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The widespread development of Permian/Triassic boundary microbialites is a key biosedimentary response to the mass extinction event. The microbial communities that precipitated these structures remain, however, poorly resolved. Here, we report abundant columnar microfossils from five Permian‐Triassic successions in South China. Four phases of the microfossil life cycle are recorded: baeocytes, juvenile column, mature column, and baeocyte‐splitting, which are interpreted as Pleurocapsales (cyanobacterial clade that reproduces by multiple fission). The great thickness (up to 13 m) of these microbial deposits and Raman spectrum analysis indicate a strong calcification capacity of the Pleurocapsales. Pronounced peaks of Mg/(Mg + Ca) ratios from microbialite and distinct Mg‐enrichment in microfossils indicate that Pleurocapsales blooms may also have facilitated dolomitization of the microbialites. Common occurrence, strong biomineralization capacity, baffling framework, and the close association with carbonate minerals suggest that Pleurocapsales were key constructors of post‐extinction microbialites. Plain Language Summary: As a biosedimentary response to the end‐Permian mass extinction, the widespread occurrence of microbialites has attracted increasing levels of interests from geologists worldwide. However, the major constituents of microbial communities that constructed the microbialites have long been disputed. Our new study describes morphological features, growth direction and reproduction characteristic of columnar microfossils obtained from the microbialites, and all characteristics indicate that these microfossils are assignable to a type of photosynthesizing bacteria, known as Pleurocapsales cyanobacteria. Both biomineralization and dolomitization of these Pleurocapsales are evaluated based on in‐situ observations and geochemical analyses. The new results show that the cluster‐branching morphologies, together with common occurrence and strong calcification capacity indicate that Pleurocapsales were key constructors of the microbialites. High Mg/(Mg + Ca) ratios coinciding with the microbialites and Mg‐enrichment in the microfossils suggest that the Pleurocapsales may have catalyzed dolomitization of the microbialites after the end‐Permian mass extinction. Key Points: Four phases of a microbe life cycle are recognized from fossils within microbialites reflecting Pleurocapsales asexual reproductionCommon occurrence and baffling framework indicate that Pleurocapsales cyanobacteria are important in constructing microbialitesPleurocapsales cyanobacteria were an important component of microbialite calcification and magnesium enrichment [ABSTRACT FROM AUTHOR]

Redox variations across the Permian‐Triassic boundary (PTB) have long been debated, especially during the proliferation of PTB microbialites. Here, we report redox fluctuations across the PTB to evaluate links between the two based on pyrite framboid analysis from basin to platform settings in South China. During the end‐Permian extinction, abundant framboids indicate a widespread anoxia that was likely a direct cause of extinction. In the earliest Triassic (Hindeodus parvus conodont zone), pyrite framboids were absent in ramp to basin and shallow, nonmicrobialite platform sections. In contrast, the coeval microbialites yield abundant framboids indicative of dysoxia. The fact that framboids were only confined to PTB microbialites and absent in other habitats indicates that microbe bloom may have stimulated dysoxic watermass and triggered the framboid growth within microbe aggregates. Thus, microbialites were not built in reducing settings, but instead, microbial proliferation caused local, dysoxia within shallow oxygenated platforms after the extinction. Plain Language Summary: The widespread occurrence of microbialites immediately after the end‐Permian extinction has attracted interest about their formation conditions. Microbialites are generally believed to be formed during microbial bloom in reducing conditions linked to the mass extinction. However, the cause‐and‐effect relationship between microbe bloom and reducing states of seawater has long been disputed. Our new results based on analyses of 17,306 pyrite framboid diameters and morphologies derived from various Permian‐Triassic boundary sections in South China show that microbialite formation (microbial bloom) was not triggered by reducing waters upwelled from deeper water masses, instead, it caused local, dysoxic areas within oxygenated platforms after the end‐Permian crisis. Key Points: A total of 17,306 pyrite framboids are analyzed in size and morphology from the Permian‐Triassic boundary beds of 26 sections, South ChinaFramboids were absent in ramp to basin and shallow, nonmicrobialite platforms after the extinction, but occurred in coeval microbialitesMicrobe bloom/microbialite growth was not triggered by reducing condition, but stimulated dysoxic water mass and triggered framboid growth [ABSTRACT FROM AUTHOR]

