Skip to main content

Main menu

  • Home
    • Journal home
    • Lyell Collection home
    • Geological Society home
  • Content
    • Online First
    • Issue in progress
    • All issues
    • Thematic Collections
    • Supplementary publications
    • Open Access
  • Subscribe
    • GSL fellows
    • Institutions
    • Corporate
    • Other member types
  • Info
    • Authors
    • Librarians
    • Readers
    • GSL Fellows access
    • Other member type access
    • Press office
    • Accessibility
    • Help
    • Metrics
  • Alert sign up
    • eTOC alerts
    • Online First alerts
    • RSS feeds
    • Newsletters
    • GSL blog
  • Submit
  • Geological Society of London Publications
    • Engineering Geology Special Publications
    • Geochemistry: Exploration, Environment, Analysis
    • Journal of Micropalaeontology
    • Journal of the Geological Society
    • Lyell Collection home
    • Memoirs
    • Petroleum Geology Conference Series
    • Petroleum Geoscience
    • Proceedings of the Yorkshire Geological Society
    • Quarterly Journal of Engineering Geology and Hydrogeology
    • Quarterly Journal of the Geological Society
    • Scottish Journal of Geology
    • Special Publications
    • Transactions of the Edinburgh Geological Society
    • Transactions of the Geological Society of Glasgow
    • Transactions of the Geological Society of London

User menu

  • My alerts
  • Log in
  • My Cart

Search

  • Advanced search
Journal of the Geological Society
  • Geological Society of London Publications
    • Engineering Geology Special Publications
    • Geochemistry: Exploration, Environment, Analysis
    • Journal of Micropalaeontology
    • Journal of the Geological Society
    • Lyell Collection home
    • Memoirs
    • Petroleum Geology Conference Series
    • Petroleum Geoscience
    • Proceedings of the Yorkshire Geological Society
    • Quarterly Journal of Engineering Geology and Hydrogeology
    • Quarterly Journal of the Geological Society
    • Scottish Journal of Geology
    • Special Publications
    • Transactions of the Edinburgh Geological Society
    • Transactions of the Geological Society of Glasgow
    • Transactions of the Geological Society of London
  • My alerts
  • Log in
  • My Cart
  • Follow gsl on Twitter
  • Visit gsl on Facebook
  • Visit gsl on Youtube
  • Visit gsl on Linkedin
Journal of the Geological Society

Advanced search

  • Home
    • Journal home
    • Lyell Collection home
    • Geological Society home
  • Content
    • Online First
    • Issue in progress
    • All issues
    • Thematic Collections
    • Supplementary publications
    • Open Access
  • Subscribe
    • GSL fellows
    • Institutions
    • Corporate
    • Other member types
  • Info
    • Authors
    • Librarians
    • Readers
    • GSL Fellows access
    • Other member type access
    • Press office
    • Accessibility
    • Help
    • Metrics
  • Alert sign up
    • eTOC alerts
    • Online First alerts
    • RSS feeds
    • Newsletters
    • GSL blog
  • Submit

Brachiopod-dominated communities and depositional environment of the Guanshan Konservat-Lagerstätte, eastern Yunnan, China

View ORCID ProfileFeiyang Chen, View ORCID ProfileGlenn A. Brock, View ORCID ProfileZhiliang Zhang, View ORCID ProfileBrittany Laing, Xinyi Ren and View ORCID ProfileZhifei Zhang
Journal of the Geological Society, 178, jgs2020-043, 18 September 2020, https://doi.org/10.1144/jgs2020-043
Feiyang Chen
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Formal analysis (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Writing - Original Draft (Lead)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Search for this author on this site
  • ORCID record for Feiyang Chen
Glenn A. Brock
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Search for this author on this site
  • ORCID record for Glenn A. Brock
Zhiliang Zhang
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Search for this author on this site
  • ORCID record for Zhiliang Zhang
Brittany Laing
2Department of Biological Sciences, Macquarie University, , , Australia
3Department of Geological Sciences, University of Saskatchewan, , , Canada
Roles: [Conceptualization (Equal)], [Formal analysis (Equal)], [Methodology (Equal)], [Writing - Original Draft (Supporting)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Search for this author on this site
  • ORCID record for Brittany Laing
Xinyi Ren
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
Roles: [Investigation (Equal)], [Methodology (Equal)], [Writing - Review & Editing (Supporting)]
  • Find this author on Google Scholar
  • Search for this author on this site
Zhifei Zhang
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Search for this author on this site
  • ORCID record for Zhifei Zhang
  • For correspondence: elizf@nwu.edu.cn zhangelle@126.com
PreviousNext
  • Article
  • Figures & Data
  • Info & Metrics
  • PDF
Loading

Abstract

The Guanshan Biota is an unusual early Cambrian Konservat-Lagerstätte from China and is distinguished from all other exceptionally preserved Cambrian biotas by the dominance of brachiopods and a relatively shallow depositional environment. However, the faunal composition, overturn and sedimentology associated with the Guanshan Biota are poorly understood. This study, based on collections through the best-exposed succession of the basal Wulongqing Formation at the Shijiangjun section, Wuding County, eastern Yunnan, China recovered six major animal groups with soft tissue preservation; brachiopods vastly outnumbered all other groups. Brachiopods quickly replace arthropods as the dominant fauna following a transgression at the base of the Wulongqing Formation. A transition from a botsfordiid-, eoobolid- and acrotretid- to an acrotheloid-dominated brachiopod assemblage occurs up-section. Four episodically repeated lithofacies reveal a relatively low-energy, offshore to lower shoreface sedimentary environment at the Shijiangjun section, which is very different from the Wulongqing Formation in the Malong and Kunming areas. Multiple event flows and rapid obrution are responsible for faunal overturn and fluctuation through the section. A detailed lithofacies and palaeontological investigation of this section provides a better understanding of the processes and drivers of faunal overturn during the later phase of the Cambrian Explosion.

Supplementary material: Composition and comparison of the Malong Fauna and the Guanshan Biota is are available at: https://doi.org/10.6084/m9.figshare.c.5080799

Discoveries of spectacular soft-bodied animal assemblages from Cambrian Konservat-Lagerstätten around the world have provided incredible insights into the anatomy, behaviour, ecology and early evolution of complex metazoans (Paterson et al. 2011; Hu et al. 2013; O'Brien and Caron 2016; Aria and Caron 2017; Hou et al. 2017; Yang et al. 2018; Liu et al. 2020; Z.F. Zhang et al. 2020a). Early Cambrian Konservat-Lagerstätten from China – such as the Niutitang Fauna, Chengjiang Biota, Guanshan Biota, Shipai Biota, Balang Fauna, Kaili Biota and the newly discovered Qingjiang Biota (Peng et al. 2005; Zhao et al. 2005; Hu et al. 2013; J. Liu et al. 2016; Hou et al. 2017; Fu et al. 2019) – span a wide range of geological time and provide a unique opportunity to map changes in early Cambrian ecological communities over time (Chen et al. 2019). The Guanshan Biota (Cambrian Series 2, Stage 4) supplementary material, one of the oldest Konservat-Lagerstätten from South China, occurs in the Wulongqing Formation (Hu et al. 2013) in eastern Yunnan. Younger than the famous Chengjiang and Malong biotas (Cambrian Series 2, Stage 3), but older than the Kaili and Burgess Shale biotas (Miaolingian Series, Wuliuan Stage), the Guanshan Biota is a significant evolutionary bridge in our understanding of the chronology of the Cambrian radiation and its aftermath (J. Liu et al. 2012a; Hu et al. 2013). Recent intensive, although preliminary, excavations reveal that the Guanshan Biota is composed of 14 major animal groups and various ichnotaxa (Hu et al. 2010, 2013; Chen et al. 2019). Uniquely, the Guanshan Biota is dominated by brachiopods, which serves to distinguish it from all other Cambrian Konservat-Lagerstätten, which are dominated (in terms of diversity and relative abundance) by euarthropod groups. Faunal overturn between the Chengjiang, Malong and Guanshan biotas suggests that the sessile benthic members of the assemblages are affected by the same factors that affect mobile trilobites (Luo et al. 2008; Chen et al. 2019). Furthermore, the Wulongqing Formation is characterized by bioturbated, thinly bedded sandstones, siltstones and mudstones, which crop out widely in eastern Yunnan, South China (Fig. 1) and represent a transgressive systems tract directly after the Hongjingshao Formation (Hu et al. 2010). Previous, very generalized, sedimentological work on the Wulongqing Formation suggests a relative shallow (shoreface to offshore transitional) depositional environment (Hu et al. 2010; Chen et al. 2019), which is distinct from the generally deeper water (in some cases slope to basin) setting of most other early Cambrian deposits that preserve soft tissues (Ivantsov et al. 2005; Collom et al. 2009; Peel and Ineson 2011).

Continuous exploration and research in the Guanshan Biota has led to the discovery of multiple new localities and increased systematic descriptions of the fossil taxa (Hu et al. 2010, 2013; J. Liu et al. 2012a, 2016; Hopkins et al. 2017; Li et al. 2017; Chen et al. 2019) (Fig. 1), including documentation of one of the oldest examples of kleptoparasitism in the fossil record (Z.F. Zhang et al. 2020a). The Wulongqing Formation is generally poorly exposed at most sites and artificial cover by urban landscaping has obscured many of the classic flat-lying sites. There has been a dearth of even basic ecological analyses of the faunal assemblages from the Guanshan Biota, and the detailed sedimentology and lithology of the succession are very poorly resolved.

Fig. 1.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 1.

Localities of the Guanshan Biota and distribution of lower Cambrian outcrops in eastern Yunnan, South China. The Shijiangjun section in Wuding county is represented by locality 1.

This paper aims to comprehensively document the lithofacies and sedimentology of the basal part of the Wulongqing Formation hosting soft-bodied fossils at the Shijiangjun section, the best-exposed succession in Wuding county, eastern Yunnan (Fig. 1). These data will help to decipher the relationships between microfacies, sedimentary events and faunal overturn after transgression and how fluctuations in depositional environments affect the faunal composition during the later stages of the Cambrian evolutionary radiation. The uppermost Hongjingshao and lower Wulongqing formations are exposed in the new section with a very clear conformable stratigraphic contact (Figs 2 and 3). This provides an opportunity to document temporal changes in the faunal composition and sedimentary environments at the centimetre scale based on lithological, sedimentological, palaeontological and ichnological evidence. This detailed study enables an interpretation of the depositional environment associated with the lower Wulongqing Formation and facilitates a better resolution of the process and drivers of faunal overturn that distinguish the Guanshan faunas from the Wuding, Malong and Kunming areas (Hu et al. 2013; Chen et al. 2019).

  • Download figure
  • Open in new tab
  • Download powerpoint
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 2.

