bioconstructions in the Gzhelian-Asselian (Late Pennsylvanian-early Permian) of Houchang, southern Guizhou, South China

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ORIGINAL PAPER

Factors controlling the development of Fomichevella coral

bioconstructions in the Gzhelian-Asselian (Late Pennsylvanian-early Permian) of Houchang, southern Guizhou, South China

Yongli Zhang¹_&Enpu Gong¹_&Wentao Huang¹_&Mark A. Wilson²_&Changqing Guan¹_&Xiao Li¹_&Lifu Wang¹_&

Junjie Wang¹_&Zhuowei Miao¹

Received: 10 October 2019 / Revised: 29 December 2019 / Accepted: 2 April 2020

#Senckenberg Gesellschaft für Naturforschung and Springer-Verlag GmbH Germany, part of Springer Nature 2020

Abstract

Bioconstructions (five biostromes, one patch reef, and one large reef) of the colonial coralFomichevellaare well exposed, if infrequent, in the Upper Pennsylvanian-lower Permian of the Houchang area (southern Guizhou, China). Though they are primarily built by the same species, different growth morphologies and internal constructions are evident, which indicate different factors controlling their formation and development. The coral biostromes alternate with shoal deposits, being attributed to level- bottom communities that were closest to the reef ecosystem. The patch reef consists of a simple coral framework and a low biodiversity community. In contrast, rigid skeletal frameworks ofFomichevellaare common and the biodiversity of the community is relatively higher in the large coral reef. Comparing these coral biostromes, the patch reef, and the large coral reef, several factors controlling their formation and development are suggested: stable substrate, water turbulence, sea-level fluctuation, and community structure. Hard substrates can provide stable habitats for the coral larvae during the initial colonisation stage, while the frequently changing sedimentary conditions were not conducive to the healthy development ofFomichevella. The simple community (e.g. the biostromes, patch reef) lacking encrusters and binders was too fragile to tolerate a turbulent environment.

In Houchang, though intrinsic and extrinsic factors influenced the development of theFomichevellabioconstructions, the extrinsic factors seem more important, which may be the key aspects controlling the Carboniferous coral bioconstructions.

Keywords Coral biostromes . Controlling factors . Gzhelian-Asselian . South China

Introduction

Reefs, the most complex and diverse marine ecosystems, both living and ancient, are extremely sensitive to environmental changes such as temperature, salinity, acidification, and depositional rate (Wood 1993; Hoegh-Guldberg et al. 2007).

Continual changes of environment are detrimental to the development of reef ecosystems. Sheehan (1985) argued that the evolutionary pattern of Phanerozoic reefs was quite similar to other marine communities. The complex, highly diverse eco- logic structure of reefs is the significant difference between

them and level-bottom communities, the re-establishment of which after an extinction is slow (Stanley 1981; Sheehan 1985). Environmental stability is necessary for sustainable reefs and the geobiological transition from shallow-water level-bottom community to the more complex reef ecosystem.

The Carboniferous was a stagnant period for reef ecosystems, especially for skeletal framework reefs, compared with the Devonian and Permian. From the global view, reefs developed throughout the Carboniferous, but lacked a consistent stable reef community (West 1988; Aretz and Vachard 2007;

Gong et al.2012; Yao et al.2016). After the mass extinction in the Devonian, though various reef types (e.g. coral reef, algal reef, microbial reef) have been documented in the Mississippian (Fang and Hou1985; West1988; Webb1998,1999; Nakazawa 2001; Aretz and Herbig 2003, 2008; Gong et al. 2012;

Rodríguez et al.2012; Chen et al.2013; Maillet et al.2020), biostromes characterised by diverse corals, calcimicrobes, calcisponges, brachiopods, and crinoids were conspicuous (Rodríguez 1996; Aretz 2001, 2010; Aretz et al. 2010;

