Microbial mat controls on infaunal abundance and diversity in modern marine microbialites

Microbialites are the most abundant macrofossils of the Precambrian. Decline in microbialite abundance and diversity during the terminal Proterozoic and early Phanerozoic has historically been attributed to the concurrent radiation of complex metazoans. Similarly, the apparent resurgence of microbialites in the wake of Paleozoic and Mesozoic mass extinctions is frequently linked to drastic declines in metazoan diversity and abundance. However, it has become increasing clear that microbialites are relatively common in certain modern shallow, normal marine carbonate environments—foremost the Bahamas. For the first time, we present data, collected from the Exuma Cays, the Bahamas, systematically characterizing the relationship between framework‐building cyanobacteria, microbialite fabrics, and microbialite‐associated metazoan abundance and diversity. We document the coexistence of diverse microbialite and infaunal metazoan communities and demonstrate that the predominant control upon both microbialite fabric and metazoan community structure is microbial mat type. These findings necessitate that we rethink prevalent interpretations of microbialite–metazoan interactions and imply that microbialites are not passive recipients of metazoan‐mediated alteration. Additionally, this work provides support for the theory that certain Precambrian microbialites may have been havens of early complex metazoan life, rather than bereft of metazoans, as has been traditionally envisaged.     

Earliest Triassic microbialites in the South China block and other areas: controls on their growth and distribution

Earliest Triassic microbialites (ETMs) and inorganic carbonate crystal fans formed after the end-Permian mass extinction (ca. 251.4 Ma) within the basal Triassic Hindeodus parvus conodont zone. ETMs are distinguished from rarer, and more regional, subsequent Triassic microbialites. Large differences in ETMs between northern and southern areas of the South China block suggest geographic provinces, and ETMs are most abundant throughout the equatorial Tethys Ocean with further geographic variation. ETMs occur in shallow-marine shelves in a superanoxic stratified ocean and form the only widespread Phanerozoic microbialites with structures similar to those of the Cambro-Ordovician, and briefly after the latest Ordovician, Late Silurian and Late Devonian extinctions. ETMs disappeared long before the mid-Triassic biotic recovery, but it is not clear why, if they are interpreted as disaster taxa. In general, ETM occurrence suggests that microbially mediated calcification occurred where upwelled carbonate-rich anoxic waters mixed with warm aerated surface waters, forming regional dysoxia, so that extreme carbonate supersaturation and dysoxic conditions were both required for their growth. Long-term oceanic and atmospheric changes may have contributed to a trigger for ETM formation. In equatorial western Pangea, the earliest microbialites are late Early Triassic, but it is possible that ETMs could exist in western Pangea, if well-preserved earliest Triassic facies are discovered in future work.     

Facies selectivity of benthic invertebrates in a Permian/Triassic boundary microbialite succession: Implications for the “microbialite refuge” hypothesis

Thrombolite and stromatolite habitats are becoming increasingly recognized as important refuges for invertebrates during Phanerozoic Oceanic Anoxic Events (OAEs); it is posited that oxygenic photosynthesis by cyanobacteria in these microbialites provided a refuge from anoxic conditions (i.e., the “microbialite refuge” hypothesis). Here, we test this hypothesis by investigating the distribution of ~34, 500 benthic invertebrate fossils found in ~100 samples from a microbialite succession that developed following the latest Permian mass extinction event on the Great Bank of Guizhou (South China), representing microbial (stromatolites and thrombolites) and non‐microbial facies. The stromatolites were the least taxonomically diverse facies, and the thrombolites also recorded significantly lower diversities when compared to the non‐microbial facies. Based on the distribution and ornamentation of the bioclasts within the thrombolites and stromatolites, the bioclasts are inferred to have been transported and concentrated in the non‐microbial fabrics, that is, cavities around the microbial framework. Therefore, many of the identified metazoans from the post‐extinction microbialites are not observed to have been living within a microbial mat. Furthermore, the lifestyle of many of the taxa identified from the microbialites was not suited for, or even amenable to, life within a benthic microbial mat. The high diversity of oxygen‐dependent metazoans in the non‐microbial facies on the Great Bank of Guizhou, and inferences from geochemical records, suggests that the microbialites and benthic communities developed in oxygenated environments, which disproves that the microbes were the source of the oxygenation. Instead, we posit that microbialite successions represent a taphonomic window for exceptional preservation of the biota, similar to a Konzentrat‐Lagerstätte, which has allowed for diverse fossil assemblages to be preserved during intervals of poor preservation.     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