Stratigraphic column, sedimentology, facies, structures, bioturbation index and pie charts of relative faunal abundance in the lower Wulongqing Formation at the Shijiangjun section. The 41 rock samples taken for cutting and polishing are marked on the left-hand side of the column. The datum point (0 cm) is the boundary between the upper Hongjingshao and lower Wulongqing formations. Four facies (F1, F2, F3 and F4) were recognized. The bioturbation index for each polished rock samples was evaluated based on the extent to which bioturbation disrupted the primary bedding. The composition of the fossil assemblage is shown by pie charts at the phylum level and the brachiopod genus level. Ass., assemblages; BI, bioturbation index; F., facies. HJS, Hongjingshao Formation; Sam., sample number; WLQ, Wulongqing Formation.

Fig. 3.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 3.

Field photographs of the lower Wulongqing Formation at the Shijiangjun section. Yellow upper case letters mark the layers yielding a fossil assemblage in accordance with Figure 2. (a) General view of the lower part of the section. The yellow line on the bottom of the section indicates the lithological contact between the Hongjingshao and Wulongqing formations. (b) Load casts at the bottom of the sandstone deposits. (c) Wavy bedding structure above layer C. (d) Normal graded bedding from fossil-yielding layer D, scale bar: 1 cm. (e) Gutter casts, lenticular bedding and wavy ripples at the Shijiangjun section. (f) Plan view of gutter casts from fossil-yielding layer F. (g) The simplified palaeoenvironmental reconstruction for the Guanshan Biota from Wuding area. HJS, Hongjingshao Formation; WLQ, Wulongqing Formation.

Materials and methods

The Shijiangjun section (25° 35′ 11″ N, 102° 22′ 22″ E) was measured through the uppermost Hongjingshao and lower Wulongqing formations and large-scale sedimentary features were noted. A total of 2988 fossil specimens were collected in one four-week field season sequentially and independently from ten contiguous siltstone and mudstone layers varying in thickness from 6 to 110 cm (Fig. 2; Table 1). Whole fossils were identified and classified to the phylum level and, where applicable, brachiopod genera were identified. Faunal relative abundances are based on all the well-preserved fossils, whereas trace fossils and fragmentary and unidentifiable specimens, as well as all shell concentrations, were excluded.

View this table:
  • View inline
  • View popup
Table 1.

Relative abundance and diversity of fossil taxa collected from the lower Wulongqing Formation at the Shijiangjun section, Wuding county

Lithological samples (n = 41) in oriented plaster jackets were collected at intervals from mudstone and sandstone layers through the section (Fig. 2; Table 2). All the samples collected for rock slabs and thin sections were cut and polished at the Shaanxi Key Laboratory of Early Life and Environments, China and revealed the vertical internal organization of the physical and biogenic sedimentary structures. Scanning of the polished slabs was achieved using an Epson V370 photo-scanner at Macquarie University, Australia. Following the methodology outlined by Dorador et al. (2014) and Dorador and Rodríguez-Tovar (2018), Adobe Photoshop was used to digitally improve the visibility (contrast) of the sedimentary and ichnological structures. Sedimentary characteristics, including grain size, lithology, sedimentary structures and vertical bioturbation intensity were recorded (Fig. 2; Table 2). The percentage bioturbation in each sample was evaluated using Adobe Photoshop (Cao et al. 2015; Gougeon et al. 2018). The bioturbation area was selected using the lasso tool and recorded through the measurement log in pixels. This was then divided by the total area in pixels to determine the percentage of bioturbation. These percentages were then used within the bioturbation index (BI) scheme of Taylor and Goldring (1993). All the rock samples and fossil specimens investigated are deposited in the Early Life Institute (ELI) and the Department of Geology, Northwest University, Xi'an China.

View this table:
  • View inline
  • View popup
Table 2.

Summary of lithology, sedimentary structures, taphonomic characters, fossil concentrations, bioturbation index and occurrence of soft tissues observed in the Wulonqging Formation from the Shijiangjun section

Results

Geological setting, locality and section

The stratigraphic section is 8 m thick and composed of distinctive intercalated beds of thin to thick (5–60 cm), very fine to very coarse sandstone, siltstone and mudstone (Fig. 3a). Rare gravels and isolated pebbles occur in sandstone samples S2, S3, S4, S5, S6, S9 and S16, in addition to two layers of purple muddy medium to coarse sandstone (S15 and S16), which contained 3–5% oolite grains (Fig. 2; Table 2). Commonly developed primary sedimentary structures include massive bedding, normal graded bedding (Fig. 3d), lenticular bedding and wavy bedding (Figs 2, 3c, e). The contacts between the sandstones and mudstones are sharp. The most common local erosion structures include gutter casts, erosional scour and low ripple marks (Figs 2, 3b, e, f). The measured section has an overall low level of bioturbation, with some highly bioturbated beds occurring in the middle part of the section (3.3–5.3 m) accompanying the only identified trace fossil Teichichnus? isp. (Fig. 2). Based on lithological, sedimentary and ichnological features, the section is divided into four distinct facies that repeat and cycle throughout the section, as shown in Figures 2–5.

Fig. 4.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 4.

Polished slabs of the lower Wulongqing Formation at the Shijiangjun section, with facies classification and sample numbers in parentheses. (a, b) Slabs of Facies 1 showing wavy laminations, graded lamination, lenticular lamination and erosive base. (c) Silty mudstone without sedimentary structures representing Facies 2. (d–g) Massive sandstone deposits representing Facies 3. (d, e) Poorly sorted, angular to sub-angular clasts with few granules. (f, g) Highly bioturbated sandstone with glauconite grains. (h) Mudstone without structures (Facies 4). Scale bars 5 mm.

Fig. 5.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 5.

Photomicrographs of thin sections from the lower Wulongqing Formation at the Shjiangjun section showing four lithofacies types. (a) Medium sorted irregular grains from Facies 1. (b) Graded laminations with an erosional base from Facies 1. (c) Mudstone with low content of well-sorted silt grains from Facies 2. (d) Poorly sorted grains with low sphericity from Facies 3. (e) Common glauconite grains within Facies 3; note the iron oxides within grains (black arrows). (f) Highly bioturbated sandstone from Facies 3. (g) Poorly sorted sandstone from Facies 3, coarse grains are irregular with low sphericity. (h) Uniform mudstone of Facies 4. All photomicrographs were taken with parallel light except (d), which is under cross-polarized light. Scale bars 1 mm.

Lithofacies identification and interpretation

Facies 1: low–medium bioturbated and interbedded mudstones–siltstones/sandstones

Facies 1 consists of thinly bedded mudstone with thin to thick laminated siltstone and/or very fine sandstone (Figs 4a, b and 5a). The silt and sand grains are medium to well-sorted, mainly angular to subrounded, low to high sphericity with increasing sphericity up-section (Fig. 5a). Fine to medium sandstone intercalations occur as lenticular and wavy bedding (Fig. 4a). Laterally discontinuous millimetre-scale (mainly 3–5 mm with some c. 1 mm) silt laminations are common. Graded laminations (4–10 mm) manifest either as a sharp horizontal contact or an erosional base (sole marks) (Figs 4b and 5b). The contact between sand and mud is nearly always sharp. Bioturbation is generally indistinct and unidentifiable, with Teichichnus? isp. documented in two samples (Fig. 2). The bioturbation index ranges from 0 to 3, with a predominant index of 0–1 (up to 4.89% disturbance). More heavily bioturbated beds exist locally (M5 and M20) with indexes of 2–3 recording up to 40% sedimentary fabric disturbance (Fig. 2; Table 2). The graded laminations and erosive bases suggest deposition from decelerating flows (Bouma 1962; Kneller 1995). The medium maturity of the sand/silt laminations probably indicates a certain degree of winnowing and transportation.

Interpretation. The interbedded mudstone and sandstone reflect an alternation of quiet water sediment fallout (low energy) combined with relatively high-energy flows (Buatois et al. 2012; Majid et al. 2017).

Facies 2: Silty mudstones

Facies 2 is represented by uniform mudstones with occasional millimetre-scale silt laminations (≤1 mm) (Fig. 4c). The silt grains are moderately sorted, angular to subrounded (low content) and of low sphericity (Fig. 5c). Interestingly, the M10 layer (Fig. 2) contains a higher concentration of muscovite than any other layer. Fragmentary shelly fossils are often present and are preserved parallel or oblique to bedding, with a particularly high concentration of trilobite fragments documented in layer M25. Bioturbation is rare (BI = 0), with the percentage bioturbation never exceeding 1% (Fig. 2; Table 2).

The absence of rheological surfaces on the silty mudstone packages indicates a relatively low-energy hydrodynamic system (Zavala et al. 2012). Abundant sub-parallel to oblique brachiopod and/or trilobite fragments within the mudstone indicate transportation by currents (Fürsich et al. 1992).

Interpretation. High rates of fallout or other unobservable environmental stressors (e.g. oxygen, salinity or temperature) may be responsible for the relative absence of bioturbation. As a result, the relatively structureless silty mudstone packages are interpreted as deposited from rapid fallout from suspension during quiet periods of fair weather conditions (Maceachern et al. 1999).

Facies 3: low to highly bioturbated glauconitic sandstones

Facies 3 consists of very fine to very coarse sandstone with rare granule- to pebble-sized clasts (Fig. 4d–g). The granules and pebbles predominately occur in samples S2–S6, S9 and S16 (Figs 2 and 4e; Table 2). The medium- to very coarse-grained sand beds from the lower and upper part of this section are characterized by very poorly to poorly sorted grains distributed within the intervals 0–2.1 m and 4.6–5.0 m (Figs 4d, e and 5d, g). Coarse grains are mainly angular to subrounded and dominated by low to medium sphericity (Fig. 5d, g). Although the very fine- to medium-grained sand beds from interval 2.2–4.2 m are mainly moderately sorted (Fig. 4f), few beds show medium to high sphericity. Two beds (S15 and S16) contain 1–5% elongate ooids. Most of the ooids are oval and few are rounded.

The sandstone beds are either characterized by a homogeneous uniform grain size or high bio-disturbance, which has destroyed the original sedimentary structures. Only levels S7 and S8 show weakly normal graded bedding. Sand beds S11–S15 show a relatively higher content of mud and a higher percentage of bio-disturbance (BI = 2–5) (Fig. 2; Table 2). The bioturbation index and bio-disturbance reach a peak of BI = 5 and 98.76% within S12 (Figs 4f and 5f), followed by S11 (80.88%) and S14 (76.16%) (Fig. 4g). However, more than half of the sandstones below S11 show scarce or no bioturbation (Fig. 2; Table 2).