Abstract A new Permian–Triassic boundary microbialite (PTBM) is described from the Xiajiacao section of western Hubei Province, South China. The new microbialite, 3.16 m thick, comprises a thin layer of stromatolite and a thick thrombolite unit. An irregular contact separates the uppermost Permian skeletal packstone from the post-extinction stromatolite, but it is not yet possible to discriminate whether it was formed by submarine solution in the wake of ocean acidification or subaerial exposure due to regional regression, or a combination of both. The stromatolite shows "cabbage-like" morphology, and the thrombolite is characterized by centimetric clotted texture. Abundant columns of the microproblematica structure Gakhumella , coccoid-like spheroids, bacterial clump-like spheroids, and hollow spheroids are recognized in both stromatolites and thrombolites and may have played an important role in accretion of the microbialites. Pyrite framboid analysis indicates that microbialites may have been affected by lower dysoxic to upper dysoxic conditions in the immediate aftermath of the Permian–Triassic extinction. Stratigraphic abundance of both high-temperature grains (β-quartz and glassy balls) peaked ~ 20 cm below the biotic extinction horizon, implying that either intensive volcanic eruption occurred only just prior to biotic extinction, or volcanism was still intensive during biotic extinction, but volcanic grains were not deposited in the Xiajiacao locality, likely due to the shallow, agitated environment caused by the regional regression. Overall, microbial bloom, indicated by the widespread PTBMs, seems to have been little affected by the contemporaneous volcanism. Highlights • Gakhumella,and coccoid-like, bacterial clump-like and hollow spheroids are crucial in accretion of microbialite; • Pyrite framboid size and morphology show that the microbialite may have been deposited in lower to upper dysoxic conditions. • Microbial bloom, indicated by the widespread PTBMs seems not to be linked with the contemporaneous volcanism. [ABSTRACT FROM AUTHOR]

Unusual sedimentary deposits such as stromatolites, thrombolites, dendrolites, and giant ooids are widely distributed in the Permian–Triassic boundary (PTB) successions of shallow platform facies worldwide. These microbial fabrics, however, are rarely present in the same section. The genesis of these PTB microbialites and the oceanographic conditions linking to the end-Permian mass extinction still remain disputed despite intense study. Here, we report a stromatolite-dendrolite-thrombolite-giant ooid association from the PTB beds of the Yudongzi section on the northwestern margin of the Yangtze Platform, South China, which was situated in low latitudes during the Permian-Triassic (P-Tr) transition. The Yudongzi microbialites rest on an irregular contact with underlying Permian bioclastic limestone and are comprised of lower bedded stromatolite, digitate dendrolite, upper bedded stromatolite, and thrombolite units. The bedded stromatolite is characterized by undulating laminated structures and various amalgamated and layered clots. The digitate dendrolite is made up of digitate-like branches, formed by clots distributed in an arched arrangement, and characterized by abundant branching dendroids. The thrombolite displays distinct clotted textures that comprise a mixture of dark-colored microbial fabric and light-colored interstitial sediment. In microbialite, columnar structures identified as Gakhumella -like calcimicrobe are very abundant. Each individual of Gakhumella -like structure has densely arranged segments forming a columnar- to fan-shaped micro-structure. Single segments are arch-shaped and form a thin chamber between segments. Coccoid-like, bacterial clump-like, and other spheroidal aggregates are also commonly present and suggestive of a biogenetic origin for the Yudongzi microbialites. Moreover, distinct aragonite crystal fans are also for the