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The Neoproterozoic–Palaeozoic transition (NPT) around 600 Ma ago was a critical time interval when the Earth experienced fundamental change, manifested as climatic extremes – ‘snowball Earth’ – followed by the emergence and rapid diversification of animals – ‘Cambrian explosion’. How animals and environments co-evolved, and what caused these fundamental changes to the Earth system during the NPT, is a great scientific puzzle, which has been a rapidly developing frontier of interdisciplinary research between bio- and geosciences. South China preserves a complete stratigraphic succession of the NPT developed in various facies ranging from shallow to deep marine realms with extraordinarily well-preserved, successive fossil biotas in various taphonomic settings (Zhu, 2010; Fig. 1), making it a key area and global focus of studies in the field over recent decades. Indeed, the current narrative of early animal evolution has largely been based on the fossil biotas from South China. These include: (1) the world's oldest microscopic animal fossils with cellular details from the early Ediacaran Weng'an biota (Doushantuo Formation); (2) putative macroscopic animal fossils preserved as carbonaceous imprints from the early Ediacaran Lantian, Wenghui and Miaohe biotas (also Doushantuo Formation); (3) typical late Ediacaran faunas, preserved in dark limestone (Shibantan biota) and as large and poorly mineralized tubular animal fossils (Gaojiashan biota), both from the Dengying Formation; (4) phosphatized small shelly and soft-bodied animal fossils from the early Cambrian Meishucun and Kuanchuanpu faunas; and (5) Cambrian fossil Lagerstätten (Chengjiang, Guanshan and Kaili faunas) with typical Burgess Shale-type soft-bodied preservation.
As a well-known phosphatized Lagerstätte, the Ediacaran Weng'an biota in central Guizhou Province of South China contains diverse acanthomorphic acritarchs, algal thalli, tubular microfossils as well as various spheroidal fossils. These fossils provide crucial palaeontological evidence for the radiation of multicellular eukaryotes after the termination of the Neoproterozoic global glaciation. While the Weng'an biota is generally considered as early Ediacaran in age on the basis of phosphorite Pb–Pb isochron ages ranging from 572 Ma to 599 Ma, the reliability and accuracy of these age data have been questioned and some geologists have proposed that the Weng'an biota may be younger than 580 Ma instead. Here we report a SIMS zircon U–Pb age of 609 ± 5 Ma for a tuffaceous bed immediately above the upper phosphorite unit in the Doushantuo Formation at Zhangcunping, Yichang, South China. Litho-, bio- and chemostratigraphic correlations suggest that the upper phosphorite unit at Zhangcunping can be well correlated with the upper phosphorite unit at Weng'an, which is the main horizon of the Weng'an biota. We therefore conclude that the Weng'an biota could be as old as 609 ± 5 Ma.
Fossiliferous Ediacaran successions of South China, the Doushantuo and Dengying formations and their equivalents, are key to understanding bio- and geological evolution at the Neoproterozoic–Cambrian transition. However, their absolute ages, especially the upper Ediacaran successions, are poorly constrained. SIMS zircon U–Pb dating results in this study suggest that ash beds at the basal and middle parts of the Jiucheng Member (middle Dengying Formation) in eastern Yunnan Province were deposited at 553.6 ± 2.7/(3.8) Ma and 546.3 ± 2.7/(3.8) Ma, respectively. These new dates indicate that the age for the base of Dengying Formation in eastern Yunnan Province is similar to the 550.55 ± 0.75 Ma date, which is from an ash bed at the top of the Miaohe Member and has been regarded as the age for the base of Dengying Formation in Yangtze Gorges area. These dates do not permit a clear test of the two correlation models for the chronostratigraphic position of the Miaohe Member (uppermost Doushantuo Formation vs. middle Dengying Formation), implying that further integrated intra-basinal stratigraphic correlations and more high-resolution chronological data from the upper Ediacaran deposits of South China are required. New dates of the Jiucheng Member constrain the age of the fossil biotas in the middle Dengying Formation and extend the stratigraphic range of Rangea, Hiemalora and Charniodiscus to 546.3 ± 2.7/(3.8) Ma. The geochronology of the Dengying Formation implies that Ediacaran-type fossils preserved in this formation are younger than the White Sea Assemblage and temporally overlapping with the Nama Assemblage.