* Wentao Huang

1310435@stu.neu.edu.cn

1 Department of Geology, Northeastern University, Wenhua Road 3-11, Heping District, Shenyang 110819, China

2 Department of Earth Sciences, The College of Wooster, Wooster, OH 44691, USA

https://doi.org/10.1007/s12549-020-00431-6

/ Published online: 20 June 2020

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Rodríguez et al.2013; Yao et al.2016; Huang et al.2018). A short proliferation phase of metazoan reef appeared in the late Viséan after the longest post-extinction reef recovery delay of the Phanerozoic (Yao et al.2019). Instead of further development of metazoan reef, algal reefs (e.g. phylloid algae reefs) became dominant after the Mississippian-Pennsylvanian boundary (West1988; Wahlman 2002; Gong et al. 2012;

Yan et al.2019), as some bioconstructions were built by chaetetid in Moscovian (Suchy and West1988,2001; West 1988; Nakazawa2001; Gong et al.2012; Zhang et al.2018).

The Carboniferous was a rebuilding stage of complex reef ecosystems after their complete collapse in the Late Devonian, forming a link in reef evolution from the Devonian to the Permian (Webb2002; Yao et al.2016).

In Guangxi and Guizhou province, South China, some Carboniferous bryozoan-coral, coral biostromes/reefs, microbial mounds, and phylloid algal reefs were documented and carefully studied (Fang and Hou1985; Gong et al.2007a,b;

Shen and Qing2010; Chen et al.2013; Yao et al.2016; Yao and Wang2016; Huang et al.2019). Gong et al. (2004) reported a rare large coral reef (Bianping coral reef) built by Fomichevella from the latest Carboniferous (Gzhelian, Triticites Zone) in the Houchang area, Guizhou Province.

An Asselian patch reef and five Gzhelian coral biostromes dominated by Fomichevella are also well developed in the Houchang area. Coral biostromes are the community closest to reef ecosystems, some of which have the potential to form reefs (Aretz and Herbig 2003; Yao et al. 2016). The Fomichevellabioconstructions of the Houchang area provide an ideal site to study the initiating conditions for a metazoan framework reef community. In this paper we focus on a continuous section containing five coral biostromes in the lower part and one patch coral reef in the upper part, and compare them with the Bianping coral reef (Triticiteszone) in the same region to reveal the initial conditions of reef community development. We also explore factors controlling theFomichevella bioconstructions in the study area and further discuss the characteristics of Carboniferous coral bioconstructions.

Geological setting and overview of the section

Geological setting

In the Pennsylvanian, shallow carbonate platforms were widespread in South China, including the Southwest, South Central, Yangtze, Longmenshan, and Western Yunnan platforms (Feng et al.1999) (Fig.1a). Our study area is situated on the margin of the Southwest platform, where Pennsylvanian and Permian strata are well preserved and crop out continu- ously (Bureau of Geology and Mineral Resources of Guizhou Province1987) (Fig.1b). The Pennsylvanian is characterised by shallow epicontinental sea deposits, comprising the

Weining and Maping Formations (Bureau of Geology and Mineral Resources of Guizhou Province 1987) (Fig.2). In the Maping Formation, many reefal buildups formed in high-energy margin environments have been studied (Gong et al.2004,2007a,b; Guan et al.2007; Sun et al.2007; Zhang et al.2007; Guan et al.2010).

Overview of the section

A continuous Maping succession (Upper Pennsylvanian- lower Permian) completely crops out near Lumazhai village, in the Houchang area (Fig.3), comprising thick-bedded lime- stones including bioclastic grainstone, bioclastic packstone, and bioclastic wackestone. The measured section (Lumazhai section) is about 41 m thick, with five coral biostromes in the lower part (distributed in 4–10 m) and a patch coral reef in the top (distributed in 37–40 m) (Figs.3and4). Benthic organisms are abundant in this section, including foraminifers (e.g.

fusulinids), brachiopods, corals,Tubiphytes, and algae. A de- tailed study of the fusulinids was carried out.Triticitesspp., Schwagerinaspp.,Quasifusulinaspp.,Schubertellaspp., and Qzawainellaspp. are recognised in the substrate and overlying beds of the biostromes that correspond to theTriticiteszone, indicating a Gzhelian age (Zhang and Zhou2004; Zhang et al.