first time found in the PTB microbialites in South China, suggesting that the fans may have resulted from high inorganic carbon concentrations, and thus high levels of carbonate dioxide in the seawater. The Yudongzi microbialites are capped by shelly grainstone and a massive giant oolite, implying that the demise of the Yudongzi microbialites may be due to destruction by strong wave currents in a high energy setting. Some ooids are coated unevenly by micritic laminae (or envelopes) indicating the possibility of a microbe-related origin. These microbial-coated giant ooids, together with the massive microbialites, may imply the long-term of microbial bloom in the oceans immediately after the end-Permian extinction. The co-existence of abundant Gakhumella-like microorganisms and oxygen-dependent invertebrates, together with some coccoid-like spheroids, aragonite cement crystal fans, and pyrite framboids indicative of oxygen-poor conditions, implies that oxygenation state of the PTB microbialites is much more complex than previously recognized, demonstrating need for continued multidisciplinary examination of facies changes associated biotic mass extinction. [Display omitted] • PTB microbialite complex comprises bedded stromatolite, digitate dendrolite, thrombolite and giant ooids. • Microbial fabrics include undulating laminated structures, digitate-like branches, and clotted textures. • Ooids are coated by micritic laminae indicating the possibility of a microbe-related origin. • Columnar structures identified as Gakhumella are very abundant and belong to the benthic Pleurocapsale cyanobacteria. • Aragonite crystal fans are distinct, and may have resulted from high bicarbonate in the seawater. [ABSTRACT FROM AUTHOR]

Abstract We report two shallow marine, ichnofauna-bivalve-microbial mat biofacies from the Lower Triassic Xiahuancang Formation of the southern Qilian area, Qinghai Province, northwestern China, which was located at moderate-high paleolatitudes on the northern margin of the Paleotethys Ocean. Paleoenvironmental analyses show that Members I and II of the Xiahuancang Formation represent a shoreface and a lower shoreface to offshore transition setting, respectively. Biofacies 1, recognized from Member I, is characterized by a diverse ichnofauna (including deep-tiers of Rosselia and Diplocraterion), Claraia -dominated bivalves, and microbially induced sedimentary structures (MISSs). Biofacies 2, in succeeding Member II, is dominated by a diverse ichnofauna, epifaunal and shallow infaunal bivalves, and wrinkle structures. Primary co-occurrences, preservational features, and palimpsest or crosscutting relationships of all components within the biofacies indicate that microbial mats, bivalves, and trace-makers actively interacted with one another during deposition. They largely represent contemporaneous biotic associations. Microbial mats are interpreted to have grown under well-oxygenated conditions after storm deposition due to the association of deep-tiering infauna and diverse epifauna as well as well-developed cross-stratification, and the top layer of microbial mats could serve as an oasis for metazoans. Microbial mats not only proliferated in harsh environments, but also coexist with epifauna and deep-tiering infauna in well-oxygenated settings following the end-Permian crisis. Their occurrences in the Early Triassic are unrelated to environmental stresses, which are coincident with their sedimentologic record from other geological time intervals. Apparently, the southern Qilian ichnofauna-bivalve-microbial mat biofacies with juxtaposed epifauna, infauna and MISS- or wrinkle-related microbial mats represents a matground-adapted benthic metazoan ecosystem in the earliest Triassic, which provided sufficient oxygen, food, and other hydrodynamic conditions hospitable for metazoans and ecosystems to recover and reached a fairly high level (recovery stage 3) in a tough time when most biotas suffered biotic depletion and environmental stress. The biotic components in unique biofacies may represent a phase shift community in siliciclastic settings during the Early Triassic. Graphical abstract Unlabelled Image Highlights • The ichnofauna-bivalve-microbial mat biofacies is characterized by diverse ichnofauna, bivalves, and MISS; • Low ichnodiversity in the Induan cannot be taken as an indicator of environmental stresses in all cases; • Microbial mats proliferated in both harsh environments and well-oxygenated settings following the end-Permian crisis; • The southern Qilian biofacies represents a matground-adapted metazoan ecosystem during the earliest Triassic; • Such unique biofacies represent community phase shifts in siliciclastic settings during the Early Triassic. [ABSTRACT FROM AUTHOR]