A palaeobotanical perspective on the great end-Permian biotic crisis

Mass extinctions are crucial to understanding changes in biodiversity through time. However, it is still disputed whether extinction dynamics in the marine and terrestrial biotas followed comparable trajectories. For instance, while marine realms have suffered five strong depletions in diversity, the so-called ‘Big Five’ mass extinctions, only the end-Permian event appears to have also resulted in a major abrupt reduction in continental diversity. However, recent evidence based on the diversity dynamics of vegetation has suggested the presence of two major episodes of extinction in the terrestrial environments, at the end-Carboniferous and the end-Permian times. This apparent contradiction is addressed in the present study. Here, we show that while the end-Carboniferous plant extinction was focused on particular environments (e.g. tropical wetlands) and affected mainly the free-sporing plant diversity (i.e. lycopsids, ferns and progymnosperms), only the end-Permian mass extinction had devastating effects on vegetation on a global scale. If we take the biosphere as a whole, the results highlight that the end-Permian biotic crisis was the only genuine global mass extinction event, affecting widely both the marine and terrestrial environments.     

Encrusting micro-organisms and microbial structures in Upper Jurassic limestones from the Southern Carpathians (Romania)

Late Jurassic–Early Cretaceous ?tramberk-type reef limestones are known from some parts of the Southern Carpathians in Romania. The Upper Jurassic deposits mainly consist of massif reef limestones including a variety of microbialites associated with micro-encrusters. They played an important role in the formation and evolution of the reef frameworks and thus are of significant importance for deciphering the depositional environments. For our study, the most important encrusting organisms are Crescentiella morronensis, Koskinobullina socialis, Lithocodium aggregatum, Bacinella-type structures, Radiomura cautica, Perturbatacrusta leini, Coscinophragma sp., and crust-forming coralline sponges such as Calcistella. Based on microscopic observations, microbial contribution to reef construction is documented by the abundance of dense micrite, laminate structures, clotted, thrombolithic or peloidal microfabrics, constructive micritic cortices, biogenic encrustations and cement crusts, as well as by other types of microbial structures and crusts. Most of the investigated carbonate deposits can be classified as “coral-microbial-microencruster boundstones” which are characteristic for the Intra-Tethyan domain. Their paleogeographical significance is indicated by the presence of many features comparable with carbonate deposits of rimmed platform systems from the Northern Calcareous Alps or Central Apennines. Based on the distribution of the facies and facies associations within the carbonate sequences under study we can distinguish slope and external shelf margin environments. The microbial crusts, the encrusting micro-organisms, and in some cases the syndepositional cements have stabilized and bound the carbonates of the slope facies types. Subsequently, the stable substrate favored the installation of coral-microbial bioconstruction levels.     

