Elsevier

Global and Planetary Change

Volume 163, April 2018, Pages 158-170
Global and Planetary Change

Prelude of benthic community collapse during the end-Permian mass extinction in siliciclastic offshore sub-basin: Brachiopod evidence from South China

https://doi.org/10.1016/j.gloplacha.2018.02.010Get rights and content

Highlights

  • Palaeocommunity turnover of brachiopod from basinal facies occurred in the upper part of the Clarkina changxingensis Zone.

  • Palaeocommunity diversity and brachiopod body size decreased during this turnover.

  • The dominant taxa of brachiopod palaeocommunity changed to more competitive genus during this turnover.

  • The palaeocommunity turnover was triggered by the decreased marine primary productivity and increased volcanic activity.

Abstract

Analysis of the Permian–Triassic palaeocommunities from basinal facies in South China provides an insight into the environmental deterioration occurring in the prelude to the mass extinction event. Quantitative and multivariate analyses on three brachiopod palaeocommunities from the Changhsingian to the earliest Triassic in basinal facies in South China have been undertaken in this study. Although the end-Permian extinction has been proved to be a one-stepped event, ecological warning signals appeared in the palaeocommunities long before the main pulse of the event. A brachiopod palaeocommunity turnover occurred in the upper part of the Clarkina changxingensis Zone, associated with a significant decrease of palaeocommunity diversity and brachiopod body size. During this turnover the dominant genera changed from Fusichonetes and Crurithyris (or/and Paracrurithyris) to the more competitive genus Crurithyris (or/and Paracrurithyris). The brachiopod palaeocommunity turnover was supposed to be triggered by the decreased marine primary productivity and increased volcanic activity. Moreover, such early warning signals are found not only in the deep-water siliceous facies, but also in the shallow-water clastic facies and carbonate rock facies in 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).

References (79)

  • D.Z. Piper et al.

    A modern vs. Permian black shale - the hydrography, primary productivity, and water column chemistry of deposition

    Chem. Geol.

    (2004)
  • K.T. Ratcliffe

    Palaeoecology, taphonomy and distribution of brachiopod assemblages from the Much Wenlock Limestone Formation of England and Wales

    Palaeogeogr. Palaeoclimatol. Palaeoecol.

    (1991)
  • N. Tribovillard et al.

    Trace metals as paleoredox and palaeoproductivity proxies: an update

    Chem. Geol.

    (2006)
  • P.B. Wignall

    Large igneous provinces and mass extinctions

    Earth Sci. Rev.

    (2001)
  • H.F. Yin et al.

    Two episodes of environmental change at the Permian–Triassic boundary of the GSSP section Meishan

    Earth Sci. Rev.

    (2012)
  • H.F. Yin et al.

    The end-Permian regression in South China and its implication on mass extinction

    Earth Sci. Rev.

    (2014)
  • D.X. Yuan et al.

    Revised conodont-based integrated high-resolution timescale for the Changhsingian Stage and end-Permian extinction interval at the Meishan sections, South China

    Lithos

    (2014)
  • Y. Zhang et al.

    Community replacement, ecological shift and early warning signals prior to the end-Permian mass extinction: a case study from a nearshore clastic-shelf section in South China

    Palaeogeogr. Palaeoclimatol. Palaeoecol.

    (2017)
  • G.A. Afanasjeva

    Changes in the communities of Paleozoic brachiopods due to their development of their filtering system

    Paleontol. J.

    (2009)
  • G.A. Afanasjeva

    Large extinctions of articulate brachiopods in the Paleozoic and their ecological and evolutionary consequences

    Paleontol. J.

    (2010)
  • S.D. Burgess et al.

    High-precision timeline for Earth's most severe extinction

    Proc. Natl. Acad. Sci.

    (2014)
  • A. Chao et al.

    Abundance-based similarity indices and their estimation when there are unseen species in samples

    Biometrics

    (2006)
  • Z.Q. Chen et al.

    The timing and pattern of biotic recovery following the end-Permian mass extinction

    Nat. Geosci.

    (2012)
  • R.K. Colwell

    Biodiversity: concepts, patterns, and measurement

    Princeton Guide Ecol.

    (2009)
  • C.P. Doncaster et al.

    Early warning of critical transitions in biodiversity from compositional disorder

    Ecology

    (2016)
  • D.H. Erwin

    The great Paleozoic crisis: life and death in the Permian

    (1993)
  • D.H. Erwin

    The Permo–Triassic extinction

    Nature

    (1994)
  • J.A. Fagerstrom

    Fossil communities in paleoecology: their recognition and significance

    Geol. Soc. Am. Bull.

    (1964)
  • S.B. Felitsyn et al.

    Mobility of phosphorus during the weathering of volcanic ashes

    Lithol. Miner. Resour.

    (2002)
  • Z.Z. Feng et al.

    Lithofacies Paleogeography of Permian of South China

    (1997)
  • W.J. Foster et al.

    Functional diversity of marine ecosystems after the Late Permian mass extinction event

    Nat. Geosci.

    (2014)
  • M.L. Fraiser et al.

    When bivalves took over the world

    Paleobiology

    (2007)
  • P. Frogner et al.

    Fertilizing potential of volcanic ash in ocean surface water

    Geology

    (2001)
  • S.J. Gould et al.

    Clams and brachiopods-ships that pass in the night

    Paleobiology

    (1980)
  • Ø. Hammer et al.

    PAST: paleontological statistics software package for education and data analysis

    Palaeontol. Electron.

    (2001)
  • W.H. He et al.

    Controls on body size during the Late Permian mass extinction event

    Geobiology

    (2010)
  • W.H. He et al.

    Systematics and palaeoecology of Changhsingian (Late Permian) Ambocoeliidae brachiopods from South China and implications for the end-Permian mass extinction

    Alcheringa

    (2012)
  • W.H. He et al.

    Changhsingian (latest Permian) deep-water brachiopod fauna from South China

    J. Syst. Palaeontol.

    (2014)
  • W.H. He et al.

    Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes

    Geobiology

    (2015)
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