A late Smithian ichnoassemblage is reported from the Lower Triassic succession from the Lichuan section, western Hubei Province, South China. This ichnoassemblage comprises 13 ichnogenera, which include simple, horizontal traces ( Cochlichnus , Cosmorhaphe , Didymaulichnus , Gordia , Circulichnis , and Planolites ), vertical traces ( Arenicolites , Laevicyclus ), oblique- or horizontal branching traces ( Palaeophycus ), slightly complex burrow networks ( Thalassinoides , and Treptichnus ), and grazing traces ( Cosmorhaphe and Gyrochorte ). These traces are also categorized into fodinichnia, domichnia, pascichnia, and repichnia ethologic types, suggesting a fairly high degree of behavioral complexity. Of these, the fodinichnia traces are most abundant. The ichnoassemblage horizons usually have rather high ichnofabric indices (ii), reaching ii 4–5. Bedding plane coverage is up to 70–90%, resulting in high bedding plane bioturbation index as well (BPBI 4–5). The Lichuan ichnoassemblage traces have mean and maximum diameters of 4.8 mm and 9 mm, respectively ( n = 342). The traces penetrate to a depth of up to 30 mm into the sediment, with an average depth of penetration of 15 mm, indicating third to fourth tiering levels (3–4). Overall, the Lichuan ichnofauna shows an abrupt increase in ichnodiversity, burrow width, behavioral complexity, and ecologic tiering level, which are indicative of recovery stages of 2–3 in the late Smithian. When compared to five coeval ichnoassemblages from other sections in South China (Yashan, Susong, Daxiakou, Gaimao, and Tianshengqiao) and coeval ichnoassemblages from elsewhere (northern Italy, Western Australia, western US), the Lichuan trace-fossil assemblage reaches similar level of diversity during this time interval. Paleoenvironmental analysis indicates that the late Smithian recovery pulse is typically confined to the offshore transition setting, whereas more distal zones record much lower levels of recovery. Therefore, time and environmental conditions were the two crucial factors controlling the rate and degree of recovery of trace-making organisms in the aftermath of the latest Permian mass extinction. [ABSTRACT FROM AUTHOR]

Two Anisian (Middle Triassic) marine ichnocoenoses are reported from the Boyun and Junmachang (JMC) sections located along the eastern and western margins of the Kamdian Continent, Yunnan Province, Southwest China, respectively. The Boyun ichnoassemblage is middle Anisian in age and is dominated by robust Rhizocorallium , while the JMC ichnoassemblage is of an early Anisian age and is characterized by the presence of Zoophycos . The ichnoassemblage horizons of the Boyun section represent an inner ramp environment, while the JMC section was likely situated in a mid-ramp setting near storm wave base as indicated by the presence of tempestites. The ichnofossil-bearing successions are usually highly bioturbated in both the Boyun (BI 3–5, BPBI 5) and JMC (BI 3–4, BPBI 3–4) sections. Three large, morphologically complicated ichnogenera: 1) Rhizocorallium ; 2) Thalassinoides ; and, 3) Zoophycos characterize the Anisian ichnocoenoses. Of these, Rhizocorallium has mean and maximum tube diameters up to 20.4 mm and 28 mm, respectively, while Thalassinoides mean and maximum tube diameters are 14.2 mm and 22 mm, respectively. Zoophycos is present in the early Anisian strata of the JMC section, and represents the oldest known occurrence of this ichnogenus following the latest Permian mass extinction. Similar to coeval ichnoassemblages elsewhere in the world, the Yunnan ichnocoenoses embrace a relatively low ichnodiversity, but their burrows usually penetrate deeply into the sediment, and include large and complex Rhizocorallium and Thalassinoides . All of these ichnologic features are indicative of recovery stage 4 after the latest Permian crisis. Anisian ichnoassemblages occur globally in six different habitat settings, and all show similar ecologic characteristics except for slightly different degrees of ichnotaxonomic richness, indicating that depositional environment is not a crucial factor shaping the recovery of the trace-makers, but may have an impact on their ichnodiversity. When compared with some important Early Triassic (mainly