Development of albian microbialites and microbialite reefs at marginal platform areas of the Vasco-Cantabrian Basin (Soba reef area,Cantabria, N. Spain)

The Middle Albian sequence from the western marginal area of the Vasco-Cantabrian Basin contains calcified microbialites in different marine depositional environments, individually well defined by microstructure, lamina characteristics and mode of formation. Microbialites may form the primary framework of reefs, which occur as composite stacks in mid to lower slope environments or as isolated bodies in small intraplatform basins. In most areas microbialite reef growth was initiated below the photic zone. Stratiform intercalations of microbialites and composite microbialite/foraminifer oncoids are restricted to well bedded carbonate platform deposits (Urgonian). Three basis types of microbialites are recognized:

Deep‐water microbialites of the Mesoproterozoic Dismal Lakes Group: microbial growth,lithification, and implications for coniform stromatolites

Offshore facies of the Mesoproterozoic Sulky Formation, Dismal Lakes Group, arctic Canada, preserve microbialites with unusual morphology. These microbialites grew in water depths greater than several tens of meters and correlate with high‐relief conical stromatolites of the more proximal September Lake reef complex. The gross morphology of these microbial facies consists of ridge‐like vertical supports draped by concave‐upward, subhorizontal elements, resulting in tent‐shaped cuspate microbialites with substantial primary void space. Morphological and petrographic analyses suggest a model wherein penecontemporaneous upward growth of ridge elements and development of subhorizontal draping elements initially resulted in a buoyantly supported, unlithified microbial form. Lithification began via precipitation within organic elements during microbialite growth. Mineralization either stabilized or facilitated collapse of initially neutrally buoyant microbialite forms. Microbial structures and breccias were then further stabilized by precipitation of marine herringbone cement. During late‐stage diagenesis, remaining void space was occluded by ferroan dolomite cement. Cuspate microbialites are most similar to those found in offshore facies of Neoarchean carbonate platforms and to unlithified, buoyantly supported microbial mats in modern ice‐covered Antarctic lakes. We suggest that such unusual microbialite morphologies are a product of the interaction between motile and non‐motile communities under nutrient‐limiting conditions, followed by early lithification, which served to preserve the resultant microbial form. The presence of marine herringbone cement, commonly associated with high dissolved inorganic carbon (DIC), low O₂ conditions, also suggests growth in association with reducing environments at or near the seafloor or in conjunction with a geochemical interface. Predominance of coniform stromatolite forms in the Proterozoic—across a variety of depositional environments—may thus reflect a combination of heterogeneous nutrient distribution, potentially driven by variable redox conditions, and an elevated carbonate saturation state, which permits preservation of these unusual microbialite forms.     

Late Smithian microbial deposits and their lateral marine fossiliferous limestones (Early Triassic,Hurricane Cliffs,Utah, USA)

Recurrent microbialite proliferations during the Early Triassic are usually explained by ecological relaxation and abnormal oceanic conditions. Most Early Triassic microbialites are described as single or multiple lithological units without detailed ecological information about lateral and coeval fossiliferous deposits. Exposed rocks along Workman Wash in the Hurricane Cliffs (southwestern Utah, USA) provide an opportunity to reconstruct the spatial relationships of late Smithian microbialites with adjacent and contemporaneous fossiliferous sediments. Microbialites deposited in an intertidal to subtidal interior platform are intercalated between inner tidal flat dolosiltstones and subtidal bioturbated fossiliferous limestones. Facies variations along these fossiliferous deposits and microbialites can be traced laterally over a few hundreds of meters. Preserved organisms reflect a moderately diversified assemblage, contemporaneous to the microbialite formation. The presence of such a fauna, including some stenohaline organisms (echinoderms), indicates that the development of these late Smithian microbial deposits occurred in normal-marine waters as a simple facies belt subject to relative sea-level changes. Based on this case study, the proliferation of microbialites cannot be considered as direct evidence for presumed harsh environmental conditions.     