2010a) (Fig.4). The fusulinids found in the patch reef are Sphaeroschwagerinaspp.,Paraschwagerina spp.,Triticites spp., andSchwagerinaspp., establishing an earliest Permian age (Zhang et al.2010a) (Fig.4). The first occurrence of the Sphaeroschwagerinaspp. is at 18.5 m marking the beginning of Sphaeroschwagerina Zone (Assleian age). Thus, the Gzhelian-Assleian boundary is situated around 18.5 m on Lumazhai section in this study, based on the fusulinids.

Materials and methods

The Lumazhai section was carefully measured (Fig. 3). The lithology, sedimentary structures, bed thickness, and compo- nents were documented in detail. About 150 representative sam- ples were collected. More than 300 thin sections and 45 polished slabs were prepared for microfacies and palaeoecological analysis. The abundance ofFomichevellais mainly based on semi- quantitative field observations. The material is housed at the Department of Geology, Northeastern University, Shenyang, China.

Fig. 1 Geological background information for the study area.aLatest Carboniferous palaeogeography of South China (after Feng et al.1999) and the location of the study site.bA geological map of the study area and the locations of coral biostromes and reefs

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Results

Microfacies analyses

The measured section is located near the entrance to Lumazhai village and comprises a series of shallow- water carbonates. Along this section, four microfacies types are distinguished.

Bioclastic grainstone

Bioclastic grainstone, widely distributed in the Lumazhai section, alternates with bioclastic packstone or wackestone (Fig.5a). It also is the substrate of coral biostromes and the overlying layers on the patch reef. Fining upward graded beds are common in the bioclastic grainstone (Fig.5a, b). Various bioclasts make up about 70–80% of the volume of the grainstone, including abundant foraminifers, mollusc shells, crinoids, algae, corals, and calcimicrobes (Fig. 5c). Most of the bioclasts are broken and rounded.

Bioclastic packstone

Bioclastic packstone contain various angular to sub- rounded skeletal fragments. Crinoids, foraminifers, and mollusc shells are the most abundant bioclasts, along with Tubiphytes, phylloid algae, corals, and bryozoans. Parts of the fossils are well preserved. It alternates with bioclastic grainstone and wackestone along the section (Fig. 5a). In Fig. 2 The stratigraphic framework of the upper Carboniferous-lower

Permian in southern Guizhou (Bureau of Geology and Mineral Resources of Guizhou Province1987; Zhang et al.2010a)

Fig. 3 Overview of the Lumazhai section and locations of coral biostromes (1–5) and the patch reef. The telegraph poles along the section are 12 m high

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some beds adjacent to bioclastic grainstone, micritic lithoclasts with medium-well rounded outlines are abundant (Fig.5d).

Bioclastic wackestone

Bioclastic wackestone is present in the section alternating with bioclastic packstone or grainstone. It also occurs in the internal parts of the coral patch reef and coral biostromes, filling in the spaces of coral frameworks (Fig. 6a). Only a few foraminifers, Tubiphytes, mollusc shells, crinoids, and calcimicrobes can be seen in the bioclastic wackestone (Fig. 6b). These fossils normally are fine fragments (< 0.5 mm). Ostracods are preserved with their valves articulated (Fig. 6c).

Coral framestone

Coral framestone is constructed byFomichevella, which can be seen in the patch reef at the top of Lumazhai section.

Fomichevellacorallites have diameters of 2–4 cm (Fig.6a).

Primary cavities with sparry calcite are common. The growth spaces of the coral are filled with bioclastic wackestone or packstone.