Spathian) ichnoassemblages worldwide, the Anisian ichnocoenoses examined for this study are slightly less diversified, and possess more or less the same maximum burrow sizes, but the penetration depth of burrows and the distribution of burrow sizes are much larger than those from the Early Triassic. It is worthy of note that the lower ichnodiversity of the Anisian ichnocoenoses may have resulted from intense bioturbation by deeper tiers, representing a taphonomic product that is totally unrelated to environmental stress. In addition, Anisian Rhizocorallium and Thalassinoides have much larger burrow sizes than the same ichnotaxa from the Lower Triassic, implying that ichnocoenoses may have recovered in Spathian, but did not stabilize until the Anisian. [ABSTRACT FROM AUTHOR]

The Meishan section, South China is the Global Stratotype Section and Point (GSSP) for the Permian–Triassic boundary (PTB), and is also well known for the best record demonstrating the Permian–Triassic mass extinction (PTME) all over the world. This section has also been studied using multidisciplinary approaches to reveal the possible causes for the greatest Phanerozoic biocrisis of life on Earth; many important scenarios interpreting the great dying have been proposed on the basis of data from Meishan. Nevertheless, debates on biotic extinction patterns and possible killers still continue. This paper reviews all fossil and sedimentary records from the Permo-Triassic (P–Tr) transition, based on previously published data and our newly obtained data from Meishan, and assesses ecologically the PTME and its aftermath to determine the biotic response to climatic and environmental extremes associated with the biocrisis. Eight updated conodont zones: Clarkina yini , Clarkina meishanensis , Hindeodus changxingensis , Clarkina taylorae , Hindeodus parvus , Isarcicella staeschei , Isarcicella isarcica , and Clarkina planata zones are proposed for the PTB beds at Meishan. Major turnover in fossil fragment contents and ichnodiversity occurs across the boundary between Bed 24e-5 and Bed 24e-6, suggesting an extinction horizon in a thin stratigraphic interval. The irregular surface in the middle of Bed 27 is re-interpreted as a firmground of Glossifungites ichnofacies rather than the previously proposed submarine dissolution surface or hardground surface. Both fossil fragment contents and ichnodiversity underwent dramatic declines in Beds 25–26a, coinciding with metazoan mass extinction. Fossil fragment content, ichnodiversity and all ichnofabric proxies (including burrow size, tiering level, bioturbation level) indicate that the P–Tr ecologic crisis comprises two discrete stages, coinciding with the first and second phases of the PTME in Meishan. Ecologic crisis lagged behind biodiversity decline during the PTME. Pyrite framboid size variations suggest that depositional redox condition was anoxic to euxinic in the latest Changhsingian, became euxinic in Beds 25–26a, turned dysoxic in Bed 27, then varied from euxinic to anoxic through most of the Griesbachian. The ~ 9 °C increase in seawater surface temperature from Bed 24e to Bed 27 at Meishan seems to result in dramatic declines in biodiversity and fossil fragment contents in Beds 25–26a, but had little effect on all ecologic proxies. Both metazoans and infauna seem not to have been affected by the pre-extinction anoxic–euxinic condition. The anoxic event associated with the PTME may have occurred in a much shorter period than previously thought and is only recorded in Beds 25–26a at Meishan. Fossil fragment contents, ichnofaunas, ichnofabrics and pyrite framboid size all show that no signs of oceanic acidification and anoxia existed in Bed 27. The early Griesbachian anoxia may have resulted in rarity of ichnofauna and metazoans in the lower Yinkeng Formation, in which the ichnofauna is characterized by small, simple horizontal burrows of Planolites , and metazoan faunas are characterized by low diversity, high abundance, opportunist-dominated communities. The rapid increase of ~ 9 °C in sea-surface temperature and a short anoxia or acidification coincided with the first-pulse biocrisis, while a prolonged and widespread anoxia probably due to a long period of high seawater temperate condition may be crucial in mortality of most organisms in the second-pulse PTME. Marine ecosystems started to recover, coupled with environmental amelioration, in the late Griesbachian. [ABSTRACT FROM AUTHOR]

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