The Asteridium–Heliosphaeridium–Comasphaeridium (AHC) acritarch assemblage is composed of common organic-walled microfossils in the basal Cambrian chert–phosphorite units in South China, indicating that the AHC assemblage can be a useful biostratigraphic tool for the Ediacaran–Cambrian boundary successions in the Yangtze Platform. To test the validity of the AHC acritarch assemblage as a biostratigraphic tool, the stratigraphic range of the AHC acritarch assemblage was confirmed, and its spatial and temporal relationships to other bio- and chemostratigraphic tools were analysed in the Yanjiahe Formation, Yangtze Gorges area, South China. The result shows that the AHC assemblage temporally correlates to the Anabarites trisulcatus–Protohertzina anabarica Assemblage Zone, and spatially correlates to the large negative carbon isotope anomaly of the lowermost Cambrian (BACE) in the Yanjiahe Formation. This implies that the radiation of phytoplankton occurred slightly before the radiation of the small shelly fossils, and the AHC acritarch assemblage can be another important chronological reference to the lowermost Cambrian successions in South China, and potentially to global correlations.
The Wa'ergang section in South China has been proposed as a potential Global Stratotype Section and Point (GSSP) for the base of Stage 10, the uppermost stage of the Cambrian System. In this study, high-resolution C-isotopic compositions are reported and we identified three large negative δ13C excursions, namely N1, N2 and N3, at Wa'ergang. The N1 is located just above the First Appearance Datum (FAD) of Lotagnostus americanus, corresponding to the possible base of the Proconodontus posterocostatus conodont Zone. The N2 was identified within the Micragnostus chuishuensis trilobite Zone and the Proconodontus muelleri conodont Zone. The N3 is located in the lowermost part of the Leiagnostus cf. bexelli – Archaeuloma taoyuanense trilobite Zone or Eoconodontus conodont Zone. The N1 and N2 can be correlated with the negative δ13C excursions preceding the Top of Cambrian Carbon Isotope Excursion (TOCE) observed globally. The N3 can be correlated with the TOCE or the HEllnmaria–Red Tops Boundary (HERB) Event. The inter-basinal correlation of N1 and L. americanus strongly supports that the base of Stage 10 may be best defined by the FAD of L. americanus. We also used a box model to quantitatively explore the genesis of the negative δ13C excursions from South China. Our numerical simulations suggest that weathering of the organic-rich sediments on the platform, probably driven by intermittent sea level fall and/or the oxygenation of the Dissolved Organic Carbon (DOC) reservoir in seawater, may have contributed to the generation of the negative δ13C excursions observed in the Stage 10 at Wa'ergang in South China.
Sedimentary rocks from the Ediacaran–Cambrian boundary record important biological, climatic and geotectonic changes during this time. To date, only few geochemical investigations on the upper Ediacaran – upper Cambrian rocks in the Tarim Basin have been carried out. Here, we report high-resolution δ13Ccarb records from the Penglaiba, the Wushi phosphorite and the Dongergou sections from Ediacaran–Cambrian Series 3 in the Keping area of the Tarim Basin. The sections display several obvious δ13Ccarb shifts; δ13Ccarb values increased from 3‰ to 6.7‰ across the Qigebulage Formation. Moreover, a negative δ13Ccarb shift across the Ediacaran–Cambrian boundary is apparent; δ13Ccarb values decreased to a minimum of −9.8‰ in the Wushi phosphorite section (−7.7‰ in Dongergou section and −5.4‰ in Penglaiba section), followed by a positive carbonate carbon isotopic excursion across the Yuertusi Formation into the middle of the overlying Xiaoerbulak Formation. Furthermore, more or less invariable positive δ13Ccarb values characterize the middle and upper Xiaoerbulak Formation. The most negative δ13Ccarb value (−14.3‰) occurred near the base of the Shayilik Formation, which is the absolute minimum value among the studied sections of the Cambrian Series 2 to Cambrian Series 3 transition in the world. The δ13C data from Keping, Tarim Basin are in good agreement with carbon isotope profiles recorded in South China, and these events may reflect the perturbation of the carbon cycle in the Tarim Basin during the Ediacaran–Cambrian and the Cambrian Series 2 – Cambrian Series 3 transitions.