Geospatial insights into the controls of microbialite formation at Laguna Negra,Argentina

Microbialites provide a record of the interaction of microorganisms with their environment constituting a record of microbial life and environments through geologic time. Our capacity to interpret this record is limited by an incomplete understanding of the microbial, geochemical, and physical processes that influence microbialite formation and morphogenesis. The modern system Laguna Negra in Catamarca Province, Argentina contains microbialites in a zone of carbonate precipitation associated with physico-chemical gradients and variable microbial community structure, making it an ideal location to study how these processes interact to drive microbialite formation. In this study, we investigated the geospatial relationships between carbonate morphology, geochemistry, and microbial community at the macro- (decimeter) to mega- (meter) scale by combining high-resolution imagery with field observations. We mapped the distribution of carbonate morphologies and allochtonously-derived volcaniclasts and correlated these with sedimentary matrices and geochemical parameters. Our work shows that the macroscale distribution of different carbonate morphologies spatially correlates with microbial mat distributions—a result consistent with previous microscale observations. Specifically, microbialitic carbonate morphologies more commonly occur associated with microbial mats while abiotically derived carbonate morphologies were less commonly associated with microbial mats. Spatial variability in the size and abundance of mineralized structures was also observed, however, the processes controlling this variability remains unclear and likely represent a combination of microbial, geochemical, and physical processes. Likewise, the processes controlling the spatial distribution of microbial mats at Laguna Negra are also unresolved. Our results suggest that in addition to the physical drivers observed in other modern environments, variability in the spatial distribution of microbialites and other carbonate morphologies at the macro- to megascale can be controlled by microbial processes. Overall, this study provides insight into the interpretation of microbialite occurrence and distributions in the geologic record and highlights the utility of geospatial statistics to probe the controls of microbialite formation in other environments.     

Phylogenetic Clustering of Origination and Extinction across the Late Ordovician Mass Extinction

Mass extinctions can have dramatic effects on the trajectory of life, but in some cases the effects can be relatively small even when extinction rates are high. For example, the Late Ordovician mass extinction is the second most severe in terms of the proportion of genera eliminated, yet is noted for the lack of ecological consequences and shifts in clade dominance. By comparison, the end-Cretaceous mass extinction was less severe but eliminated several major clades while some rare surviving clades diversified in the Paleogene. This disconnect may be better understood by incorporating the phylogenetic relatedness of taxa into studies of mass extinctions, as the factors driving extinction and recovery are thought to be phylogenetically conserved and should therefore promote both origination and extinction of closely related taxa. Here, we test whether there was phylogenetic selectivity in extinction and origination using brachiopod genera from the Middle Ordovician through the Devonian. Using an index of taxonomic clustering (R_CL) as a proxy for phylogenetic clustering, we find that A) both extinctions and originations shift from taxonomically random or weakly clustered within families in the Ordovician to strongly clustered in the Silurian and Devonian, beginning with the recovery following the Late Ordovician mass extinction, and B) the Late Ordovician mass extinction was itself only weakly clustered. Both results stand in stark contrast to Cretaceous-Cenozoic bivalves, which showed significant levels of taxonomic clustering of extinctions in the Cretaceous, including strong clustering in the mass extinction, but taxonomically random extinctions in the Cenozoic. The contrasting patterns between the Late Ordovician and end-Cretaceous events suggest a complex relationship between the phylogenetic selectivity of mass extinctions and the long-term phylogenetic signal in origination and extinction patterns.     

Genesis of microbialites as contemporaneous framework components of deglacial coral reefs, Tahiti (IODP 310)

Deglacial reefs from Tahiti (IODP 310) feature a co-occurrence of zooxanthellate corals with microbialites that compose up to 80 vol% of the reef framework. The notion that microbialites tend to form in more nutrient-rich environments has previously led to the concept that such encrustations are considerably younger than the coral framework, and that they have formed in deeper storeys of the reef edifice, or that they represent severe disturbances of the reef ecosystem. As indicated by their repetitive interbedding with coralline red algae, the microbialites of this reef succession of Tahiti, however, formed immediately after coral growth under photic conditions. Clearly, the deglacial reef microbialites present in the IODP 310 cores did not follow disturbances such as drowning or suffocation by terrestrial material, and are not “disaster forms”. Given that the corals and the microbialites developed in close spatial proximity, highly elevated nutrient levels caused by fluvial or groundwater transport from the volcanic hinterland are an unlikely cause for the exceptionally voluminous development of microbialites. That voluminous deglacial reef microbialites generally are restricted to volcanic islands, however, implies that moderately, and possibly episodically elevated nutrient levels favored this type of microbialite formation.     

