Elsevier

Precambrian Research

Volume 346, 15 August 2020, 105801
Precambrian Research

An assemblage of macroscopic and diversified carbonaceous compression fossils from the Tonian Shiwangzhuang Formation in western Shandong, North China

https://doi.org/10.1016/j.precamres.2020.105801Get rights and content

Highlights

  • A Tonian assemblage of macroscopic carbonaceous compressions from North China.

  • In total, twelve genera, sixteen species and an unnamed filaments, are described.

  • Three taxa are multicellular and seven taxa are likely coenocytic algae.

  • The Tonian diversity of macroalgae is probably underestimated.

Abstract

Macroalgae play a crucial ecological and biogeochemical role in modern marine ecosystems. Although eukaryotic algae have a deep history within the Precambrian, most pre-Cryogenian alga fossils are microscopic and macroalga fossils are poorly documented during this period. Here we report a well-preserved assemblage of macroscopic and diversified carbonaceous compression fossils from the Tonian Shiwangzhuang Formation in western Shandong, North China. This assemblage consists of seventeen distinct morphotaxa, including eleven macroalgae, showing remarkable taxonomic diversity that is more diverse than many other Tonian macrofossil assemblages all over the world. Three taxa from the Shiwangzhuang assemblage show evidence of simple multicellularity with two of them consisting of giant cells and comparable to giant-celled green algae, and seven taxa with diagnostic transverse annulations are likely coenocytic algae. The Shiwangzhuang assemblage indicates that some eukaryotic clades had achieved macroscopic growth through simple multicellularity or coenocytism, paving the way, either ecologically or phylogenetically, for the eventual appearance of complex multicellularity. In addition, there are some biostratigraphically significant genera, including the Chuaria-Tawuia and Sinosabellidites-Protoarenicola-Pararenicola assemblages, indicating a Tonian age for the Shiwangzhuang Formation. This study also suggests that the Tonian diversity of multicellular or coenocytic macroalgae is probably underestimated, pending for a systematic and thorough investigation of Tonian macroscopic carbonaceous compressions. Macroalgae may have an initial ecological expansion, at least locally if not globally, in the Tonian Period.

Introduction

Macroalgae, which are usually multicellular or coenocytic eukaryotes, play an important role, both ecologically and biogeochemically, in modern marine ecosystems (Xiao and Dong, 2006). Molecular clock studies suggest that eukaryotic algae have an evolutionary deep history tracing back to the Paleoproterozoic to Mesoproterozoic (Parfrey et al., 2011, Sánchez-Baracaldo et al., 2017). However, fossil biomarkers indicate that prokaryotes were the only notable primary producers in pre-Cryogenian oceans and it was not until the Cryogenian that eukaryotic algae rose to ecological prominence (Brocks et al., 2017, Hoshino et al., 2017). Although there are a handful of macrofossils interpreted as multicellular or coenocytic algae in the Tonian (Butterfield, 2004, Butterfield et al., 1994, Dong et al., 2008, Qian et al., 2009, Sharma et al., 2009, Tang et al., 2017, Tang et al., 2020), the fossil record of macroalgae during this period is poorly documented, hampering our ability to evaluate the potential ecological and geobiological importance of early macroalgae.

Carbonaceous compressions, including Burgess Shale-type macrofossils and organic-walled microfossils, provide an important window into our understanding of the origin and early evolution of eukaryotes. Most macroalga fossils are preserved as carbonaceous compressions throughout the Proterozoic (Hofmann, 1994, Xiao and Dong, 2006). However, compared with Ediacaran macroalga assemblages (Xiao et al., 2002, Yuan et al., 2013), the taxonomic diversities of Tonian macroalga assemblages are usually considered much lower and their morphology relatively simple (Butterfield, 2009, Xiao, 2013). In this study, we describe an assemblage of macroscopic and diversified carbonaceous compression fossils from the Tonian Shiwangzhuang Formation (~910–720 Ma) of the Tumen Group from western Shandong, North China (Fig. 1, Fig. 2). The Shiwangzhuang assemblage shows notable taxonomic diversity which is not common in the Tonian. It consists of seventeen distinct morphotaxa, including eleven macroalgae. Among them, three taxa show evidence of simple multicellularity with two of them consisting of giant cells, and seven tubular taxa are likely coenocytic algae with diagnostic transverse annulations.