Non-biomineralizing Ediacaran macrofossils are rare in carbonate facies, but they offer valuable information about their three-dimensional internal anatomy and can broaden our view about their taphonomy and palaeoecology. In this study, we report a new Ediacaran fossil, Curviacus ediacaranus new genus and species, from bituminous limestone of the Shibantan Member of the Dengying Formation in the Yangtze Gorges area of South China. Curviacus is reconstructed as a benthic modular organism consisting of serially arranged and crescent-shaped chambers. The chambers are confined by chamber walls that are replicated by calcispars, and are filled by micritic sediments. Such modular body construction is broadly similar to the co-occurring Yangtziramulus zhangii and other Ediacaran modular fossils, such as Palaeopascichnus. The preservation style of Curviacus is similar to Yangtziramulus, although the phylogenetic affinities of both genera remain unresolved. The new fossil adds to the diversity of Ediacaran modular organisms.
Enigmatic chambered structures have been reported forming reef frames in Cryogenian interglacial carbonates, prior to the commonly acknowledged microbial-metazoan reefs at the terminal Ediacaran, and interpreted as fossils of possible sponge-grade organisms. A better constraint on the affinity of these structures is partly hindered by few analogues in other time periods. This study describes similar structures from peritidal dolostones of the Ediacaran Denying Formation from Yunnan, China. Samples were investigated using optical microscopy and three-dimensional (3-D) reconstruction based on grinding tomography. The Dengying chambered structures are comparable with Cryogenian structures in basic construction, but are not frame building, and show variations in overall shape and inhabiting facies. Two-dimensional (2-D) cross-sections show that thin, homogeneous micritic laminae are the basic building blocks of the chamber walls. Thick walls represent parallel accretion of these laminae, and thin walls developed from the angular growth of a single lamina or merging of multiple laminae. In 3-D space, the laminae primarily correspond to continuous surfaces which sometimes contain sub-circular holes, while a few represent filamentous elements connected to the surfaces. The morphological features and growth pattern of the Dengying chambered structures indicate that they are likely to be calcified microbial constructions rather than skeletal remains of basic metazoans. However, aside from the Cryogenian and Dengying examples, comparable chambered constructions with laminae-based architecture are yet unknown from other fossil or extant microbialites. Further work investigating related structures is needed to determine the microbial consortia and controlling environmental factors that produced these chambered structures.
Microbially induced sedimentary structures (MISS) are microbial traces in sandy deposits. They have been formed by various modes of microbial behaviour in response to the prevailing physical dynamics in shallow-marine environments since early Archaean time. An association of fossils, phosphatic structures and grains, stromatolites and microbial laminated levelling structures is documented from the Zhongyicun Member of Lower Cambrian strata in the Baideng section dated close to 535 Ma. SEM examination demonstrates that microbial laminated levelling structures are the result of the development of microbial mats composed of filamentous structures. We propose that there was a short hiatus after the dolomite layer deposited when the dolomite layer was weathered to form centimetre-scale valleys firstly, and then microbes accumulated in these valleys and formed the microbial laminated levelling structures.
The earliest fossil record of animal biomineralization occurs in the latest Ediacaran Period (c. 550 Ma). Cloudina and Sinotubulites are two important tubular taxa among these earliest skeletal fossils. The evolutionary fate of Cloudina-type fossils across the Ediacaran–Cambrian transition, however, remains poorly understood. Here we report a multi-layered tubular microfossil Feiyanella manica gen. et sp. nov. from a phosphorite interval of the lowest Cambrian Kuanchuanpu Formation, southern Shaanxi Province, South China. This newly discovered fossil is a conical tube with a ‘funnel-in-funnel’ construction, showing profound morphological similarities to Cloudina and Conotubus. On the other hand, the outer few layers, and particularly the outermost layer, of Feiyanella tubes are regularly to irregularly corrugated, a feature strikingly similar to the variably folded/wrinkled tube walls of Sinotubulites. The Feiyanella tubes additionally exhibit two orders of dichotomous branching, similar to branching structures reported occasionally in Cloudina and possibly indicative of asexual reproduction. Owing to broad similarities in tube morphology, tube wall construction and features presumably indicative of asexual reproduction, Cloudina, Conotubus, Sinotubulites and the here described Feiyanella may thus constitute a monophyletic group traversing the Ediacaran–Cambrian boundary. The tube construction and palaeoecological strategy of Feiyanella putatively indicate evolutionary continuity in morphology and palaeoecology of benthic metazoan communities across the Ediacaran–Cambrian transition.
