian Epoch 2 Chengjiang Lagerstätte

图5 陕南灯影组西蒿坪段始圆货贝Eoobolus incipiens 形态及结构 a-d—腹壳,箭头标示微弱发育的顶肌痕,ELI S3 AM-04;e,f—背壳ELI XYB S3 AM-04;g,h—较大的背壳发育倒三角形中脊,ELI S3 AK-09,ELI S3 AK-08;i—变态壳的圆坑状结构,ELI XYB S3 AM-02;j—后变态壳发育瘤点纹饰,ELI XYB S3 AM-03;k—放大显示瘤点构造,ELI XYB S3 AT 013。

Fig.5 Morphology and structure of Eoobolus incipiens from the Xihaoping Member, Dengying Formation of southern Shaanxi

图6 滇东地区梅树村剖面黑林铺组玉案山段底部磷酸钙质壳腕足动物Botsfordiidae gen.et sp.indet.B(博茨傅徒贝科未定属种) a,b—酸蚀处理获得的博茨傅徒贝破碎壳体,ELI-MSC-2-01 V-80;b—图a中的瘤状壳体纹饰的放大显示;c—博茨傅徒贝的瘤点纹饰,直径约30 μm,ELI-MSC-2-01 V-77;d—纵切面显示壳体结构,可见微弱的棒状结构,ELI-MSC-3-02 V-92。

Fig.6 Botsfordiidae gen. et sp. indet. B, a representative of phosphatic-shelled brachiopod from the bottom of the Yu’anshan Member, Heilinpu Formation of the Meishucun section in eastern Yunnan

图7 华南寒武纪第二世磷酸钙质壳腕足动物的形态与结构多样性 a—Eohadrotreta zhenbaensis,ELI-AJH S05 I143;b—Palaeobolus?liantuoensi,ELI-XYB S4-3 BQ-10;c—Botsfordia minuta ELI-XYB S4-2 BO-08;d—Eohadrotreta zhenbaensis,ELI S05 E-85;e—Spinobolus popovi,AEI-AJH S05 CA-04;f—Botsfordia minuta ELI-XYB S4-3 CK-19;g—Eohadrotreta zhenbaensis,ELI-AJH S05 D-94;h—Spinobolus popovi,ELI-AJH S05 64;i—Botsfordia minuta,ELI-XYB 13 CK-07;j—Eoobolus incipiens,ELI-XYB S3 AM-06;k—Lingulellotreta yuanshanensis,ELI-XYB S5-4 CH-03;l—Botsfordia minuta,ELI-XYB 13 CK-04;m—Eohadrotreta zhenbaensis,ELI-AJH 8-2-3 AC-11。

Fig.7 Morphology and structural diversity of phosphatic-shelled brachiopods from Cambrian Epoch 2 in South China

图8 寒武系水井沱组Eohadrotreta zhenbaensis的3个个体发育阶段(据文献[57]) a,b—肉茎孔形成阶段T1,ELI-AJH 8-2-1 AD2-15;c,d—肉茎孔封闭阶段T2,ELI-AJH 8-2-1 AE-03;e—交互沟伸长阶段T3,ELI-AJH 8-2-3 AD2-08。

Fig.8 Three ontogenetic stages of Eohadrotreta zhenbaensis from the Cambrian Shuijingtuo Formation. Adapted from [57].

图9 磷酸钙质壳腕足动物的生命周期,显示重要的最早期发育阶段(据文献[16]略作修改) a—现代舌形贝科Lingula,示次级幼虫的生命周期;b—寒武纪早期乳孔贝科Eohadrotreta原始幼虫,示变态发育完成于固着后不久。bs—幼年壳;es—胚胎期形成壳(embryoic shell);lse—幼虫刚毛;mas—成年壳;ms—变态壳(幼壳);mse—外套膜边缘刚毛;ns—少年壳;pr—胎壳;te—触手。

Fig.9 Life cycle of phosphatic-shelled brachiopods, demonstrating important earliest ontogenetic stage. Modified after [16].

图10 滇东、陕南与峡东地区寒武纪第二世的地层对比(据文献[10]略作修改)

Fig.10 Stratigraphic correlation of Cambrian Series 2 in eastern Yunnan Province (a), southern Shaanxi Province (b) and eastern Yangtze Gorges (c). Modified after [10].

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华南寒武纪碳酸盐岩中最早腕足动物的辐射、发育与分布

The earliest phosphatic-shelled brachiopods from the carbonates of South China: their diversification, ontogeny and distribution

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