Bioclastic rudstone

Bioclastic rudstone occurs as the substrate of the coral patch reef in the top of the section. This microfacies contains abundant brachiopods of different sizes (1–5 cm) and lithoclasts.

The brachiopod shells are fragmented, and poorly sorted Fig. 4 The stratigraphic section

of the Lumazhai section.aThe whole section.bDetails of the lower part (grey area) in diagram a, showing the five coral biostromes and cyclic deposits

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(Fig.7a). Lithoclasts show angular outlines (Fig.7b). Calcite cavities are common in this microfacies.

Cyclic deposits

The lower part of Lumazhai section is characterised by subtidal cyclic deposits. The different microfacies types tend to be stacked in the same vertical order (Fig.8). Cycles fine upwards with a thickness of 0.5–1 m. Coarse bioclastic grainstone forms the base of the cycle, which is followed by

finer packstone. The top of the cycle comprises bioclastic wackestone with relatively high carbonate mud content. In some bioclastic wackestone beds, biostromes constructed by Fomichevellaare well preserved.

Coral bioconstructions

In the Houchang area, variousFomichevellabioconstructions (biostromes, patch reef, and large reef) are well presented in the lower Maping Formation (Gzhelian-Asselian). The Fig. 5 Photographs of bioclastic grainstone and packstone.aField

photograph of the cyclic deposits with graded beds in the Lumazhai section, showing the alternation of coarse grainstone and finer wackestone-packstone.bPolished slabs of the graded beds.cThin-

section photograph of bioclastic grainstone: fusulinids (Fu), foraminifers (Fo), crinoids (Cr).dThin-section photograph of bioclastic packstone with abundant micritic grains and fusulinids (Fu)

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biostromes and patch reef are explored in the Lumazhai section. The large reef is located in Bianping village, about 2.5 km from the Lumazhai section (Fig.1b).

Coral biostromes

Five coral biostromes are present in lower part of the Lumazhai section (Figs.3and4). These biostromes occur in the bioclastic wackestone. The second and third coral biostromes extend laterally, with 15–20 m length and 0.3–

0.5 m thickness, overlying bioclastic packstone-grainstone.

The other three coral biostromes are in smaller scale, 5–

10 m long and 0.2–0.4 m thick. The biodiversity of the coral layers is low, only a few foraminifers, algae,Ivanovia, and crinoids have been recognised. The well preserved colonial coralFomichevellais the main builder of the biostromes, with abundance about 20%, locally up to 30% (Fig.9a, b) of rock volume. Most of the corals rest on their sides, or even parallel to the substrate. Only a few corallites grow at a small angle to the bioclastic packstone-grainstone. Smaller corallites are usually distributed around the larger ones, and the spaces in the coral framework are filled with bioclastic wackestone.

Fig. 6 Photographs of bioclastic wackestone.aPolished slabs of coral framework and infilling wackestone:Fomichevella(F).bThin-section photograph of bioclastic wackestone in coral biostromes, with fine

crinoids (Cr).cThin-section photograph of bioclastic wackestone in coral frameworks, showing the well-preserved valves of the ostracod

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Ivanoviacan be seen in some biostromes with very low abundance, normally in small scale (< 15 cm). The beds overlying the coral layers are thick-bedded bioclastic packstone- grainstone containing abundant foraminifers and a few Fomichevellafragments.

Coral patch reef

TheFomichevellapatch reef, which has distinct topographic relief, crops out about 20 m wide and 10 m thick in the field, near the end of the section (Figs.3,4, and10a). The substrate is composed of bioclastic grainstone and rudstone (Fig.10b).

Fomichevella, phacelloid and with an erect growth habit, form the reef framework. Asexual reproductive budding of Fomichevellais visible. The reef community is simple and foraminifers and crinoids are common fossils. The spaces in the coral framework are filled with bioclastic wackestone- packstone. Bioclastic grainstone covers the coral patch reef, in which some coral debris are seen.