The occurrence of syngenetic glauconite grains within the sandstones of Facies 3 is unique (Fig. 4f, g; Table 2). These grains were identified based on their green colour, random microcrystalline internal texture and aggregate polarization (Baioumy and Boulis 2012). They are, in some instances, coated and replaced by iron oxides (mostly hematite and goethite). These grains occur in every sandstone interbed at relatively low contents (Fig. 2; Table 2). The grains are usually medium sorted, subrounded to rounded and of medium sphericity (Fig. 5e). Although glauconite cannot be used as a specific environmental indicator (Mcrae 1972; Chafetz and Reid 2000; Chafetz 2007), it is commonly associated with transgressive systems tracts (Delamette 1989; Garzanti et al. 1989; Amorosi et al. 2012; Banerjee et al. 2012, 2017; Rudmin et al. 2017). Different types of glauconite (i.e. autochthonous, parautochthonous and detrital) can be determined based on the criteria proposed by Amorosi (1997). The glauconite that usually occurs in detrital granular and sand facies lacks a diffuse green pigmentation, which often alternates between glauconite-rich and glauconite-free layers, and can be interpreted to indicate an allochthonous (e.g. parautochthonous or detrital) origin (Amorosi 1997; Baioumy and Boulis 2012). By contrast, the low compositional and structural maturity of Facies 3, as well as a lack of glauconite in the older Hongjingshao Formation, implies a parautochthonous origin (Amorosi 1997; Baioumy and Boulis 2012), in which the autochthonous glauconites have been transported a short distance from their original location by waves, storm currents and/or gravity flow processes.

Local observations of Facies 3 show that these sandstones have a low compositional and textural maturity, which suggests that the sediments were deposited with minimal traction and clast collisions from a proximal sediment source. Therefore the clasts retain their immature, angular texture (Henstra et al. 2016). Storm deposits are generally understood to consist of well-sorted sand with a fining-upwards sequence that reflects the waning storm waves (Swift et al. 1983; Saito 1989; Duke et al. 1991; Cheel and Leckie 1993; Meldahl 1993). The storm flow usually converts to a turbidity current as the power of the storm flow weakens near the storm wave base, resulting in the suspended mud and gravel depositing together with fine suspended sediments during recessive periods (X. Liu et al. 2012b).

Interpretation. The common occurrence of poor bedding and disordered accumulation indicate fairly rapid suspension fall out without winnowing (Stow 2005), probably affected by gravity flow deposition in relatively deeper water (Wu et al. 2016). The sharp contacts at the lower and upper boundaries between the sandstones and mudstones show that each sandstone layer represents a single event. However, the changing grain size inside the thin sandstone units shows an unstable hydrodynamic environment. Facies 3 is interpreted to have been deposited within lower shoreface zone formed near the storm wave base and was affected by multiple pulses of gravity flows.

Facies 4: mudstones

Facies 4 represents mudstones with occasional interbedded wisps of silt (Figs 4h and 5h). The mud layers are considerably thicker (2.5–3 cm) than in other facies (Fig. 5h). The silt laminations are fairly thin (0.3–1 mm) with sharp erosive bases and a crudely micro-graded lower part and structureless upper part. Shelly fossils preserved within Facies 4 are usually parallel to sub-parallel to the bedding plane. The bio-disturbance within Facies 4 is the lowest among the four facies, only up to 0.15%, resulting in a low bioturbation index (BI = 0) (Fig. 2; Table 2).

Interpretation. These sedimentary features, along with the soft tissue preservation associated with Facies 4, suggest a mainly rapid deposition (obrution) of suspended muds settling from weak storm flows in a relatively low-energy environment (Zhu et al. 2001).

Composition and relative abundance of fossil assemblages

Thousands of well-preserved fossils spanning six key animal groups (n = 2988) were collected from the lower Wulongqing Formation at the Shijiangjun section during one four-week field season. The taxa include brachiopods, arthropods, hyoliths, priapulids, vetulicolians and anomalocaridiids in descending order of rank abundance (Table 1). All these taxa are also found in the Wulongqing Formation from the Kunming and Malong areas (Hu et al. 2013; Chen et al. 2019). Brachiopods, arthropods and hyoliths form the three main components, with up to 98.9% of the total number of specimens (Table 1; Fig. S1). Even though the anomalocaridiids, vetulicolians and priapulids are rare in this section, they are very important elements of Cambrian Burgess Shale-type Lagerstätten (Zhao et al. 2010; Paterson et al. 2011; Smith 2015). Four genera of organophosphatic brachiopods, including Neobolus, Eoobolus, Westonia (Luo et al. 2008), Linnarssonia (Duan et al. 2020) and two calcareous taxa (Kutorgina and Nisusia) occur throughout the section. Neobolus is the most abundant genus (40.2%), followed by Eoobolus (28.9%) and Westonia (27.2%). However, arthropods remain the most diverse group, composed of trilobites, bradoriids, Guangweicaris, Panlongia, Isoxys, Tuzoia and Leanchoilia. Among these, trilobites are the most abundant taxon (82%) (Table 1).

Fossil data from every mudstone layer was obtained during four weeks of intensive fieldwork in 2018 (Table 1). The fossil composition within assemblages A and B is similar, consisting of five animal groups, while faunal diversity decreases in assemblages C–F. This is followed by an increased diversity associated with faunal assemblages G–J. Faunal assemblage I has the highest diversity, with almost all taxa known from the entire section concentrated in this assemblage. Assemblage F has the greatest abundance of fossils (n = 748) accounting for 25% the total number of individuals, followed by assemblages C, B, G and J (Fig. 2; Table 1).

The relative abundance of individual specimens from ten sampling layers was obtained (Fig. 2; Table 1) to gauge the baseline assemblage structure. Assemblages A and B are dominated by arthropods, accounting for 63.6 and 59.8%, respectively. Brachiopods dominate all other assemblages from layers C–J, with some fluctuation of composition in the relative abundance between brachiopod taxa. The abundance of brachiopods reaches a peak within assemblage F. Hyoliths, a common early Cambrian group, occur throughout the entire section, except for assemblage G. Anomalocaridiids, vetulicolians and priapulids are interspersed irregularly within the assemblages.

The relative abundance of six genera of brachiopods throughout the section is very instructive (Fig. 2; see also Fig. 7b). Assemblage A is composed, almost equally, of three genera (Neobolus, 36.4%; Eoobolus, 36.4%; and Linnarssonia, 27.2%), whereas assemblage B contains a higher proportion of Neobolus (51.1%), with the relative abundance of the remaining two taxa 26.1 and 22.8%, respectively. Westonia occurs as a small proportion of assemblage C, whereas Neobolus and Eoobolus together exceed 97%. Assemblages C and D are mainly composed of Eoobolus (20.5 and 62.8%, respectively) and Neobolus (77.3 and 26.7%, respectively) with minor Westonia. By contrast, Westonia reaches a higher relative abundance (26.2%) in assemblage F. Eoobolus dominates assemblages G and H (52.4 and 64.2%, respectively), where Westonia also reaches a higher proportion of the assemblage (44.7% in G). Assemblages I and J are both dominated by Westonia, with 61.5 and 88% relative abundance, respectively; Eoobolus (24 and 9.6%, respectively) ranks second in these assemblages. The rare calcareous brachiopods Kutorgina and Nisusia are restricted to the upper part of the section in assemblages I and J (Figs 6g and 7b).

Fig. 6.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 6.

Exquisitely preserved soft-bodied fossils from the lower Wulongqing Formation at the Shijiangjun section. (a) Brachiopod Linnarssonia concentration from assemblage A (ELI-SJJ-001). (b) Trilobite Palaeolenus concentration from assemblage B (ELI-SJJ-002). (c, d) Brachiopods Neobolus and Westonia concentrations from assemblage F (ELI-SJJ-003, ELI-SJJ-003-2). (e) Brachiopod Neobolus with well-preserved parasitic tubeworms, indicated by arrows (assemblage B, ELI-SJJ-004). (f) Brachiopod Westonia preserved with mantle canals (assemblage F, ELI-SJJ-005). (g) Rare brachiopod Nisusia sp. (assemblage J, ELI-SJJ-006). (h) Well-preserved coiled palaeoscolecidan (assemblage F, ELI-SJJ-007). (i) Posterior part of an indeterminate vetulicolian (assemblage A, ELI-SJJ-008). (j) Trilobite Palaeolenus preserved with the rare digestive system (assemblage B, ELI-SJJ-009). Scale bars: (a, e–h and j) 2 mm; (b–d, i) 1 cm.

Fig. 7.
  • Download figure
  • Open in new tab
  • Download powerpoint
Fig. 7.

Stratigraphic fluctuation in the relative abundance of the community at the (a) phylum level and (b) brachiopod genus level from the Shijiangjun section.

The lower part of the Wulongqing Formation (0–6 m) at the Shijiangjun section also contains distinctive brachiopod and trilobite fossil concentrations (Fig. 2; Table 2). The concentrations preserved in coarser sandy deposits (especially Facies 3) are highly fragmented (although also fragile and thin) and moderately well-sorted, which indicates a relatively high level of energy and transportation (Table 2). Some well-preserved shell concentrations are also preserved within thin mud beds (e.g. Facies 1 and 4), occasionally restricted to single bedding planes, and in a relative sense these thin shells are characterized by low levels of fragmentation, poor sorting, low to medium disarticulation, and occur sub-parallel to bedding planes with a high ratio (>50%) of conjoined brachiopod shells with more or less soft tissue preservation. These taphonomic proxies indicate a relatively rapid obrution deposit and minimal transportation (Figs 2 and 6a–d; Table 2). The shell concentrations from the Shijiangjun section are either monospecific or paucispecific, dominated by brachiopods or trilobites (Fig. 6a–d). These concentrations are nearly always restricted to specific layers. For example, abundant Palaeolenus are exclusively found within layer M6 in assemblage B (Fig. 6b), whereas a concentration of Linnarssonia shells is known within layer M3 in assemblage A (Fig. 6a). The brachiopod concentrations from assemblage F are most abundant and mainly composed of monospecific layers of Neobolus (Fig. 6c) or Westonia (Fig. 6d), respectively. The Eoobolus and Westonia shell concentrations extend to the upper part of the section (Fig. 2). Throughout the section, brachiopod concentrations are completely restricted to Facies 1 and 2, whereas trilobite concentrations are mainly associated with Facies 4, which is restricted to assemblage B (Fig. 2).

Remarkable soft tissue preservation occurs in all assemblages except D and E, demonstrating the high preservation potential within facies at the Shijiangjun section of the Wulongqing Formation in the Wuding area. Tube-dwelling organisms encrusting to Neobolus shells (see Z.F. Zhang et al. 2020a) (with exceptionally preserved setae and soft viscera) are fairly common within the lower part of the section within mudstone beds (layers A, B, C and F) (Fig. 6e). Abundant specimens of Westonia display high-quality soft tissue preservation from assemblage F, including setal fringes and mantle canals (Fig. 6f). Palaeoscolecidan worms, as an important component of lower Paleozoic soft-bodied assemblages, were found throughout the section, except for assemblage C (Fig. 6h). Relatively rare vetulicolians occur at the base and in the upper part of the section (assemblages A, B, I and J) (Figs 2 and 6i). Anomalocaridiids are the rarest element in the section, only preserved as isolated frontal appendages in assemblages I and J. The rare oldest known digestive system of trilobites (Hopkins et al. 2017) have also been preserved in the Wuding area, but only in assemblage B (Fig. 6j).