A New Genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for Timing the Recovery of Brachiopoda After the End-Permian Mass Extinction

A new genus, Meishanorhynchia , is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic–Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1 ± 0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3 ± 0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4 ± 0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.     

Growth models of Bajocian coral-microbialite reefs of Chargey-lès-Port (eastern France): palaeoenvironmental interpretations

Very large amount of microbialites, up to 70% of the reef volume takes part in the edification of Lower Bajocian coral reefs in the Chargey-lès-Port quarry (Haute-Saône, France). Such high amounts of microbialites were unknown within bioconstructions of Middle Jurassic age. Along the 16 m-thick section, seven successive biohermal or biostromal units developed on a shallow platform. Bioconstructions display a first coral growth phase with either constratal or superstratal growth fabrics. Coral fauna is relatively poorly diversified and is dominated by massive forms (Isastrea, Thamnasteria, and Periseris) or branched phaceloid (Cladophyllia) and ramose (Dendraraea) colonies. Corals can be heavily encrusted by microbialites of diverse forms and fabrics (leiolitic, thrombolitic, and stromatolitic). According to the coral growth fabrics, microbialite crusts developed on top of or at the underside of coral colonies, forming a coral-microbialite elementary unit. Microbialites show a multiphase development: (i) directly at the coral surface, a first and mm-scale microbialite layer locally developed; (ii) a second, cm-scale microbialite layer (up to 8 cm thick) covered the entire coral reef framework and assumed the main building role; and (iii) a third, mm- to cm-scale, laminated microbialite layer may also be observed onlapping previous reef structures, before having been progressively buried under sediments. Contemporaneously to the coral growth phase, the first microbialite layer developed on dead portions of coral colonies. The transition between coral growth and microbialite development (i.e., second layer of microbialites) is interpreted as a result of a coral reef crisis, probably reflecting more nutrient-rich conditions. The passage to a stromatolitic (third) layer suggests a control of the accumulation rate. Composition and architecture of coral-microbialite reef units of Chargey-lès-Port highlight the relations between high-frequency fluctuating environmental factors (mainly accumulation rate and trophic conditions) and reef development.     

Palaeoecology Of A Late Devonian Back Reef: Canning Basin, Western Australia

Back‐reef ecologies within the celebrated mixed carbonate‐siliciclastic Late Devonian (late Frasnian) Pillara Limestone of Windjana Gorge, in the Canning Basin, Western Australia, are re‐interpreted as being dominated by microbial communities. Proposed microbialites are expressed as weakly‐laminated, fenestral micrite, that show unsupported primary voids, peloidal textures, disseminated bioclastic debris, and traces of microfilaments. These grew as either extensive free‐standing mounds or columns, often intergrown with encrusting metazoans, or thick post‐mortem encrustations upon skeletal benthos. In some cases, microbial encrustations are inferred to have developed in protected cavities formed by progressive burial of the reef. The calcimicrobe Shuguria also shows a preferentially cryptic habit, encrusting either primary cavities formed by skeletal benthos, microbialite, or the ceilings of mm‐sized fenestrae within microbialite. A further calcimicrobe, Rothpletzella, formed columns up to 0.3 m high in areas enriched by very coarse siliciclastic sediment. Stromatoporoid sponges with a diverse range of morphologies also formed in situ growth fabrics. Monospecific thickets of closely‐aggregating dendroid stromatoporoid sponges (Stachyodes costulata), and platy‐laminar forms (?Hermatostroma spp.) were common, as were remarkably large stromatoporoids (Actinostroma spp.) that grew as isolated individuals up to 5 m in diameter. Such sponges showed impressive powers of regeneration from partial mortality, and individual clones may have been capable of substantial longevities of up to 500 years. Actinostroma spp. showed highly complex growth forms including platy‐multicolumnar (A. windjanicum), and a hitherto undescribed inferred whorl‐forming foliaceous morphology (Actinostroma sp.) reminiscent of the modern photosymbiotic coral Acropora palmata. These complex morphologies formed abundant primary cavities, previously thought to be only rarely developed in association with stromatoporoids.key words : Late Devonian, Canning Basin, reefs, palaeoecology, microbialite.     