Large coral reef

The largeFomichevellareef, 80–100 m high and about 700 m long, is in theTriticiteszone and located in Bianping village (Fig.11a) (Gong et al.2004; Zhang et al.2010b). Bioclastic grainstone with foraminifers, crinoids, and brachiopod shells is distributed at the base of the coral reef (Fig. 11b) (Gong et al.2004; Zhang et al. 2010b). Several patch reefs were discovered in the lower part, including a phylloid algae reef, lime mound, and anIvanoviareef (Gong et al.2004; Zhang et al.2010b).Fomichevellain phacelloid form is the main reef framework and its corallites are pillared (Fig.11c, d) (Gong et al. 2004). They are arranged tightly side by side, and the open spaces are filled with micrite (Gong et al.2004). The reef assemblage was divided into six communities by Guan et al.

(2004), including Ivanovia, Eugonophyllum, microbes, Tubiphytes, fusulinids (e.g. Triticites, Schwagerina, Quasifusulina), brachiopods (e.g. Choristites, Squamularia, Echinaria),Fomichevella, foraminifers (e.g.Textularia), crinoids, gastropods, and bryozoans (Guan et al.2004).

Fig. 7 Photographs of bioclastic rudstone.aPolished slabs of bioclastic rudstone, with abundant brachiopod shells.b,cThin-section photographs of bioclastic rudstone: brachiopod (B), fusulinids (Fu), lithoclast (L), gastropod (G)

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Interpretation and discussion

In this study area, various coral bioconstructions (biostromes, patch reef, and large reef) are well presented in the lower Maping Formation (Gzhelian-Asselian).

These reefs are infrequent globally during the Gzhelian- Asselian. All coral bioconstructions in the study were mainly built by Fomichevella. Fomichevella preferred to colonise hard substrates, which was critical for the coral polyps to maintain a normal growth habit and resist waves (Bolton and Driese 1990; Taylor and Wilson 2003; Zhang et al. 2010b). Furthermore, marine environments with clean sea-water, medium water energy, and nutrient-poor water were more beneficial for the development of Fomichevella buildups (Zhang et al.

2010b). The different morphologies and the internal or- ganisations of the bioconstructions suggest differences in physical and biological factors controlling their formation and development (Kershaw and Keeling 1994).

Depositional environment

The study area was on a shallow carbonate platform environment in the Pennsylvanian, when many biodetritus banks, mounds, phylloid algae reefs, and coral reefs were present (Bureau of Geology and Mineral Resources of Guizhou Province 1987; Fan and Rigby 1994; Gong et al. 2004,

2007a; Zhang et al. 2007; Huang et al. 2019). Coral biostromes mainly occur in bioclastic wackestone, alternating with bioclastic grainstones and packstones. The fine skeletal grains and relatively high carbonate mud content in the wackestone suggests a low-energy depositional environment within deeper water (Egenhoff et al. 1999). Widespread bioclastic grainstone and packstone are the substrate and overlying bed of the biostromes. They are common shoal deposits, formed in shallow water environment with high water energy conditions (Flügel 2004; Bádenas and Aurell2010). Cyclic deposits comprising bioclastic grainstone, packstone, and wackestone in this section may indicate the sea-level fluctuations and frequent changes of sedimentary environments between stable and turbulent periods (Flügel2004; Bádenas and Aurell 2010). Fomichevella biostromes in bioclastic wackestone beds are interpreted as products of relatively stable intervals (Fig.12).

The patch reef developed on bioclastic rudstones. Poorly sorted bioclastic debris and angular lithoclasts in the rudstones suggest transport from the interior platform and rapid deposi- tion in a low-energy environment. The patch reef was formed when the depositional environment was relatively stable with clean seawater and medium water energy beneficial to Fomichevella. In the overlying bed of the reef, the common distribution of bioclastic packstone and grainstone implies shallowing water and an increase in water energy and depositional rate which may be responsible for the end of reef development.