Discussion

Depositional environment

Heterolithic successions consisting of sandstone beds interbedded with mudstones are usually deposited below the fair weather wave base and above the storm wave base (Dott and Bourgeois 1982; Myrow and Southard 1996; Dumas and Arnott 2006; Bullimore et al. 2008; Buatois and Mángano 2011; Eide et al. 2015). These beds are commonly described as tabular (Elliott 1978; Coe et al. 2003) and often show abundant erosive gutter casts (Eide et al. 2015). The alternation of mudstone (Facies 1, 2 and 4) and sandstone (Facies 3) layers – in addition to graded lamination/bedding, wavy bedding, ripple marks and gutter casts from the Shijiangjun section – suggests a depositional environment close to the storm wave base, which underwent multiple depositional events and episodic cycles (Shanmugam 2002; Hu et al. 2010; Buatois and Mángano 2011).

Previous studies have interpreted the sedimentary environment associated with the Guanshan Biota as mainly offshore transition with common storm events (Hu et al. 2010; Chen et al. 2019), which is comparable with the Cambrian Stage 4 Emu Bay Shale from Australia (Paterson et al. 2016) and the Ordovician Fezouata Biota (Martin et al. 2016). However, typical storm-generated structures such as hummocky cross-stratification, an indicator of oscillatory combined flows reflecting deposition under high-energy storm conditions (Arnott and Southard 1990; Cheel 1990; Southard et al. 1990; Cheel and Leckie 1993; Yokokawa et al. 1999; Dumas et al. 2005) are absent in the Wuding succession.

The occurrence of erosive bases, ripple marks, wavy bedding, fine-graded bedding, gutter casts and multiple massive fine to coarse deposits indicates a complex hydrodynamic environment, with less frequent waves and distal storms (Zhu et al. 2001; Buatois and Mángano 2011; X. Liu et al. 2012; Michalík et al. 2013). Periodic subaqueous gravity flows resulted in the deposition of distinctive centimetre-scale sandstone interbeds (Facies 3) at the Shijiangjun section. Hence the sedimentary environment of the lower Wulongqing Formation in the Wuding area is largely the result of fluctuating wave energy, distal storms and gravity flows.

The centimetre-scale conglomerates characterized by high sphericity reported from the Wulongqing Formation at Malong and Kunming represent high-energy channels, probably proximal to the shoreface (Hu et al. 2010; Chen et al. 2019). The absence of basal conglomerates and the occurrence of medium to very coarse sandstones with few granules at the base of the Wulongqing Formation in the Wuding area (Fig. 4e) suggest a relatively deeper and low-energy clastic sedimentary environment than that in the Malong and Kunming areas (Hu et al. 2010; Chen et al. 2019), although this remains to be tested because detailed continuous successions of the Wulongqing Formation have not been studied sedimentologically. Overall, the depositional environment here is interpreted as offshore to lower shoreface as defined by Martin et al. (2016) and the offshore zone of Buatois and Mángano (2011), which slightly extends below the storm wave base (Fig. 3g).

Faunal overturn and assemblage composition

The baseline time series of the fossil data recovered from the lower Wulongqing Formation at the Shijiangjun section reveals a unique transition in the structure of the benthic community over time (Fig. 2; Table 1). The relative abundance of six key animal groups, including six brachiopod genera, from ten sampled layers demonstrates gradual replacement, overturn and fluctuation in the faunal composition (Figs 2 and 7). Although arthropods dominate the base (0–1.1 m) of the section (assemblages A and B), the proportion of brachiopods gradually increases, replacing arthropods as the dominant fauna in assemblages C–J, reaching peak abundance (97.99%) within assemblage F (Figs 2 and 7a). Although there is a fluctuation in the relative abundance of brachiopods through assemblages G–J (c. 60–80%), arthropods maintain a relatively low, but stable, percentage.

There is no doubt that trilobites dominated early Cambrian benthic communities in terms of diversity and abundance, which is demonstrated well in the older Chengjiang Lagerstätte (Zhao et al. 2010; Hou et al. 2017; Paterson et al. 2019) and the Malong Fauna (Luo et al. 2008; Chen et al. 2019). The latter is characterized by extremely abundant and diverse trilobites yielding from the underlying Hongjingshao Formation (Fig. S1; Table S1). However, detailed fossil data from the Guanshan Biota in Wuding and Malong areas reveals a community structure that is unique for early Cambrian Konservat-Lagerstätten, with brachiopods dominating the benthic community in abundance, if not diversity, and often forming distinctive concentrations of shell beds in the lower Cambrian Stage 4 of the Wulongqing Formation (Figs 2 and 7a; Fig. S1). The ecological transition from trilobite- to brachiopod-dominated communities occurs widely across shallow marine clastic environments across the South China Platform (Fig. 1), coinciding with well-documented transgression events during Cambrian Age 4 (Luo et al. 2008; Hu et al. 2010, 2013; Chen et al. 2019). Thus organophosphatic brachiopods diversify and become superabundant across the broad ‘shallow’ shelf of the Yangtze Platform during the final stage of the Cambrian Explosion (Z.F. Zhang et al. 2020a). The rise of organophosphatic brachiopods as the numerically dominant element in the lower Cambrian Stage 4 Wulongqing Formation is the oldest brachiopod-dominated soft substrate community known in the fossil record and represents a precursor to more complex community tiering and brachiopod-dominant benthic communities during the Great Ordovician Biodiversification Event (Bassett et al. 2002; Servais and Harper 2018; Topper et al. 2018; Z.L. Zhang et al. 2018, 2020b; Z.F. Zhang et al. 2020a).

The brachiopods recovered from the section include lingulides (Eoobolus, Neobolus and Westonia), an acrotretide (Linnarssonia) and calcareous kutorginides (Kutorgina and Nisusia) (Table 1). Lingulides occur in high abundance and also form many shell concentrations within several assemblages (Fig. 2). The number of brachiopod concentrations (at least ten thin mud beds) far exceeds those produced by trilobites (only one mud bed). The composition of brachiopod taxa within each assemblage shows a rapid transition through time (Figs 2 and 7b). Neobolus is predominant in the lower part of the section (assemblages A–C, E and F), with Eoobolus (lingulides) and acrotretides common, but subordinate (Figs 2 and 7b). The relative abundance of the acrotheloid brachiopods, earlier referred to as ‘Westonia’ gubaiensis, increases gradually up-section, replacing, in part, the lingulides (Eoobolus and Neobolus) and acrotretides. This is partly attributed to the fact that the brachiopods of Eoobolus and Linnarssonia had a much smaller shell (c. 2–5 mm in maximum length) than Westonia. In addition, Westonia has a very wide and circular shell in outline, which is potentially adapted to the shallowing seawater environment. In general, the linguliform (e.g. lingulides and acrotretides) brachiopods show a strong control on assemblage dominance, whereas calcareous forms (kutorginides) remain rare (Fig. 7b).

Fossil concentrations, although common throughout geological time (Li and Droser 1997; Damborenea and Lanés 2007; Mancosu et al. 2015; El-Sabbagh and El Hedeny 2016; García-Ramos and Zuschin 2019), are rarely reported from Burgess Shale-type Lagerstätten (Han et al. 2006). The dominance of brachiopods within the Guanshan Biota, compared with other Cambrian Lagerstätten, is unique (Luo et al. 2008; Zhao et al. 2010; Paterson et al. 2016; Strang et al. 2016; Fu et al. 2019). The in situ preserved brachiopod concentrations in the Wuding area also occur in the Malong and Kunming areas, which indicates a wide geographical distribution (c. 6000 km2) after the rapid transgression at the base of the Wulongqing Formation (Hu et al. 2013; Chen et al. 2019; Z.F. Zhang et al. 2020a).

Overall, the fossil data show that brachiopods quickly replaced arthropods as the dominant fauna following a transgression that led to the deposition of the Wulongqing Formation at Wuding (Figs 2 and 7a; Fig. S1). Different brachiopod genera dominated different assemblages and, in places, formed distinctive shell concentrations (Figs 2 and 7b).

Guanshan Biota and its environment

The Guanshan Biota is an exceptionally preserved Konservat-Lagerstätte, uniquely characterized by brachiopod-dominated early Cambrian communities, substantially different from the arthropod-dominated Konservat-Lagerstätten such as the Chengjiang and Burgess Shale biotas (Zhao et al. 2010; O'Brien and Caron 2016; Hou et al. 2017; Nanglu et al. 2020). Although the preservation of soft tissues within biomineralized and sclerotized exoskeletons is common, which is at least partly attributable to the high number of brachiopods, trilobites and hyoliths, completely soft-bodied organisms (e.g. ctenophores) are absent in the Shijiangjun section, which is similar to the Ordovician Fezouata Biota (Saleh et al. 2020). This phenomenon is possibly related to preservation bias because the brachiopods, trilobites and hyoliths are more resistant to decay and much more readily preserved within this Konservat-Lagerstätte, which might lead to an underestimation of the diversity (Saleh et al. 2020) of the Guanshan Biota in the Wuding area. The lack of completely soft-bodied taxa may be due to the lack of an exaerobic preservational trap that typifies the Burgess Shale-type deposits (Gaines and Droser 2005).

The relatively shallow sedimentary environment (lower offshore in Martin et al. 2016 or offshore in Buatois and Mángano 2011) of the Guanshan Biota also separates it from most other Cambrian Lagerstätten worldwide except, perhaps, for the early Cambrian Emu Bay Shale from Australia, which is interpreted to have been deposited in a nearshore micro-basin setting adjacent to an active tectonic margin that generated continual syndepositional faulting and slumping (Paterson et al. 2016). The Guanshan Biota is also comparable with the Ordovician Fezouata Biota, both in terms of depositional environment and shelly faunal composition (Van Roy et al. 2015; Saleh et al. 2018). The latter was deposited mainly in an offshore to lower shoreface setting (Martin et al. 2016).

Gravity, traction and turbiditic flows are responsible for the transitions from arthropod- to brachiopod-dominated assemblages from the lower part of the Wulongqing Formation at the Shijiangjun section. The depositional environment between the fair weather wave base and the storm wave base is usually affected by frequent event flows, such as oscillatory and gravity flows (Kooi and Groen 2001; Majid et al. 2017), which helps to mix oxygen-enriched surface water with stagnant bottom water, providing favourable nutrient-rich conditions for the development of the benthic community (X. Liu et al. 2012b). Transportation from a nearby source, rapid fall out from suspension and the resuspension of seston provides a high nutrient load for suspension feeders such as brachiopods to flourish.