Microbialites and global environmental change across the Permian-Triassic boundary: a synthesis

Permian-Triassic boundary microbialites (PTBMs) are thin (0.05-15 m) carbonates formed after the end-Permian mass extinction. They comprise Renalcis-group calcimicrobes, microbially mediated micrite, presumed inorganic micrite, calcite cement (some may be microbially influenced) and shelly faunas. PTBMs are abundant in low-latitude shallow-marine carbonate shelves in central Tethyan continents but are rare in higher latitudes, likely inhibited by clastic supply on Pangaea margins. PTBMs occupied broadly similar environments to Late Permian reefs in Tethys, but extended into deeper waters. Late Permian reefs are also rich in microbes (and cements), so post-extinction seawater carbonate saturation was likely similar to the Late Permian. However, PTBMs lack widespread abundant inorganic carbonate cement fans, so a previous interpretation that anoxic bicarbonate-rich water upwelled to rapidly increase carbonate saturation of shallow seawater, post-extinction, is problematic. Preliminary pyrite framboid evidence shows anoxia in PTBM facies, but interbedded shelly faunas indicate oxygenated water, perhaps there was short-term pulsing of normally saturated anoxic water from the oxygen-minimum zone to surface waters. In Tethys, PTBMs show geographic variations: (i) in south China, PTBMs are mostly thrombolites in open shelf settings, largely recrystallised, with remnant structure of Renalcis-group calcimicrobes; (ii) in south Turkey, in shallow waters, stromatolites and thrombolites, lacking calcimicrobes, are interbedded, likely depth-controlled; and (iii) in the Middle East, especially Iran, stromatolites and thrombolites (calcimicrobes uncommon) occur in different sites on open shelves, where controls are unclear. Thus, PTBMs were under more complex control than previously portrayed, with local facies control playing a significant role in their structure and composition.     

Stromatolite-dominated microbialites at the Permian–Triassic boundary of the Xikou section on South Qinling Block,China

Permian–Triassic boundary microbialites (PTBMs) are organosedimentary carbonates formed immediately after the end-Permian mass extinction. All those reported PTBMs constrained by convincing conodont biozones are present stratigraphycally not higher than the Hindeodus parvus zone and most of them are dominated by thrombolites. This paper provides the first record of a brief, but spectacular development of stromatolite-dominated PTBMs within the basal Isarcicella isarcica conodont zone of the earliest Triassic from the Xikou section of South Qinling Block that was at the margin of the North China Block during the Permian–Triassic transition and was geographically separated from the major occurrence of post-extinction microbialites in the South China Block. This stromatolite cap overlies a 3.7-m-thick oolitic limestone and is composed of a lower 0.2-m-thick bed and an upper 0.5-m-thick bed, separated by a 0.2-m-thick greyish green siliciclastic mudstone. These two stromatolite beds mainly consist of columnar stromatolites with subordinate domal stromatolites. The intercolumn and interstitial spaces within the stromatolites are filled with oolitic grainstones. At the microscopic scale, laminoid structures in stromatolites comprise wavy, millimetric-domical and tangled laminae. The increased grain and fossil contents and/or bioturbation in the domical and tangled laminae indicate that the formation of these laminae is likely related to an increase in the populations and the disruptions by benthic metazoans, as well as an influx of sediment grains. The δ¹³C_carb values fluctuate between 2‰ and 3‰ in the uppermost Permian strata; a distinct negative shift of 1.9‰ occurs at the topmost oolitic grainstone, just below the lower stromatolite bed, and the lowest value of −0.1‰ is located at the base of the upper stromatolite bed. The stratigraphic succession from stromatolites to thrombolites of the PTBMs may represent a transgressive succession and/or a transient ecosystem recovery immediately after the end-Permian mass extinction. The thrombolites-dominated PTBMs mainly developed in near-equator shallow marine geographic locations, and stromatolite-dominated PTBMs mainly developed at higher latitude settings, which probably indicates that a relatively lower diversity and abundance of marine benthic metazoans existed at higher latitudes after the end-Permian mass extinction.     