The large coral reef was first reported by Gong et al. (2004) and further studied by Guan et al. (2004), Sun et al. (2007), and Zhang et al. (2010b). It was interpreted as a platform marginal reef with clean sea-water and medium water energy conditions (Gong et al.2004; Zhang et al.2010b).

Substrate

Bioclasts (e.g. brachiopod shells, crinoids), lithoclasts, rocks, and synsedimentary hardgrounds are common hard substrates for Fomichevella (Wilson and Palmer 1992; Taylor and Wilson2003; Zhang et al.2010b). Coral biostromes occur in bioclastic wackestone, forming interlayers in shallow-water shoal deposits. The substrates and overlying layers of these biostromes are mainly composed of bioclastic packstone- grainstone. The bioclastic packstone-grainstone underlying the biostromes in this study has little synsedimentary cement, implying that synsedimentary diagenesis was weak in the coral biostromes substrate (Wilson and Palmer1992; Taylor and Wilson2003; Zhang et al.2010b) and thus could not provide stable substrates forFomichevella. The available hard substrate forFomichevellalarvae to colonise was limited to the bioclasts and lithoclasts, which may have led to a low abundance of settling Fomichevellalarva when the environment was appropriate. As Fomichevella developed, previous Fig. 8 Model of cyclic deposits, consisting of bioclastic grainstone,

bioclastic packstone, and bioclastic wackestone

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corallites of the coral and the coral fragments could have provided the hard substrate for the settling larva. However, the development of Fomichevella communities was often interrupted by environmental changes (e.g. sea-level drop).

In this case, it was difficult to construct a reef complex.

In the biostromes, most of theFomichevellaare not in their growth position, seemingly resulted from storms. The fine skeletal grains and relatively high carbonate mud content in the wackestone suggests a low-energy depositional environment (Egenhoff et al.1999). TheFomichevellaskeletons are not fragmented and the corallite intervals are filled with abundant mud, which also argues against the storm influence. The life strategies and modes of attachment of the corals, the hydrodynamic environment in which the corals live, post- mortem sedimentologic and hydrodynamic factors, and the conditions of the substrate on which the corals colonise may control the preserved orientations of the corals (Elias1984;

Elias et al.1987,1988; Neuman1988; Speyer and Brett1988;

Bolton and Driese1990). The odd orientations of the corallites in the Fomichevella biostromes in this study could be the result of unstable substrate and weak current reworking (Bolton and Driese1990).

TheFomichevellalarge framework reef is a platform marginal reef complex (Gong et al. 2004; Zhang et al.2010b), developing on a hard substrate comprising several patch reefs, bioclastic grainstone, and massive brachiopod rudstone (Fig.13a). The strategy employed byFomichevellato secure a sufficient area of hard substrate was by larval aggregation (Fig.13b) (Zhang et al.2010b). Sufficient hard substrate con- tributes to effective connectivity between larvae and coral density, which are potential drivers of recovery and reassembly of reef communities (Zhang et al. 2010b; Bonuso et al.

2018; Gouezo et al.2019).Fomichevella individuals grew erect and rapidly occupied the available living space to build framework when the environment was stable (Fig. 13c) (Zhang et al.2010b).

Fig. 9 Photographs of coral biostromes.aField photograph of the third coral biostrome; the geological hammer is about 30 cm.bThe well- preservedFomichevella(F) in the coral biostromes, and the smaller coral

individuals (white arrow) around the larger ones.c,dCross section of Fomichevellaunder polarised microscope

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Community structure

The communities in coral biostromes are simple, mainly containingFomichevella, Ivanovia, foraminifers, and a few molluscs. Encrusters (e.g. microbes, algae) are absent.

Crinoids and algae debris normally are present as granular deposits. Microbial activities are few. Coral biostromes are designated as level-bottom community. The patch reef also represents a simple community. Fomichevella and Tubiphytesare builders and binders, respectively. Crinoids, foraminifers, and a few molluscs are the common reef- associated organisms.