Appendage anatomy contributes crucial data for understanding the evolution and ecology of Euarthropoda. The Palaeozoic trilobites show a great diversity of exoskeletons in the fossil record. However, soft parts, especially appendages, have only been discovered from a few trilobite species. Here we report extraordinarily preserved appendages in the trilobite species Hongshiyanaspis yiliangensis Zhang & Lin in Zhang et al. 1980 (Redlichiida, Metadoxididae) from a single mudstone layer of the Xiazhuang fossil assemblage within the Hongjingshao Formation (Cambrian Series 2, Stage 3) near Kunming, Yunnan, SW China. The appendages exhibit the common architecture revealed by other trilobites and artiopods by consisting of a pair of uniramous antennae followed by a series of paired homonomous biramous limbs. The antennae in holaspid individuals comprise up to 27 spinous podomeres and their ontogeny occurs by lengthening of the podomeres. The post-antennal biramous limbs are similar to those in other polymerid trilobites and artiopods by having a single-segmented protopodite and an endopodite comprising seven segments, but possess a unique wide tripartite exopodite with long setae. Sophisticated appendage anatomy, including the body–limb junction, fine setae, putative muscle bundles and duct-type tissues, are also revealed. Appendages of trilobites, artiopods and other upper stem-group euarthropods are compared and summarized. The H. yiliangensis appendages highlight the high morphological disparity of exopodites and the conservativeness of endopodites in trilobites and artiopods. This morphological pattern, together with similar body patterning seen in crustaceans but not in chelicerates, supports the mandibulate affinities of trilobites and at least some artiopods.
A well-preserved fossil priapulid worm, Xiaoheiqingella sp., is reported from the early Cambrian Guanshan Lagerstätte (Cambrian Series II, Stage 4) near Kunming City, Yunnan Province, SW China. The body of the animal consists of four sections: a swollen introvert, a constricted neck, a finely annulated trunk and a caudal appendage. The body configuration exhibits a close resemblance to that of the crown group priapulid Xiaoheiqingella peculiaris from the early Cambrian Chengjiang Lagerstätte. The new discovery provides another striking example of crown group priapulids, representing the third occurrence of crown group fossil priapulids after the Chengjiang Lagerstätte (Cambrian Series II, Stage 3) and the Mazon Creek Lagerstätte (late Moscovian Stage, Pennsylvanian). The discovery also sheds new light on the early diversity and evolution of priapulid worms.
The skeleton of a new vauxiid sponge (Order Verongida), Angulosuspongia sinensis gen. et sp. nov., described on the basis of material from calcareous mudstones of the Kaili Formation (Cambrian Stage 5), Jianhe area, Guizhou, South China, is composed of two layers of fused spicules outlining hexagonal or polygonal openings. These vauxiid remains are the first reported from outside Laurentia, and represent only the second genus attributed to the family. Its age is close to but still slightly older than the Burgess Shale Biota, and it appears to be a primitive relative of other members of the Vauxiidae. The morphological differences between Chinese and Laurentian vauxiid sponges may be a result of vicariance. These specimens not only extend the geographic distribution of vauxiids, but also help to fill a chronostratigraphical gap between North Greenland and North American material and provide additional evidence for understanding the evolutionary history of the Demospongiae.
Nitrogen is an essential element for biological activity, and nitrogen isotopic compositions of geological samples record information about both marine biological processes and environmental evolution. However, only a few studies of N isotopes in the early Cambrian have been published. In this study, we analysed nitrogen isotopic compositions, as well as trace elements and sulphur isotopic compositions of cherts, black shales, carbonaceous shales and argillaceous carbonates from the Daotuo drill core in Songtao County, NE Guizhou Province, China, to reconstruct the marine redox environment of both deep and surface seawater in the study area of the Yangtze shelf margin in the early Cambrian. The Mo–U covariation pattern of the studied samples indicates that the Yangtze shelf margin area was weakly restricted and connected to the open ocean through shallow water flows. Mo and U concentrations, δ15Nbulk and δ34Spy values of the studied samples from the Yangtze shelf margin area suggest ferruginous but not sulphidic seawater and low marine sulphate concentration (relatively deep chemocline) in the Cambrian Fortunian and early Stage 2; sulphidic conditions (shallow chemocline and anoxic photic zone) in the upper Cambrian Stage 2 and lower Stage 3; and the depression of sulphidic seawater in the middle and upper Cambrian Stage 3. Furthermore, the decreasing δ15N values indicate shrinking of the marine nitrate reservoir during the middle and upper Stage 3, which reflects a falling oxygenation level in this period. The environmental evolution was probably controlled by the changing biological activity through its feedback on the local marine environment.