The limited amount of bioturbation throughout most of the section seems to indicate conditions unfavourable for burrowing, resulting from high turbidity, high or low salinity, or the relatively low oxygen content, perhaps explaining the dominance of relatively small, physiological simple filter-feeding brachiopods. The increase in the bioturbation index in the middle part of the section (3–6 m above the basal contact) is coincident with assemblages G–I, indicating more favourable conditions, probably a result of the relatively shallower depositional environment or fluctuating oxic conditions (Gaines and Droser 2005). The frequent overturn of fossil assemblages, especially brachiopods, may be attributed to frequent environmental fluctuations and the episodic input of coarser sediments, which probably periodically interrupt the benthic suspension assemblages.

Detailed analysis of the sedimentology, lithology and structures facilitates the identification of distinct lithofacies associated with transgressive systems tracts that directly affected the composition, diversity and relative abundance of faunal assemblages in the transition from the Hongjingshao to the Wulongqing deposits (Luo et al. 2008; Chen et al. 2019). Microfacies analysis, the degree of bioturbation and the faunal composition at the lower part of the Wulongqing Formation provide a new understanding of how fluctuations in the depositional environment influenced the faunal overturn in the Guanshan Biota across the Yangtze Platform in eastern Yunnan.

Conclusions

This is the first detailed report of the lithofacies, depositional environments and associated relative faunal abundance in the Cambrian Age 4 Guanshan Biota. The new Shijiangjun section through the basal part of the Wulongqing Formation in the Wuding area, eastern Yunnan reveals fossil assemblages composed of six bilaterian groups (Brachiopoda, Arthropoda, Hyolitha, Priapulida, Vetulicola and Anomalocaridiids). Detailed sedimentological, lithological and ichnological characteristics of the section indicate that: (1) hydrodynamic conditions are fluctuating, with episodic changes in energy and current regimes producing periodically coarse sand beds (Facies 3); (2) the sediments are derived from a relatively nearby source and accumulated rapidly; (3) the environment is affected by multi-period hydrodynamic events, such as storm and gravity flows forming obrution deposits (Zhang et al. 2019); and (4) the overall sedimentary environment in the Wuding area represents a deeper offshore to lower shoreface than the Wulongqing Formation outcropping in the Malong and Kunming areas.

The community transitioned from arthropod- to brachiopod-dominated for the first time at the base of the Wulongqing Formation in the Shijiangjun section. Within the brachiopod communities, a lingulate-dominated assemblage transitioned to an acrotheloid-dominated assemblage with the new occurrence of calcareous kutorginides up-section. The detailed study and documentation of this transition provides a better understanding of the differences in faunal composition and overturn between the Malong Fauna and Guanshan Biota (Luo et al. 2008; Chen et al. 2019). The unstable sedimentary environment with periodically sandy depositional inputs and muddy obrution deposits is probably closely associated with the observed succession of community assemblages. Brachiopods from the Guanshan Biota generally show a preference for such a fluctuating environment and adapt well to this environmental setting during the final stage of Cambrian evolutionary radiation.

Acknowledgements

The authors thank David Mathieson for helpful suggestions and Yue Liang, Yazhou Hu and Xiaolin Duan from Northwest University for constructive discussions and help in the field. Farid Saleh and one anonymous reviewer and Editor Xiaoya Ma are acknowledged for their helpful comments, which greatly improved this paper.

Author contributions

FC: conceptualization (equal), data curation (equal), formal analysis (equal), funding acquisition (equal), investigation (equal), methodology (equal), writing – original draft (lead), writing – review and editing (equal); GAB: conceptualization (equal), data curation (equal), funding acquisition (equal), methodology (equal), supervision (equal), writing – review and editing (equal); Z-LZ: conceptualization (equal), conceptualization (equal), data curation (equal), data curation (equal), formal analysis (equal), funding acquisition (equal), funding acquisition (equal), investigation (equal), investigation (equal), methodology (equal), methodology (equal), supervision (equal), writing – original draft (supporting), writing – review and editing (equal), writing – review and editing (equal); BL: conceptualization (equal), formal analysis (equal), methodology (equal), writing – original draft (supporting), writing – review and editing (equal); XR: investigation (equal), methodology (equal), writing – review and editing (supporting); Z-FZ: conceptualization (equal), conceptualization (equal), data curation (equal), data curation (equal), formal analysis (equal), funding acquisition (equal), funding acquisition (equal), investigation (equal), investigation (equal), methodology (equal), methodology (equal), supervision (equal), writing – original draft (supporting), writing – review and editing (equal), writing – review and editing (equal).

Funding

This work represents a contribution to the programs supported by the National Natural Science Foundation of China (41425008, 41720104002, 41621003 and 41890844), the Strategic Priority Research Program of the Chinese Academy of Sciences and the 111 project (D17013), the 1000 Talent Shaanxi Province Fellowship (GAB), the Macquarie University Research Fellowships to Z-LZ (2019 MQRF) and iMQRES from Macquarie University to BL. FC warmly acknowledges the China Scholarship Council (CSC 201806970026) for 14 months of research as an Exchange PhD student with GAB at Macquarie University.

Data availability statement

All data generated or analysed during this study are included in this published article (and its supplementary information files).

Conflicts of interest

The authors declare no known conflicts of interest associated with this publication.

Scientific editing by Xiaoya Ma

  • © 2020 The Author(s). Published by The Geological Society of London
http://creativecommons.org/licenses/by/4.0/

This is an Open Access article distributed under the terms of the Creative Commons Attribution 4.0 License (http://creativecommons.org/licenses/by/4.0/)

