Prelude of benthic community collapse during the end-Permian mass extinction in siliciclastic offshore sub-basin: Brachiopod evidence from South China
Introduction
The end-Permian mass extinction, caused the demise of over 90% of all marine taxa and 70% of all terrestrial taxa (Sepkoski, 1981; Erwin, 1993, Erwin, 1994; Jin et al., 2000; Xiong and Wang, 2011). It also caused a significant change in palaeoecosystem structure – from the Palaeozoic-type communities to the Mesozoic–Cenozoic-type communities (e.g., dominance of marine benthonic organisms from brachiopod fauna to molluscan fauna) (Gould and Calloway, 1980; Sepkoski, 1981; Fraiser and Bottjer, 2007; Chen and Benton, 2012; Payne et al., 2014). A considerable, and still expanding, body of literature has been devoted to the causes, patterns and timing of the mass extinction event itself (e.g., Knoll et al., 2007; Shen et al., 2011; Yin et al., 2012; Sun et al., 2012; Burgess et al., 2014; Song et al., 2014; Wang et al., 2014) and ecosystem reconstruction in the aftermath (e.g., Twitchett et al., 2004; Payne, 2005; Chen et al., 2005a, Chen et al., 2007; Brayard et al., 2006; Chen et al., 2010; Pietsch and Bottjer, 2010; Algeo et al., 2011; Dineen et al., 2014; Foster and Twitchett, 2014; Petsios and Bottjer, 2016).
The end-Permian extinction has been proved to be a short-time event and happen between 251.941 ± 0.037 Mya and 251.880 ± 0.031 Mya (Shen et al., 2011; Burgess et al., 2014). There has been limited research on the palaeocommunities during the pre-extinction interval before the end-Permian mass extinction (Shen and Shi, 1996; Wang and Sugiyama, 2000). Recently, a section-based case study reported the benthic palaeocommunity transition in the pre-extinction interval (Changhsingian) in the nearshore clastic-shelf Zhongzhai section of South China (Zhang et al., 2017). This kind of study on the marine palaeocommunity change prior to the end-Permian mass extinction, especially on section-based quantitative fossil data from varied depositional environments, could provide a new perspective for the mechanism of the mass extinction event.
Herein, this study aims to investigate:
- 1)
if the pre-extinction benthic palaeocommunity transition also occurred in siliciclastic offshore sub-basin;
- 2)
when the pre-extinction benthic palaeocommunity transition occurred (in which conodont zone);
- 3)
the potential causes of pre-extinction benthic palaeocommunity transition in siliciclastic offshore sub-basin.
This section-based study presents the turnover of benthic brachiopod palaeocommunities from the Changhsingian (latest Permian) to the earliest Triassic at a high stratigraphic resolution (based on conodont and ammonoid biostratigraphy and eventostratigraphy). The comprehensive, statistical analyses of the brachiopod fossil data and the refined stratigraphic correlation contribute significantly to our understanding of the end-Permian mass extinction event. It also contributes to our knowledge on the environmental deterioration affecting the marine palaeocommunity transition prior to the mass extinction, and finally leading up to the major crisis itself.
Section snippets
Depositional setting and age correlation
Palaeogeographically, the South China Basin was divided into deep-water siliceous sub-basin, shallow-water carbonate platform and shallow-water clastic shelf in the Lopingian (Fig. 1.4; Yang et al., 1987; Feng et al., 1997). The deep-water sub-basin of South China is mainly distributed in Guangxi, Guizhou, Hunan, Anhui, Jiangxi Province and Chongqing (Fig. 1.4), and is characterised by grey black thin-bedded siliceous mudstone and shale, intercalated with muddy limestone. It predominantly
Materials and methods
Brachiopods analysed in this paper were systematically sampled at regular stratigraphical intervals throughout the three studied sections. There were 3390 brachiopod specimens collected from the upper Dalong Formation to the lower Yinkeng Formation at MJS, 2209 brachiopod specimens collected from the upper Dalong Formation to the lower Daye Formation at RCP and 3598 brachiopod specimens collected from the upper Dalong Formation to the lower Luolou Formation at XM (Fig. 1, Fig. 2). When counting
Division of Changhsingian brachiopod associations in three studied sections
In the Q-mode cluster analysis, samples are grouped into clusters with similar occurrence and abundance of taxa. At MJS, three associations (A, B and C) are recognised (Fig. 3). The association A is represented by samples from beds 3 to 6 (equivalent to the ammonoid Tapashanites Zone). The association B is recognised in beds 12 to 16 (equivalent to the conodont Clarkina changxingensis Zone or/and C. yini Zone). The association C is recognised in beds 18 to 20 (equivalent to the conodont
Taphonomy of brachiopod faunas from the three studied sections
The pre-burial transportation of loose shells by strong ocean currents normally only occurs along shores and in the shallow-water part of marine environments (Fagerstrom, 1964). It has been also proved that long-distance transport of skeletal remains is not very common in the basin benthic environment (Stanton Jr., 1976; Warme et al., 1976). In the three studied sections, most of the thin-shelled single valves of the dominant taxa, Fusichonetes and Crurithyris, are still very complete and not
Conclusion
In summary, our evidence suggested that the brachiopod palaeocommunity changes occurred earlier than the end-Permian mass extinction, which is consistent with the environmental deterioration prior to the main pulse of crisis in the three main sedimentary facies (including shallow-water clastic shelf, shallow-water carbonate platform and deep-water siliceous rock sub-basin) in South China. In particular, the changes of brachiopod palaeocommunity present at least three main features, as follows:
Acknowledgement
We thank G. R. Shi and Z. Q. Chen for helpful suggestions on the paper, Y. Zhang for very constructive help on revising the paper, Matthew E. Clapham and anonymous reviewers for their helpful reviews. This paper is supported by the Natural Science Foundation of China (Grant Nos. 41772016, 41372030, 41602017, 41730320), the Ministry of Education of China (B08030 of 111 Project), the Foundation of the Geological Survey of China (No. 121201102000150012).
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