The early Cambrian Period was a key interval in Earth history with regard to changes in both ocean chemistry and animal evolution. Although increasing ocean ventilation has been widely assumed to have played a key role in the rapid appearance, diversification and spatial colonization of early animals, this relationship is in fact not firmly established. Here, we report a high-resolution Fe-C-S-Al-Ti geochemical study of the lower Cambrian Wangjiaping section from an outer-shelf setting of the Yangtze Sea of South China. Iron speciation data document a redox transition from dominantly euxinic to ferruginous conditions during Cambrian Age 3 (c. 521–514 Ma). Interpretation of coexisting pyrite sulphur isotope (δ34Spy) records from Wangjiaping reveals relatively high marine sulphate availability at Wangjiaping. Furthermore, Wangjiaping section shows lower δ34Spy (‒2.1±5.3‰) and lower TOC (2.4±1.1%) values but higher positive correlation (R2 = 0.66, p < 0.01) between TOC and Fepy/FeHR relative to deeper sections reported previously, suggesting that euxinia developed at Wangjiaping in response to increasing marine productivity and organic matter-sinking fluxes. Our reconstructed redox conditions and fossils at Wangjiaping in comparison with previously well-studied strata in the inner-shelf Xiaotan and Shatan sections suggest that planktonic and benthic planktonic trilobites with bioturbation appeared in the oxic water columns, whereas only planktonic trilobites without bioturbation occurred within the anoxic (even euxinic) water columns during Cambrian Age 3. This finding indicates that spatial heterogeneity of redox conditions in the shelves had an important effect on early animal distribution in the Yangtze Block.
The Fulu iron formation (IF) is an iron-rich unit in the Neoproterozoic glacial successions, South China. The major element data suggest that the iron sources of the Fulu IF are derived from binary mixing from hydrothermal and detrital loads. The Fulu IF is characterized by slightly positive Eu anomalies similar to other Neoproterozoic IFs, indicating that a high-temperature hydrothermal input may contribute little to Neoproterozoic IFs. A shift from non-existent to slightly negative Ce anomalies of the Fulu IF indicates that the IF precipitated across an iron chemocline separating a weakly oxic surface ocean from an oxygen-depleted deep ocean.
The Ediacaran Doushantuo Formation at Weng'an, South China hosts well-preserved phosphatized microfossils known as the Weng'an biota. A laser ablation ICP-MS analysis of rare earth element (REE) characteristics of the fossil-bearing black phosphorite unit of the Doushantuo Formation at Weng'an was conducted, with the aim of unravelling the depositional conditions and diagenetic processes during formation of the phosphorites. Spherical phosphatized microfossils and phosphatic clasts were analysed, and the REE data display middle REE (MREE) -enriched shale-normalized REE patterns. The spherical phosphatized microfossils show an increase in total REE contents (∑REE) from core to rim. Negative correlations between ∑REE and the extent of MREE enrichment over the other REE (indicated by LaN/SmN, YbN/SmN) are observed for analysed spots within individual phosphatic grains, which may be due to complex diagenetic history of the phosphatic grains, with fluctuations in REE sources and chemical parameters in a high-energy shallow-water environment being additional factors. The LaN/YbN and LaN/SmN characteristics of the phosphatic grains suggest they were mostly influenced by early diagenetic alteration rather than late extensive recrystallization. The negative Ce anomalies in the samples suggest they formed under oxic-bottom-water conditions. Positive Eu anomalies are present in all samples, and are likely to reflect involvement of hydrothermal fluids rather than changes in redox conditions of porewater. Overall this study has major implications for phosphorites as important archives for the study of geochemical proxies, the Ediacaran period and also evolutionary changes.