References

  1. ↵
    1. Amorosi, A.
    1997. Detecting compositional, spatial, and temporal attributes of glaucony: a tool for provenance research. Sedimentary Geology, 109, 135–153, https://doi.org/10.1016/S0037-0738(96)00042-5
    OpenUrlCrossRefWeb of Science
  2. ↵
    1. Amorosi, A.,
    2. Guidi, R.,
    3. Mas, R. and
    4. Falanga, E.
    2012. Glaucony from the Cretaceous of the Sierra de Guadarrama (Central Spain) and its application in a sequence-stratigraphic context. International Journal of Earth Sciences, 101, 415–427, https://doi.org/10.1007/s00531-011-0675-x
    OpenUrl
  3. ↵
    1. Aria, C. and
    2. Caron, J.B.
    2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature, 545, 89–92, https://doi.org/10.1038/nature22080
    OpenUrl
  4. ↵
    1. Arnott, R.W. and
    2. Southard, J.B.
    1990. Exploratory flow-duct experiments on combined-flow bed configurations, and some implications for interpreting storm-event stratification. Journal of Sedimentary Research, 60, 211–219, https://doi.org/10.1306/212F9156-2B24-11D7-8648000102C1865D.
    OpenUrlAbstract/FREE Full Text
  5. ↵
    1. Baioumy, H. and
    2. Boulis, S.
    2012. Glauconites from the Bahariya Oasis: an evidence for Cenomanian marine transgression in Egypt. Journal of African Earth Sciences, 70, 1–7, https://doi.org/10.1016/j.jafrearsci.2012.05.001
    OpenUrlCrossRefWeb of Science
  6. ↵
    1. Banerjee, S.,
    2. Chattoraj, S.L.,
    3. Saraswati, P.,
    4. Dasgupta, S.,
    5. Sarkar, U. and
    6. Bumby, A.
    2012. The origin and maturation of lagoonal glauconites: a case study from the Oligocene Maniyara Fort Formation, western Kutch, India. Geological Journal, 47, 357–371, https://doi.org/10.1002/gj.1345
    OpenUrl
  7. ↵
    1. Banerjee, S.,
    2. Bansal, U. and
    3. Thorat, A.V.
    2017. A review on palaeogeographic implications and temporal variation in glaucony composition. Journal of Palaeogeography, 5, 43–71, https://doi.org/10.1016/j.jop.2015.12.001
    OpenUrl
  8. ↵
    1. Bassett, M.G.,
    2. Popov, L.E. and
    3. Holmer, L.E.
    2002. Brachiopods: Cambrian–Tremadoc precursors to Ordovician radiation events. Geological Society, London, Special Publications, 194, 13–23, https://doi.org/10.1144/GSL.SP.2002.194.01.02
  9. ↵
    1. Bouma, A.H.
    1962. Sedimentology of Some Flysch Deposits. A Graphic Approach to Facies Interpretation. Elsevier.
  10. ↵
    1. Buatois, L.A. and
    2. Mángano, M.G.
    2011. Ichnology: Organism–Substrate Interactions in Space and Time. Cambridge University Press.
  11. ↵
    1. Buatois, L.A.,
    2. Santiago, N.,
    3. Herrera, M.,
    4. Plink-Björklund, P.,
    5. Steel, R.,
    6. Espin, M. and
    7. Parra, K.
    2012. Sedimentological and ichnological signatures of changes in wave, river and tidal influence along a Neogene tropical deltaic shoreline. Sedimentology, 59, 1568–1612, https://doi.org/10.1111/j.1365-3091.2011.01317.x
    OpenUrlCrossRefWeb of Science
  12. ↵
    1. Bullimore, S.A.,
    2. Helland-Hansen, W.,
    3. Henriksen, S. and
    4. Steel, R.J.
    2008. Shoreline trajectory and its impact on coastal depositional environments: an example from the Upper Cretaceous Mesaverde Group, NW Colorado, USA. SEPM, Special Publications, 90, 209–236, https://doi.org/10.2110/pec.08.90.0209
  13. ↵
    1. Cao, Y.M.,
    2. Curran, A.H. and
    3. Glumac, B.
    2015. Testing the use of Photoshop and ImageJ for evaluating ichnofabrics. Geological Society of America, Abstracts with Programs, 47, 343.
    OpenUrl
  14. ↵
    1. Chafetz, H.S.
    2007. Paragenesis of the Morgan Creek Limestone, Late Cambrian, central Texas: constraints on the formation of glauconite. Deep Sea Research Part II: Topical Studies in Oceanography, 54, 1350–1363, https://doi.org/10.1016/j.dsr2.2007.04.002
    OpenUrl
  15. ↵
    1. Chafetz, H. and
    2. Reid, A.
    2000. Syndepositional shallow-water precipitation of glauconitic minerals. Sedimentary Geology, 136, 29–42, https://doi.org/10.1016/S0037-0738(00)00082-8
    OpenUrlCrossRefWeb of Science
  16. ↵
    1. Cheel, R.J.
    1990. Horizontal lamination and the sequence of bed phases and stratification under upper-flow-regime conditions. Sedimentology, 37, 517–529, https://doi.org/10.1111/j.1365-3091.1990.tb00151.x
    OpenUrlCrossRefWeb of Science
  17. ↵
    1. Cheel, R.J. and
    2. Leckie, D.A.
    1993. Hummocky cross-stratification. Sedimentology Review, 1, 103–122, https://doi.org/10.1002/9781444304534.ch7
    OpenUrl
  18. ↵
    1. Chen, F.,
    2. Zhang, Z.,
    3. Betts, J.M.,
    4. Zhang, Z. and
    5. Liu, F.
    2019. First report on Guanshan Biota (Cambrian Stage 4) at the stratotype area of Wulongqing Formation in Malong County, Eastern Yunnan, China. Geoscience Frontiers, 10, 1459–1476, https://doi.org/10.1016/j.gsf.2018.09.010
    OpenUrl
  19. ↵
    1. Coe, A.,
    2. Bosence, D.,
    3. Church, K.,
    4. Flint, S.,
    5. Howell, J. and
    6. Wilson, R.
    2003. The sedimentary record of sea-level change. Cambridge University Press.
  20. ↵
    1. Collom, C.,
    2. Johnston, P. and
    3. Powell, W.
    2009. Reinterpretation of ‘Middle’ Cambrian stratigraphy of the rifted western Laurentian margin: Burgess Shale Formation and contiguous units (Sauk II megasequence), Rocky Mountains, Canada. Palaeogeography, Palaeoclimatology, Palaeoecology, 277, 63–85, https://doi.org/10.1016/j.palaeo.2009.02.012
    OpenUrlCrossRef
  21. ↵
    1. Damborenea, S. and
    2. Lanés, S.
    2007. Early Jurassic shell beds from marginal marine environments in southern Mendoza, Argentina. Palaeogeography, Palaeoclimatology, Palaeoecology, 250, 68–88, https://doi.org/10.1016/j.palaeo.2007.03.002
    OpenUrl
  22. ↵
    1. Delamette, M.
    1989. Trace fossil assemblages from the Albian phosphate-rich sandstones of the Helvetic Shelf (western Alps). Cretaceous Research, 10, 207–219, https://doi.org/10.1016/0195-6671(89)90018-9
    OpenUrl
  23. ↵
    1. Dorador, J. and
    2. Rodríguez-Tovar, F.J.
    2018. High-resolution image treatment in ichnological core analysis: initial steps, advances and prospects. Earth-Science Reviews, 177, 226–237, https://doi.org/10.1016/j.earscirev.2017.11.020
    OpenUrl
  24. ↵
    1. Dorador, J.,
    2. Rodríguez-Tovar, F.J. and
    3. IODP Expedition 339 Scientists
    2014. Digital image treatment applied to ichnological analysis of marine core sediments. Facies, 60, 39–44, https://doi.org/10.1007/s10347-013-0383-z
    OpenUrlCrossRefWeb of Science
  25. ↵
    1. Dott, R. Jr. and
    2. Bourgeois, J.
    1982. Hummocky stratification: significance of its variable bedding sequences. Geological Society of America Bulletin, 93, 663–680, https://doi.org/10.1130/0016-7606(1982)93<663:HSSOIV>2.0.CO;2
    OpenUrlAbstract/FREE Full Text
  26. ↵
    1. Duan, X.,
    2. Liang, Y.,
    3. Holmer, L.E. and
    4. Zhang, Z.
    2020. First report of acrotretoid brachiopod shell beds in the lower Cambrian (Stage 4) Guanshan Biota of eastern Yunnan, South China. Journal of Paleontology, corrected proof online September 9, 2020, https://doi.org/10.1017/jpa.2020.66
  27. ↵
    1. Duke, W.L.,
    2. Arnott, R. and
    3. Cheel, R.J.
    1991. Shelf sandstones and hummocky cross-stratification: new insights on a stormy debate. Geology, 19, 625–628, https://doi.org/10.1130/0091-7613(1991)019<0625:SSAHCS>2.3.CO;2
    OpenUrlAbstract/FREE Full Text
  28. ↵
    1. Dumas, S. and
    2. Arnott, R.
    2006. Origin of hummocky and swaley cross-stratification—the controlling influence of unidirectional current strength and aggradation rate. Geology, 34, 1073–1076, https://doi.org/10.1130/G22930A.1
    OpenUrlAbstract/FREE Full Text
  29. ↵
    1. Dumas, S.,
    2. Arnott, R. and
    3. Southard, J.B.
    2005. Experiments on oscillatory-flow and combined-flow bed forms: implications for interpreting parts of the shallow-marine sedimentary record. Journal of Sedimentary Research, 75, 501–513, https://doi.org/10.2110/jsr.2005.039
    OpenUrlAbstract/FREE Full Text
  30. ↵
    1. Eide, C.H.,
    2. Howell, J.A. and
    3. Buckley, S.J.
    2015. Sedimentology and reservoir properties of tabular and erosive offshore transition deposits in wave-dominated, shallow-marine strata: Book Cliffs, USA. Petroleum Geoscience, 21, 55–73, https://doi.org/10.1144/petgeo2014-015
    OpenUrl
  31. ↵
    1. Elliott, T.
    1978. Deltas. In: Reading, H.G. (ed.) Sedimentary Environments and Facies. 2nd edn. Blackwell Scientific, 113–154.
  32. ↵
    1. El-Sabbagh, A.M. and
    2. El Hedeny, M.M.
    2016. A shell concentration of the middle Miocene Crassostrea gryphoides (Schlotheim, 1813) from Siwa oasis, Western Desert, Egypt. Journal of African Earth Sciences, 120, 1–11, https://doi.org/10.1016/j.jafrearsci.2016.04.007
    OpenUrl
  33. ↵
    1. Fu, D.,
    2. Tong, G. et al.
    2019. The Qingjiang biota—a Burgess Shale-type fossil lagerstätte from the early Cambrian of South China. Science, 363, 1338–1342, https://doi.org/10.1126/science.aau8800
    OpenUrlAbstract/FREE Full Text
  34. ↵
    1. Fürsich, F.,
    2. Oschmann, W.,
    3. Singh, I. and
    4. Jaitly, A.
    1992. Hardgrounds, reworked concretion levels and condensed horizons in the Jurassic of western India: their significance for basin analysis. Journal of the Geological Society, London, 149, 313–331, https://doi.org/10.1144/gsjgs.149.3.0313
    OpenUrlAbstract/FREE Full Text
  35. ↵
    1. Gaines, R.R. and
    2. Droser, M.L.
    2005. New approaches to understanding the mechanics of Burgess Shale-type deposits: from the micron scale to the global picture. The Sedimentary Record, 3, 4–8, http://www.sepm.org/pages.aspx?pageid=37, https://doi.org/10.2110/sedred.2005.2.4
    OpenUrl
  36. ↵
    1. García-Ramos, D.A. and
    2. Zuschin, M.
    2019. High-frequency cycles of brachiopod shell beds on subaqueous delta-scale clinoforms (early Pliocene, south-east Spain). Sedimentology, 66, 1486–1530, https://doi.org/10.1111/sed.12541
    OpenUrl
  37. ↵
    1. Garzanti, E.,
    2. Haas, R. and
    3. Jadoul, F.
    1989. Ironstones in the Mesozoic passive margin sequence of the Tethys Himalaya (Zanskar, Northern India): sedimentology and metamorphism. Geological Society, London, Special Publications, 46, 229–244, https://doi.org/10.1144/GSL.SP.1989.046.01.20
  38. ↵
    1. Gougeon, R.C.,
    2. Mángano, M.G.,
    3. Buatois, L.A.,
    4. Narbonne, G.M. and
    5. Laing, B.A.
    2018. Early Cambrian origin of the shelf sediment mixed layer. Nature Communications, 9, 1–7, https://doi.org/10.1038/s41467-018-04311-8
    OpenUrl
  39. ↵
    1. Han, J.,
    2. Shu, D.,
    3. Zhang, Z.,
    4. Liu, J.,
    5. Zhang, X. and
    6. Ya, Y.
    2006. Preliminary notes on soft-bodied fossil concentrations from the Early Cambrian Chengjiang deposits. Chinese Science Bulletin, 51, 2482–2492, https://doi.org/10.1007/s11434-005-2151-0
    OpenUrlCrossRefWeb of Science
  40. ↵
    1. Henstra, G.A.,
    2. Grundvåg, S.-A. et al.
    2016. Depositional processes and stratigraphic architecture within a coarse-grained rift-margin turbidite system: the Wollaston Forland Group, east Greenland. Marine and Petroleum Geology, 76, 187–209, https://doi.org/10.1016/j.marpetgeo.2016.05.018
    OpenUrl
  41. ↵
    1. Hopkins, M.J.,
    2. Chen, F.,
    3. Hu, S. and
    4. Zhang, Z.
    2017. The oldest known digestive system consisting of both paired digestive glands and a crop from exceptionally preserved trilobites of the Guanshan Biota (Early Cambrian, China). PloS One, 12, e0184982, https://doi.org/10.1371/journal.pone.0184982
    OpenUrl
  42. ↵
    1. Hou, X.,
    2. Siveter, D.J. et al.
    2017. The Cambrian Fossils of Chengjiang, China: the Flowering of Early Animal Life. 2nd edn. Blackwell, https://doi.org/10.1002/9780470999950
  43. ↵
    1. Hu, S.,
    2. Zhu, M.,
    3. Steiner, M.,
    4. Luo, H.,
    5. Zhao, F. and
    6. Liu, Q.
    2010. Biodiversity and taphonomy of the Early Cambrian Guanshan biota, eastern Yunnan. Science China Earth Sciences, 53, 1765–1773, https://doi.org/10.1007/s11430-010-4086-9
    OpenUrl
  44. ↵
    1. Hu, S.,
    2. Zhu, M. et al.
    2013. The Guanshan Biota. Yunnan Science and Technology Press.
  45. ↵
    1. Ivantsov, A.Y.,
    2. Zhuravlev, A.Y.,
    3. Leguta, A.V.,
    4. Krassilov, V.A.,
    5. Melnikova, L.M. and
    6. Ushatinskaya, G.T.
    2005. Palaeoecology of the early Cambrian Sinsk biota from the Siberian platform. Palaeogeography, Palaeoclimatology, Palaeoecology, 220, 69–88, https://doi.org/10.1016/j.palaeo.2004.01.022
    OpenUrlCrossRefWeb of Science
  46. ↵
    1. Kneller, B.
    1995. Beyond the turbidite paradigm: physical models for deposition of turbidites and their implications for reservoir prediction. Geological Society, London, Special Publications, 94, 31–49, https://doi.org/10.1144/GSL.SP.1995.094.01.04
  47. ↵
    1. Kooi, H. and
    2. Groen, J.
    2001. Offshore continuation of coastal groundwater systems; predictions using sharp-interface approximations and variable-density flow modelling. Journal of Hydrology, 246, 19–35, https://doi.org/10.1016/S0022-1694(01)00354-7
    OpenUrlCrossRefWeb of Science
