Keratose sponge fabrics from the lowermost Triassic microbialites in South China: Geobiologic features and Phanerozoic evolution

https://doi.org/10.1016/j.gloplacha.2022.103787Get rights and content

Highlights

  • •
    Keratose sponge consortia from the microbialite deposits near the P-Tr boundary in South China.
  • •
    Calcified sponge skeleton might have firmed the framework of sponge-microbial build-ups.
  • •
    Deep-time keratose sponges favorite low-latitude regions and overall flourished in the aftermaths of major extinctions.

Abstract

The Permian-Triassic mass extinction (PTME) strongly devastated marine ecosystems, and, consequently, sponges, especially the reef-building clades, suffered dramatic losses in biodiversity. The Early Triassic therefore was believed to be an evolutionary gap for sponges. Microbialites spread over shallow marine carbonate settings across the entire low-latitude Tethys region following the PTME and occupied the ecospace that the pre-extinction metazoan reefs left. Here, we report putative keratose sponge consortia from the microbialites near the Permian-Triassic boundary in the Xiushui, Laolongdong, and Dongwan sections, South China. The putative keratose sponges exhibit vermiform, filamentous textures forming maze-like networks. Within the keratose sponge-microbial fabrics, the calcified sponge skeleton might firm the overall framework of microbialite, promoting construction of the sponge-microbial build-ups. The coeval occurrence of putative keratose sponges in both eastern Palaeotethys and central Neotethys regions indicates that sponges may have widely spread and played important roles in constructing metazoan-microbial reefs in earliest Triassic. Besides, several characteristics are conducive to keratose sponge surviving the stressful environments after the PTME. Global dataset shows that keratose sponges mostly confined to tropic and temperate zones. Keratose sponges overall flourished in coincidence with occurrence abundance peaks of microbial reefs during the Phanerozoic history, and they seem to be particularly abundant and widespread in the Cambrian-Ordovician, Late Devonian and the aftermath of the end-Permian mass extinction.

Introduction

Marine ecosystem suffered a dramatic loss in metazoan biodiversity from the Permian-Triassic mass extinction (PTME; Erwin, 2006), which gave opportunities for microbial community to bloom in the earliest Triassic oceans (Chen and Benton, 2012). Consequently, microbialite deposits spread over shallow carbonate platform settings in low-latitude Tethys region immediately after the PTME, and they usually addle the Permian-Triassic boundary (PTB) in South China (Yang et al., 2011; Kershaw et al., 2012; Chen et al., 2019). As a crucial link between biotic extinction and recovery processes, the PTB microbialite has attracted increasing interests from palaeontologists, geobiologists, and sedimentologists (Baud et al., 1997; Yang et al., 2011; Kershaw et al., 2012; Jiang et al., 2014; Baud, 2018; Chen et al., 2019). However, only few studies concerned the relationship between the interior architecture and potential constructor of the microbialite. Microorganisms may have acted as the key constructors of the PTB microbialites since microbes and their remains have been widely documented from these biosedimentary deposits (Ezaki et al., 2003, Ezaki et al., 2008; Wang et al., 2005; Yang et al., 2011, Yang et al., 2019; Li et al., 2015; Adachi et al., 2017; Fang et al., 2017; Tang et al., 2017; Wu et al., 2017; Bagherpour et al., 2017; Heindel et al., 2018; Chen et al., 2019; Pei et al., 2019). Except for microbial communities, metazoans are also rather abundant in the PTB microbialites, including a diverse community of ostracods, microconchids, foraminifers, gastropods, and bivalves (Yang et al., 2011, Yang et al., 2015a, Yang et al., 2015b; Forel et al., 2013; Hautmann et al., 2015; Wu et al., 2017; Forel, 2015; Foster et al., 2018, Foster et al., 2019; Su et al., 2021), but they were dwellers instead of builders in the microbialite ecosystem.
Three-Dimension reconrtruction has unraveled that the some vermiform- or network- or dot-like clump-shaped fabrics within the carbonates actually belong to keratose sponge (demosponge) (Luo and Reitner, 2014). Later, the same fabrics have been widely recognized from the PTB microbialites in Armenia, Central Iran, and southern Turkey of the Neotethys region, and these microbialites therefore were termed the sponge-microbial build-ups (Friesenbichler et al., 2018; Heindel et al., 2018; Baud et al., 2021). In contrast, although South China records most abundant PTB microbialites (PTBMs) in the world (Chen et al., 2019), the sponge-microbial fabrics have been rarely reported from PTBMs in this region. So far, only two suspicious cases were mentioned from the PTB microbialites in South China (Ezaki et al., 2008; Baud et al., 2013), and sponge-microbial fabrics of the PTB microbialites remain unclear in terms of preservation and spatial distributions.
Besides, as common reef builders, sponges and microbes usually dominate carbonate factories alternately or cooperatively during the Phanerozoic history (Chen et al., 2019). For instance, ancestral sponges and calcimicrobes constructed the first metazoan-dominating reef in the early Cambrian (Riding and Zhuravlev, 1995). Since then, sponge-bearing microbialite deposits have been well recorded (Olivier et al., 2004; Brayard et al., 2011; Lee et al., 2014), particularly in the aftermaths of major mass extinctions (Delecat et al., 2011; Friesenbichler et al., 2018; Kershaw et al., 2021). Some suspicious biostructures in the Cambrian microbialites that were thought to be of a microbial origin have also been re-assigned to sponges (Lee and Riding, 2021a). The alternation and cooperation of sponges and microbes in constructing carbonate build-ups might be related to various palaeoenvironmental factors. Recently, increasing attention has been paid to the importance of sponge in marine ecosystem due to their ability in activating bottom water and translating dissolved nutrients to organic particles that are available to organisms at higher trophic levels (Goeij et al., 2013). Thus, detailing structures of sponge-microbial build-ups may enable us to better understand the initiating engineering mechanism of ecosystem recovery after major mass extinctions.
Recently, we also have found some filamentous or vermiform, maze-like network or dot-like clump structures, identical to keratose sponge fabrics (Luo and Reitner, 2014; Friesenbichler et al., 2018) within the frameworks of the PTB microbialites in South China, strengthening the universality of the sponge-microbial mode in the PTB microbialites. Except for water depth, the newly found sponge-microbial build-ups have no essential difference from the ones in Central Neotethys. Thus, this paper aims to document geobiologic features of keratose sponge consortia from three PTB microbialite deposits in South China (Fig. 1), which may provide insights into metazoan resilience in devastated environments immediately after the PTME. Finally, a global dataset of deep-time keratose sponges is updated to provide insights into the spatiotemporal distributions and Phanerozoic evolution of this unique metazoan clade.