In contrast, the large coral reef has a relatively more complex community (Gong et al.2004; Guan et al.2004; Zhang et al.2010b). A relatively stable microenvironment was cre- ated in the reef as theFomichevellaframework formed, which was favourable to reef dwellers. The biodiversity of the reef

community was improved by the settlement of brachiopods, gastropods, foraminifers,Tubiphytes, bryozoans, microbes, crinoids, and various algae (Gong et al. 2004; Guan et al.

2004; Zhang et al. 2010b).Tubiphytes and microbes were commonly associated with whole reef growth and fulfilled vital functions, which could bind the corals and fix sediments, contributing to the stabilisation of substrate and framework (Fig.13) (Zhang et al.2010b). Various algae were producers and promoted the material and energy cycles in the reef complex. Meanwhile, brachiopods, gastropods, fusulinids, foraminifers, and bryozoans were consumers in the community.

Framework reefs result from rapid skeletal production and the interdependent functional success of the constructor, baffler, and binder guilds (Fagerstrom 1988). These diverse organisms, with different roles in the large coral reef, interacted with each other to maintain the balance of the reef-building ecosystem. The healthy benthic community in the large coral reef Fig. 10 Field photographs of the coral patch reef.aOverview of the coral

patch reef. The person is 1.70 m high.bThe substrate of the patch reef consisting of abundant brachiopods (B) and gastropods (G).cField

photograph of the well-preservedFomichevella(F) in the patch reef, commonly with 3–4 cm transection in the field

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may have helped the reef ecosystem buffer external threats (DeCarlo et al.2017).

Comparisons

Comparing coral biostromes, patch reef, and large coral reef, the relatively stable environment, stable substrate, and healthy community structure are the key factors in the development of the large coral reef (Zhang et al.2010b). Though the lack of synsedimentary hardgrounds restricted the settling of Fomichevellalarvae in the initial stage of the biostromes, the frequent changes of circumstance (e.g. water depth, water energy, turbidity) seems more important in controlling the development of biostromes. The patch reef lay on the bioclastic grainstone and rudstone beds that provided enough hard substrate for theFomichevellalarvae when the environment was appropriate. The bioclastic wackestone-packstone in the patch reef suggested a relatively stable environment with medium

water energy during reef formation. However, the duration of stable environmental conditions was too short to build a complex reef system, resulting in a low biodiversity community in the patch reef. As the water shallowed and hydrodynamic energy increased, the patch reef was terminated.

Following the Late Devonian massive extinction, there was a global recession of skeletal reef systems in the Carboniferous.

Based on the Paleoreefs database of Kiessling (2003), reports of Carboniferous coral bioconstructions are limited and mainly concentrated in the Viséan (Fang and Hou1985; West1988;

Rodríguez1996; Aretz 2001; Aretz and Herbig2003; Aretz et al.2010; Rodríguez et al.2012; Somerville et al.2012; Chen et al.2013; Rodríguez et al.2013; Yao et al.2016; Huang et al.

2018), just a few of which are coral bioconstructions in the Pennsylvanian when algal reefs were the dominant type (Gong et al. 2004; Chang et al. 2012; Yang et al. 2013).

Most Carboniferous coral bioconstructions show biostrome or patch reef outlines while large coral reefs are scarce.

Fig. 11 Field photographs of the large coral reef.aOverview of the large coral reef; the two exposures are separated by an apparent fault; they thus likely represent the same reef complex; approximately 700 m long (Zhang et al.2010b).bThe substrate of the large coral reef containing

abundant brachiopods (B).c,dWell-preservedFomichevella(F) in the reef core. Asexual reproductive budding ofFomichevellais visible (white arrow). The marking pen in picturecis 14 cm

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Numerous intrinsic and extrinsic factors (e.g. palaeobiology and palaeoenvironment) influence the formation and development of bioconstructions (Aretz and Chevalier2007). Viséan coral biostromes distributed in China, Europe, and North Africa, show some similarities and differences in aspects of the dominant builders (Yao et al.2016), while their growth and demise are interpreted to have been mainly controlled by sea-level fluctuations on the basis of facies analysis (Aretz 2001; Aretz and Herbig 2003; Aretz et al.2010; Yao et al.