Microbial response to limited nutrients in shallow water immediately after the end-Permian mass extinction

Previous work indicates that a variety of microbes bloomed in the oceans after the end-Permian faunal mass extinction, but evidence is sporadically documented. Thus, the nature and geographic distribution of such microbes and their associations are unclear, addressed in this study using a series of biomarker groups. On the basis of microbial biomarker records of the 2-methylhopane index, evidence is presented for cyanobacterial blooms in both the western and eastern Tethys Sea and in both shallow and deep waters, after the mass extinction. The enhanced relative abundance of C(28) (expressed by the C(28) /C(29) ratio of) regular steranes suggests a bloom of prasinophyte algae occurred immediately after the end-Permian faunal extinction, comparable with those observed in some other mass extinctions in Phanerozoic. Significantly, cyanobacteria and prasinophyte algae show a synchronized onset of bloom in the shallow water Bulla section, north Italy, inferring for the first time their coupled response to the biotic crisis and the associated environmental conditions. However, in Meishan of Zhejiang Province in south China, the bloom declined earlier than in Bulla. The association of increased 2-methylhopane index with a negative shift in the nitrogen isotope composition infers a scenario of enhanced nitrogen fixation by cyanobacteria immediately after the faunal mass extinction. N(2) fixation by cyanobacteria is here interpreted to have provided prasinophyte algae with ammonium in nutrient-limited shallow waters, and thus caused their associated blooms.     

Lipid biomarkers recording marine microbial community structure changes through the Frasnian-Famennian mass extinction event

Studying the response and recovery of marine microbial communities during mass extinction events provides an evolutionary window through which to understand the adaptation and resilience of the marine ecosystem in the face of significant environmental disturbances. The goal of this study is to reconstruct changes in the marine microbial community structure through the Late Devonian Frasnian-Famennian (F-F) transition. We performed a multiproxy investigation on a drill core of the Upper Devonian New Albany Shale from the Illinois Basin (western Kentucky, USA). Aryl isoprenoids show green sulfur bacteria expansion and associated photic zone euxinia (PZE) enhancement during the F-F interval. These changes can be attributed to augmented terrigenous influxes, as recorded collectively by the long-chain/short-chain normal alkane ratio, carbon preference index, C₃₀ moretane/C₃₀ hopane, and diahopane index. Hopane/sterane ratios reveal a more pronounced dominance of eukaryotic over prokaryotic production during the mass extinction interval. Sterane distributions indicate that the microalgal community was primarily composed of green algae clades, and their dominance became more pronounced during the F-F interval and continued to rise in the subsequent periods. The 2α-methylhopane index values do not show an evident shift during the mass extinction interval, whereas the 3β-methylhopane index values record a greater abundance of methanotrophic bacteria during the extinction interval, suggesting enhanced methane cycling due to intensified oxygen depletion. Overall, the Illinois Basin during the F-F extinction experienced heightened algal productivity due to intensified terrigenous influxes, exhibiting similarities to contemporary coastal oceans that are currently undergoing globalized cultural eutrophication. The observed microbial community shifts associated with the F-F environmental disturbances were largely restricted to the extinction interval, which suggests a relatively stable, resilient marine microbial ecosystem during the Late Devonian.