  48. ↵
    1. Li, X. and
    2. Droser, M.L.
    1997. Nature and distribution of Cambrian shell concentrations; evidence from the Basin and Range Province of the Western United States (California, Nevada, and Utah). Palaios, 12, 111–126, https://doi.org/10.2307/3515301
    OpenUrlAbstract/FREE Full Text
  49. ↵
    1. Li, J.,
    2. Liu, J. and
    3. Ou, Q.
    2017. New observations on Vetulicola longbaoshanensis from the lower Cambrian Guanshan Biota (series 2, stage 4), South China. Science China Earth Sciences, 60, 1795–1804, https://doi.org/10.1007/s11430-017-9088-y
    OpenUrl
  50. ↵
    1. Liu, J.,
    2. Ou, Q. et al.
    2012a. New occurrence of the Cambrian (Stage 4, Series 2) Guanshan Biota in Huize, Yunnan, South China. Bulletin of Geosciences, 87, 125–132, https://doi.org/10.3140/bull.geosci.1229
    OpenUrl
  51. ↵
    1. Liu, X.,
    2. Zhong, J.,
    3. Grapes, R.,
    4. Bian, S. and
    5. Liang, C.
    2012b. Late Cretaceous tempestite in northern Songliao Basin, China. Journal of Asian Earth Sciences, 56, 33–41, https://doi.org/10.1016/j.jseaes.2012.02.007
    OpenUrl
  52. ↵
    1. Liu, J.,
    2. Han, J. et al.
    2016. New localities and palaeoscolecid worms from the Cambrian (Stage 4, Series 2) Guanshan Biota in Kunming, Yunnan, South China. Acta Geologica Sinica – English Edition, 90, 1939–1945, https://doi.org/10.1111/1755-6724.13013
    OpenUrl
  53. ↵
    1. Liu, F.,
    2. Skovsted, C.B.,
    3. Topper, T.P.,
    4. Zhang, Z.F. and
    5. Shu, D.G.
    2020. Are hyoliths Palaeozoic lophophorates? National Science Review, 7, 453–469, https://doi.org/10.1093/nsr/nwz161
    OpenUrl
  54. ↵
    1. Luo, H.L.,
    2. Li, Y.,
    3. Hu, S.X.,
    4. Fu, X.P.,
    5. Hou, S.G. and
    6. Liu, X.R.
    2008. Early Cambrian Malong Fauna and Guanshan Fauna from Eastern Yunnan, China. Yunnan Science and Technology Press [in Chinese with English summary].
  55. ↵
    1. Maceachern, J.A.,
    2. Stelck, C.R. and
    3. Pemberton, S.G.
    1999. Marine and Marginal Marine Mudstone Deposition: Paleoenvironmental Interpretations Based on the Integration of Ichnology, Palynology and Forarniniferal Paleoecology. Special Publication-SEPM, 4, 205–225.
    OpenUrl
  56. ↵
    1. Majid, M.F.A.,
    2. Ismail, M.S.,
    3. Rahman, A.H.A. and
    4. Mohamed, M.A.
    2017. Facies distribution and petrophysical properties of shoreface-offshore transition environment in Sandakan Formation, NE Sabah Basin. IOP Conference Series: Earth and Environmental Science, 88, 012023, https://doi.org/10.1088/1755-1315/88/1/012023
    OpenUrl
  57. ↵
    1. Mancosu, A.,
    2. Nebelsick, J.H.,
    3. Kroh, A. and
    4. Pillola, G.L.
    2015. The origin of echinoid shell beds in siliciclastic shelf environments: three examples from the Miocene of Sardinia, Italy. Lethaia, 48, 83–99, https://doi.org/10.1111/let.12090
    OpenUrl
  58. ↵
    1. Martin, E.L.,
    2. Pittet, B. et al.
    2016. The Lower Ordovician Fezouata Konservat-Lagerstätte from Morocco: age, environment and evolutionary perspectives. Gondwana Research, 34, 274–283, https://doi.org/10.1016/j.gr.2015.03.009
    OpenUrl
  59. ↵
    1. Mcrae, S.
    1972. Glauconite. Earth-Science Reviews, 8, 397–440, https://doi.org/10.1016/0012-8252(72)90063-3
    OpenUrl
  60. ↵
    1. Meldahl, K.H.
    1993. Geographic gradients in the formation of shell concentrations: Plio-Pleistocene marine deposits, Gulf of California. Palaeogeography, Palaeoclimatology, Palaeoecology, 101, 1–25, https://doi.org/10.1016/0031-0182(93)90149-D
    OpenUrlCrossRefWeb of Science
  61. ↵
    1. Michalík, J.,
    2. Lintnerová, O. et al.
    2013. Paleoenvironments during the Rhaetian transgression and the colonization history of marine biota in the Fatric Unit (Western Carpathians). Geologica Carpathica, 64, 39–62, https://doi.org/10.2478/geoca-2013-0003
    OpenUrl
  62. ↵
    1. Myrow, P.M. and
    2. Southard, J.B.
    1996. Tempestite deposition. Journal of Sedimentary Research, 66, 875–887, https://doi.org/10.1306/D426842D-2B26-11D7-8648000102C1865D.
    OpenUrlAbstract/FREE Full Text
  63. ↵
    1. Nanglu, K.,
    2. Caron, J.B. and
    3. Gaines, R.R.
    2020. The Burgess Shale paleocommunity with new insights from Marble Canyon, British Columbia. Paleobiology, 46, 58–81, https://doi.org/10.1017/pab.2019.42
    OpenUrl
  64. ↵
    1. O'Brien, L.J. and
    2. Caron, J.B.
    2016. Paleocommunity analysis of the Burgess Shale Tulip Beds, Mount Stephen, British Columbia: comparison with the Walcott Quarry and implications for community variation in the Burgess Shale. Paleobiology, 42, 27–53, https://doi.org/10.1017/pab.2015.17
    OpenUrlAbstract/FREE Full Text
  65. ↵
    1. Paterson, J.R.,
    2. García-Bellido, D.C.,
    3. Lee, M.S.,
    4. Brock, G.A.,
    5. Jago, J.B. and
    6. Edgecombe, G.D.
    2011. Acute vision in the giant Cambrian predator Anomalocaris and the origin of compound eyes. Nature, 480, 237–240, https://doi.org/10.1038/nature10689
    OpenUrlCrossRefPubMedWeb of Science
  66. ↵
    1. Paterson, J.R.,
    2. García-Bellido, D.C.,
    3. Jago, J.B.,
    4. Gehling, J.G.,
    5. Lee, M.S. and
    6. Edgecombe, G.D.
    2016. The Emu Bay Shale Konservat-Lagerstätte: a view of Cambrian life from East Gondwana. Journal of the Geological Society, London, 173, 1–11, https://doi.org/10.1144/jgs2015-083
    OpenUrlCrossRef
  67. ↵
    1. Paterson, J.R.,
    2. Edgecombe, G.D. and
    3. Lee, M.S.
    2019. Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences, USA, 116, 4394–4399, https://doi.org/10.1073/pnas.1819366116
    OpenUrlAbstract/FREE Full Text
  68. ↵
    1. Peel, J.S. and
    2. Ineson, J.R.
    2011. The extent of the Sirius Passet Lagerstätte (early Cambrian) of North Greenland. Bulletin of Geosciences, 86, 535–543, https://doi.org/10.3140/bull.geosci.1269
    OpenUrl
  69. ↵
    1. Peng, J.,
    2. Zhao, Y.,
    3. Wu, Y.,
    4. Yuan, J. and
    5. Tai, T.
    2005. The Balang Fauna—a new early Cambrian fauna from Kaili City, Guizhou Province. Chinese Science Bulletin, 50, 1159–1162, https://doi.org/10.1360/982005-183
    OpenUrlCrossRefWeb of Science
  70. ↵
    1. Rudmin, M.,
    2. Banerjee, S. and
    3. Mazurov, A.
    2017. Compositional variation of glauconites in Upper Cretaceous–Paleogene sedimentary iron-ore deposits in south-eastern Western Siberia. Sedimentary Geology, 355, 20–30, https://doi.org/10.1016/j.sedgeo.2017.04.006
    OpenUrl
  71. ↵
    1. Saito, Y.
    1989. Modern storm deposits in the inner shelf and their recurrence intervals, Sendai Bay, northeast Japan. In: Taira, A. and Masuda, F. (eds) Sedimentary Facies in the Active Plate Margin. Terra Scientific, 331–344.
  72. ↵
    1. Saleh, F.,
    2. Candela, Y.,
    3. Harper, D.A.,
    4. Polechová, M.,
    5. Lefebvre, B. and
    6. Pittet, B.
    2018. Storm-induced community dynamics in the Fezouata Biota (Lower Ordovician, Morocco). Palaios, 33, 535–541, https://doi.org/10.2110/palo.2018.055
    OpenUrl
  73. ↵
    1. Saleh, F.,
    2. Antcliffe, J.B. et al.
    2020. Taphonomic bias in exceptionally preserved biotas. Earth and Planetary Science Letters, 529, 115873, https://doi.org/10.1016/j.epsl.2019.115873
    OpenUrl
  74. ↵
    1. Servais, T. and
    2. Harper, D.A.
    2018. The great Ordovician biodiversification event (GOBE): definition, concept and duration. Lethaia, 51, 151–164, https://doi.org/10.1111/let.12259
    OpenUrlCrossRef
  75. ↵
    1. Shanmugam, G.
    2002. Discussion on Mulder et al.(2001, Geo-Marine Letters 21: 86–93) Inversely graded turbidite sequences in the deep Mediterranean. A record of deposits from flood-generated turbidity currents? Geo-Marine Letters, 22, 108–111, https://doi.org/10.1007/s00367-002-0100-3
    OpenUrl
  76. ↵
    1. Smith, M.R.
    2015. A palaeoscolecid worm from the Burgess Shale. Palaeontology, 58, 973–979, https://doi.org/10.1111/pala.12210
    OpenUrl
  77. ↵
    1. Southard, J.B.,
    2. Lambie, J.M.,
    3. Federico, D.C.,
    4. Pile, H.T. and
    5. Weidman, C.R.
    1990. Experiments on bed configurations in fine sands under bidirectional purely oscillatory flow, and the origin of hummocky cross-stratification. Journal of Sedimentary Research, 60, 1–17, https://doi.org/10.1306/212F90F7-2B24-11D7-8648000102C1865D.
    OpenUrlAbstract/FREE Full Text
  78. ↵
    1. Stow, D.A.
    2005. Sedimentary Rocks in the Field: A Color Guide. Gulf Professional Publishing, https://doi.org/10.1016/j.pgeola.2010.07.004
  79. ↵
    1. Strang, K.M.,
    2. Armstrong, H.A.,
    3. Harper, D.A. and
    4. Trabucho-Alexandre, J.P.
    2016. The Sirius Passet Lagerstätte: silica death masking opens the window on the earliest matground community of the Cambrian explosion. Lethaia, 49, 631–643, https://doi.org/10.1111/let.12174
    OpenUrl
  80. ↵
    1. Swift, D.J.,
    2. Figueiredo, A.G.,
    3. Freeland, G. and
    4. Oertel, G.
    1983. Hummocky cross-stratification and megaripples; a geological double standard? Journal of Sedimentary Research, 53, 1295–1317, https://doi.org/10.1306/212F8369-2B24-11D7-8648000102C1865D.
    OpenUrlAbstract/FREE Full Text
  81. ↵
    1. Taylor, A.M. and
    2. Goldring, R.
    1993. Description and analysis of bioturbation and ichnofabric. Journal of the Geological Society, London, 150, 141–148, https://doi.org/10.1144/gsjgs.150.1.0141
    OpenUrlAbstract/FREE Full Text
  82. ↵
    1. Topper, T.P.,
    2. Zhang, Z.,
    3. Gutiérrez-Marco, J.C. and
    4. Harper, D.A.
    2018. The dawn of a dynasty: life strategies of Cambrian and Ordovician brachiopods. Lethaia, 51, 254–266, https://doi.org/10.1111/let.12229
    OpenUrl
  83. ↵
    1. Van Roy, P.,
    2. Briggs, D.E. and
    3. Gaines, R.R.
    2015. The Fezouata fossils of Morocco; an extraordinary record of marine life in the Early Ordovician. Journal of the Geological Society, London, 172, 541–549, https://doi.org/10.1144/jgs2015-017
    OpenUrl
  84. ↵
    1. Wu, P.,
    2. Liu, S.,
    3. He, B. and
    4. Dou, G.
    2016. Stratigraphic records of the dynamic uplift of the Emeishan large igneous province. International Geology Review, 58, 112–130, https://doi.org/10.1080/00206814.2015.1065515
    OpenUrl
  85. ↵
    1. Yang, Y.,
    2. Zhang, X.,
    3. Zhao, Y.,
    4. Qi, Y. and
    5. Cui, L.
    2018. New paleoscolecid worms from the early Cambrian north margin of the Yangtze Platform, South China. Journal of Paleontology, 92, 49–58, https://doi.org/10.1017/jpa.2017.50
    OpenUrl
  86. ↵
    1. Yokokawa, M.,
    2. Masuda, F.,
    3. Sakai, T.,
    4. Endo, N. and
    5. Kubo, Y.
    1999. Sedimentary structures generated in upper-flow-regime with sediment supply: antidune cross-stratification (HCS mimics) in a flume. Land-Sea Link in Asia. STA (JISTEC) & Geological Survey of Japan, 409–414.
  87. ↵
    1. Zavala, C.,
    2. Arcuri, M. and
    3. Blanco Valiente, L.
    2012. The importance of plant remains as diagnostic criteria for the recognition of ancient hyperpycnites. Revue de Paléobiologie, 11, 457–469.
    OpenUrl
  88. ↵
    1. Zhang, Z.L.,
    2. Zhang, Z.F.,
    3. Holmer, L.E. and
    4. Chen, F.Y.
    2018. Post-metamorphic allometry in the earliest acrotretoid brachiopods from the lower Cambrian (Series 2) of South China, and its implications. Palaeontology, 61, 183–207, https://doi.org/10.1111/pala.12333
    OpenUrl
  89. ↵
    1. Zhang, P.,
    2. Kuang, H.,
    3. Liu, Y.,
    4. Meng, Z.,
    5. Peng, N. and
    6. Xu, H.
    2019. Sedimentary characteristics and provenance of the basal conglomerate of the Late Jurassic–Early Cretaceous Jiaolai Basin, eastern China and their implications for the uplift of the Sulu Orogenic Belt. International Geology Review, 61, 521–538, https://doi.org/10.1080/00206814.2018.1437786
    OpenUrl
  90. ↵
    1. Zhang, Z.F.,
    2. Strotz, L.C. et al.
    2020a. An encrusting kleptoparasite-host interaction from the early Cambrian. Nature Communications, 11, 1–7, https://doi.org/10.1038/s41467-019-13993-7
    OpenUrl
  91. ↵
    1. Zhang, Z.L.,
    2. Chen, F.Y. and
    3. Zhang, Z.F.
    2020b. The earliest phosphatic-shelled brachiopods from the carbonates of South China—their diversification, ontogeny and distribution. Earth Science Frontiers, 27, 127–151, https://doi.org/10.13745/j.esf.sf.2020.6.4
    OpenUrl
  92. ↵
    1. Zhao, Y.,
    2. Zhu, M. et al.
    2005. Kaili Biota: a taphonomic window on diversification of metazoans from the basal Middle Cambrian: Guizhou, China. Acta Geologica Sinica – English Edition, 79, 751–765, https://doi.org/10.1111/j.1755-6724.2005.tb00928.x
    OpenUrl
  93. ↵
    1. Zhao, F.,
    2. Zhu, M. and
    3. Hu, S.
    2010. Community structure and composition of the Cambrian Chengjiang biota. Science China Earth Sciences, 53, 1784–1799, https://doi.org/10.1007/s11430-010-4087-8
    OpenUrl
  94. ↵
    1. Zhu, M.,
    2. Zhang, J. and
    3. Li, G.
    2001. Sedimentary environments of the early Cambrian Chengjiang biota: sedimentology of the Yu'anshan Formation in Chengjiang County, eastern Yunnan. The Cambrian of South China. Acta Palaeontologica Sinica, 40, 80–105.
    OpenUrl
View Abstract
PreviousNext
Back to top