2016).

The Pennsylvanian-early Permian was the primary interval of the late Palaeozoic ice age, with dramatic climate changes and sea-level fluctuations (Isbell et al. 2003;

Fielding et al.2008; Isbell et al. 2012). The contempora- neous organic carbonate bioconstructions mostly lack large massive reef-building organisms (Wahlman 2002).

Throughout the Gzhelian-Asselian, phylloid algae dominated most tropical shallow-water bioconstructions, which may have benefitted from its rapid growth and recovery rates. The coral bioconstructions are rarely documented.

These coral bioconstructions (biostromes, patch reef, and large reef) in the Houchang area imply that coral had the potential for building framework reef in this interval.

However, the durations of living coral communities were usually too short to form complex bioconstructions, due to the frequent depositional environment changes. In this study, the extrinsic factors (e.g. sea-level changing, hard substrate, hydrodynamic energy) are considered as key aspects controlling the development of the bioconstructions.

Carboniferous coral bioconstructions were formed in a se- vere setting when profound global changes happened frequently: greenhouse to icehouse climates, supercontinent formation, frequent sea level changes (Isbell et al. 2003;

Aretz and Vachard2007; Isbell et al.2012).

Conclusions

1. Fomichevella is the primary builder of the metazoan bioconstructions that are well exposed in the Maping Formation (Gzhelian-Asselian) in the Houchang area, Guizhou, China. Though these bioconstructions are built byFomichevella, the difference in intrinsic and extrinsic factors controlling the formation and development of the bioconstructions shaped them into different morphologies: biostromes, patch reef, and large framework reef.

2. These coral bioconstructions formed in different depositional environment settings. Coral biostromes alternate with shoal deposits and are the products of short-term relatively stable intervals. The coral patch reef and large reef are the platform marginal reefs.

3. Available suitable substrate, hydrodynamic energy, depositional rate, sea-level change, and community structure stand out as significant developmental factors. Sufficient hard substrate is the initial condition for the colonisation of Fomichevella and the concentrated Fomichevella growth pattern is the prior condition to forming an effi- cient framework, while the frequently changing environment was detrimental to its healthy development. The relatively stable environment was a crucial condition for the transition from simple coral level-bottom (biostromes) or patch reef community to a more complex and diverse reef ecosystem.

Fig. 12 The formation and development model of coral biostromes.a Very shallow water with high energy conditions resulting in the shoal deposits lacking stable substrate.bFomichevellalarvae colonised when the environment was suitable.cThe highly frequent changes of the environment terminate the further development of the low density coral community

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Acknowledgements We thank Prof. Michal Zaton and Prof. Markus Aretz for their constructive comments on this manuscript.

Funding information This study was supported by the National Natural Science Foundation of China (Grant Nos. 41972002, 41202018, and 41572004).

Data availability The datasets used during this study are available from the corresponding author on request.

Compliance with ethical standards

Conflict of interest The authors declare that they have no conflict of interest.

Fig. 13 The formation and development model of the large coral reef.aThe substrate of the coral reef consists of three patch reefs and abundant bioclastic rudstone.bFomichevellalarvae colonised the hard substrates, with effective connectivity that is beneficial for the formation of functional frameworks. The microbial filaments can help to reinforce the framework and deposits.cFomichevellagrew rapidly to form reef framework.

The settlement of abundant organisms contributed to the reef complex ecosystem

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bioconstructions in the Gzhelian-Asselian (Late Pennsylvanian-early Permian) of Houchang, southern Guizhou, South China

Factors controlling the development of Fomichevella coral

bioconstructions in the Gzhelian-Asselian (Late Pennsylvanian-early Permian) of Houchang, southern Guizhou, South China

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