Section snippets

Geological and stratigraphic settings

Three PTB microbialite-bearing sections in South China were situated in low-latitude eastern Palaeotethys region during the Permian-Triassic (P-Tr) transition (Fig. 1). Pronounced microbialite deposits developed along the margins of the Yangtze Platform or on some isolated platforms within the intra-platform basin (i.e. the Nanpanjiang basin) (Chen et al., 2019). Location and stratigraphy of these three sections are introduced as below.
The Xiushui section (Fig. 2, Fig. 3), ~2 km northwest of

Material and method

After examining 600 thin-sections from 30 PTB microbialites in South China (Chen et al., 2019), we identified these keratose sponges under microscope from the Xiushui, Dongwan, and Laolongdong sections. In each section, more than 30 rock samples were collected, slabbed, and thin-sectioned using conventional petrologic techniques, from the uppermost Changhsingian reefal carbonate to the muddy limestone capping the PTB microbialite. Each microbialite is sampled intensively at an interval of

Sponge fabrics in the Xiushui section

In Xiushui, the putative keratose sponge consortia were identified from a thin-bedded, spar-cemented oolitic limestone below the basal microbialite unit (Fig. 2). It overlies unconformably above the uppermost Changhsingian bioclastic limestone (Fig. 3B-C; Wu et al., 2014; Wu et al., 2017). Above this oolitic limestone layer, the thrombolite is characterized by a clotted texture of reddish sparitic patches embedded in a grey micritic matrix (Fig. 6B), whereas the overlying dendrolite texture

Identification of keratose sponge

The sparitic vermiform or dot-shaped structures are often present in micritic clumps within deep-time microbialites, and they often have been treated as microbial bioconstructors (Kershaw et al., 2002, Kershaw et al., 2021). However, the reconstructed 3-D architectures of automicritic clumps with anastomosing fibrous structures documented from the Devonian and Triassic microbialites show that these filamentous networks are morphologically akin to that of keratose demosponges. These vermiform or

Conclusion

Putative keratose sponge consortia are documented from the strata immediately above the end-Permian mass extinction horizon in three localities: Xiushui, Laolongdong, and Dongwan in South China. The keratose sponge consortium shows vermiform, filamentous and dot-like aggregates forming maze-like networks. The calcification of spongin skeleton might have promoted construction of the keratose sponge-microbial build-ups. The coeval occurrences of keratosa in both Neotethys and Palaeotethys regions

Declaration of Competing Interest

We declare that we do not have any commercial or associative interest that represents a conflict of interest in connection with the work submitted.

Acknowledgments

We thank Cui Luo, Joachim Reitner, and Yu Pei for constructive discussions on keratose sponges. Guest editor Stephen Grasby, Jeong-Hyun Lee, and two anonymous reviewers are also acknowledged for their critical comments and constructive suggestions, which have greatly improved the quality of the paper. This study is supported by NSFC grants (41821001, 41930322) and the 111 Project of China (BP0820004). Siqi Wu is grateful to Chinese Scholarship Council for providing her an overseas visiting

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