In this issue

Journal of the Geological Society: 178 (1)
Journal of the Geological Society
Volume 178, Issue 1
January 2021
  • Table of Contents
  • About the Cover
  • Index by author
Alerts
Sign In to Email Alerts with your Email Address
Citation tools

Brachiopod-dominated communities and depositional environment of the Guanshan Konservat-Lagerstätte, eastern Yunnan, China

Feiyang Chen, Glenn A. Brock, Zhiliang Zhang, Brittany Laing, Xinyi Ren and Zhifei Zhang
Journal of the Geological Society, 178, jgs2020-043, 18 September 2020, https://doi.org/10.1144/jgs2020-043
Feiyang Chen
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Formal analysis (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Writing - Original Draft (Lead)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • ORCID record for Feiyang Chen
Glenn A. Brock
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • ORCID record for Glenn A. Brock
Zhiliang Zhang
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
2Department of Biological Sciences, Macquarie University, , , Australia
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • ORCID record for Zhiliang Zhang
Brittany Laing
2Department of Biological Sciences, Macquarie University, , , Australia
3Department of Geological Sciences, University of Saskatchewan, , , Canada
Roles: [Conceptualization (Equal)], [Formal analysis (Equal)], [Methodology (Equal)], [Writing - Original Draft (Supporting)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • ORCID record for Brittany Laing
Xinyi Ren
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
Roles: [Investigation (Equal)], [Methodology (Equal)], [Writing - Review & Editing (Supporting)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
Zhifei Zhang
1State Key Laboratory of Continental Dynamics, Shaanxi Key Laboratory of Early Life & Environments and Department of Geology, Northwest University, , , China
Roles: [Conceptualization (Equal)], [Data curation (Equal)], [Funding acquisition (Equal)], [Investigation (Equal)], [Methodology (Equal)], [Supervision (Equal)], [Writing - Review & Editing (Equal)]
  • Find this author on Google Scholar
  • Find this author on PubMed
  • Search for this author on this site
  • ORCID record for Zhifei Zhang
  • For correspondence: elizf@nwu.edu.cn zhangelle@126.com

Citation Manager Formats

  • BibTeX
  • Bookends
  • EasyBib
  • EndNote (tagged)
  • EndNote 8 (xml)
  • Medlars
  • Mendeley
  • Papers
  • RefWorks Tagged
  • Ref Manager
  • RIS
  • Zotero
Permissions
View PDF
Share

Brachiopod-dominated communities and depositional environment of the Guanshan Konservat-Lagerstätte, eastern Yunnan, China

Feiyang Chen, Glenn A. Brock, Zhiliang Zhang, Brittany Laing, Xinyi Ren and Zhifei Zhang
Journal of the Geological Society, 178, jgs2020-043, 18 September 2020, https://doi.org/10.1144/jgs2020-043
del.icio.us logo Digg logo Reddit logo Twitter logo CiteULike logo Facebook logo Google logo Mendeley logo
Email to

Thank you for sharing this Journal of the Geological Society article.

NOTE: We request your email address only to inform the recipient that it was you who recommended this article, and that it is not junk mail. We do not retain these email addresses.

Enter multiple addresses on separate lines or separate them with commas.
Brachiopod-dominated communities and depositional environment of the Guanshan Konservat-Lagerstätte, eastern Yunnan, China
(Your Name) has forwarded a page to you from Journal of the Geological Society
(Your Name) thought you would be interested in this article in Journal of the Geological Society.
CAPTCHA
This question is for testing whether or not you are a human visitor and to prevent automated spam submissions.
Print
Download PPT
  • Tweet Widget
  • Facebook Like
  • Google Plus One
  • Article
    • Abstract
    • Materials and methods
    • Results
    • Discussion
    • Conclusions
    • Acknowledgements
    • Author contributions
    • Funding
    • Data availability statement
    • Conflicts of interest
    • References
  • Figures & Data
  • Info & Metrics
  • PDF

Related Articles

Similar Articles

Cited By...

More in this TOC Section

  • The Xigaze ophiolite: fossil ultraslow-spreading ocean lithosphere in the Tibetan Plateau
  • Microfossil and strontium isotope chronology used to identify the controls of Miocene reefs and related facies in NW Cyprus
  • U–Pb isotopic ages and provenance of some far-travelled exotic pebbles from glaciogenic sediments of the Polonez Cove Formation (Oligocene, King George Island)
Show more: Research article
  • Most read
  • Most cited
Loading
  • Geological Society of London Scientific Statement: what the geological record tells us about our present and future climate
  • The Nonesuch Formation Lagerstätte: a rare window into freshwater life one billion years ago
  • The Shibantan Lagerstätte: insights into the Proterozoic–Phanerozoic transition
  • Linking surface and subsurface volcanic stratigraphy in the Turkana Depression of the East African Rift system
  • Terrestrial stratigraphical division in the Quaternary and its correlation
More...

Journal of the Geological Society

  • About the journal
  • Editorial Board
  • Submit a manuscript
  • Author information
  • Supplementary Publications
  • Subscribe
  • Pay per view
  • Alerts & RSS
  • Copyright & Permissions
  • Activate Online Subscription
  • Feedback
  • Help

Lyell Collection

  • About the Lyell Collection
  • Lyell Collection homepage
  • Collections
  • Open Access Collection
  • Open Access Policy
  • Lyell Collection access help
  • Recommend to your Library
  • Lyell Collection Sponsors
  • MARC records
  • Digital preservation
  • Developing countries
  • Geofacets
  • Manage your account
  • Cookies

The Geological Society

  • About the Society
  • Join the Society
  • Benefits for Members
  • Online Bookshop
  • Publishing policies
  • Awards, Grants & Bursaries
  • Education & Careers
  • Events
  • Geoscientist Online
  • Library & Information Services
  • Policy & Media
  • Society blog
  • Contact the Society

Published by The Geological Society of London, registered charity number 210161

Print ISSN 
0016-7649
Online ISSN 
2041-479X

Copyright © 2021 Geological Society of London