Section snippets

Geological setting

Meso-Neoproterozoic successions in Shandong Province are represented by the Tumen Group in Luxi (western Shandong) region and the Penglai Group in Ludong (eastern Shandong) region (Fig. 1), which are geographically separated by the Tanlu Fault (Bureau, 1991, Bureau, 1996, Song and Wang, 2003, You and Zhai, 1988, Zhai et al., 2007, Zhao and Zhai, 2013). The Tumen Group in Luxi region is mainly composed of pebbly sandstones, sandstones, shales, and limestones, and its exposure in Luxi region is

Materials and methods

Specimens were collected from a fossiliferous horizon (~74 m below the top of the Shiwangzhuang Formation) in the middle of the Tonian Shiwangzhuang Formation of the Tumen Group at the Baishicun section (36°30′39″N, 119°07′39″E), Anqiu region, Shandong Province, North China (Fig. 1, Fig. 2B). Macroscopic fossils are preserved on the bedding surface of argillaceous limestones, mostly as carbonaceous compressions, sometimes pyritized. Carbonaceous compressions present more details than pyritized

Summary of the Shiwangzhuang assemblage

In a recent study focused on the paleontology of the underlying Tongjiazhuang Formation of the Tumen Group, we have mentioned the occurrences of Tawuia dalensis Hofmann in Hofmann and Aitken, 1979 and Sinosabellidites huainanensis Zheng, 1980 from the Shiwangzhuang Formation (Li et al., 2019). Our continuous excavation has recovered more than 6000 specimens of carbonaceous compression fossils from the same fossiliferous horizon of the Shiwangzhuang Formation at the Baishicun section (Fig. 2,

Systematic paleontology

Genus Anqiutrichoides n. gen.

Type species.—Anqiutrichoides constrictus n. gen. and sp.

Diagnosis.Diagnosis is as for the type species.

Etymology.Derived from the Anqiu region, in which the fossil locality is located, and from Greek trichoma (hair), with reference to the filamentous habit.

Anqiutrichoides constrictus n. gen. and sp.

Figure 14

Diagnosis.Unbranched, uniseriate, multicellular trichomes with cell length less than or equal to cell width. Trichomes 400 μm to several hundred μm in width,

Biological affinities of the Shiwangzhuang assemblage

Four morphotaxa within the Shiwangzhuang assemblage, including Glomulus filamentum, Mucoplagum primitivum, Siphonophycus punctatum, and the unnamed filaments, are likely of cyanobacterial origin and can form colonies (Fig. 16); however, they do not exhibit cellular preservation. Filaments of S. punctatum are commonly considered as empty sheath of filamentous cyanobacteria (Butterfield et al., 1994, Schopf, 1968, Tang et al., 2013). Aggregated filaments of G. filamentum are interpreted as

Conclusions

Macroscopic carbonaceous compression fossils from the Shiwangzhuang Formation of the Tumen Group are delicately preserved and diverse. They are characterized by abundant eukaryotic macroalgae and filamentous cyanobacteria. A total of seventeen taxa have been recognized from the Shiwangzhuang Formation, including a new genus and six new species. The Shiwangzhuang assemblage is more diverse than many other Tonian macrofossil assemblages all over the world. Among them, three taxa show evidence of

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements

This research was supported by the National Key Research and Development Program of China (2016YFC0601001 and 2017YFC0603101), National Natural Science Foundation of China (41921002, 41502019, and 41602007), Chinese Academy of Sciences (QYZDJ-SSW-DQC009, XDB26000000, and XDB18000000), Shandong University of Science and Technology (SDUST) Research Fund (2015RCJJ009, 2015YQJH102, and 2015TDJH101), Taishan Scholars Project, and State Key Laboratory of Palaeobiology and Stratigraphy (20201102 and

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      Besides the compressions and impressions, a few calcite-filled or pyritized fossils are three-dimensionally preserved as spheroids or compressed spheroids (Hofmann, 1985; Zhang et al., 1991; Yuan et al., 2001; Wang et al., 2022). Moreover, circular, elliptical or oval compressions of Chuaria specimens were also reported from carbonate facies, e.g., the muddy dolomite of Tuanshanzi Formation near Jixian, northern China (Hofmann and Chen, 1981), dolostone of the Wynniatt Formation in Minto Inlier on Victoria Island, northwestern Canada (Hofmann and Rainbird, 1994) and argillaceous limestone of Shiwangzhuang Formation in Shandong, northern China (Li et al., 2020). For Chuaria from the Wynniatt Formation, specimens from the dolostone are somewhat bigger than those from the black shale, but they have the same form (Hofmann and Rainbird, 1994).

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