The importance of groundwater flow to the formation of modern thrombolitic microbialites

Modern microbialites are often located within groundwater discharge zones, yet the role of groundwater in microbialite accretion has yet to be resolved. To understand relationships between groundwater, microbialites, and associated microbial communities, we quantified and characterized groundwater flow and chemistry in active thrombolitic microbialites in Lake Clifton, Western Australia, and compared these observations to inactive thrombolites and lakebed sediments. Groundwater flows upward through an interconnected network of pores within the microstructure of active thrombolites, discharging directly from thrombolite heads into the lake. This upwelling groundwater is fresher than lake water and is hypothesized to support microbial mat growth by reducing salinity and providing limiting nutrients in an osmotically stressful and oligotrophic habitat. This is in contrast to inactive thrombolites that show no evidence of microbial mat colonization and are infiltrated by hypersaline lake water. Groundwater discharge through active thrombolites contrasts with the surrounding lakebed, where hypersaline lake water flows downward through sandy sediments at very low rates. Based on an appreciation for the role of microorganisms in thrombolite accretion, our findings suggest conditions favorable to thrombolite formation still exist in certain locations of Lake Clifton despite increasing lake water salinity. This study is the first to characterize groundwater flow rates, paths, and chemistry within a microbialite‐forming environment and provides new insight into how groundwater can support microbial mats believed to contribute to microbialite formation in modern and ancient environments.     

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.     

Coexisting living stromatolites and infaunal metazoans

Microbialites, bioaccretionary structures formed during the growth and metabolism of microorganisms (principally cyanobacteria) were the dominant lifeform in shallow late-Archean and Proterozoic oceans. During the Cambrian radiation of metazoan life, which began ~540 Mya, microbialite abundance and diversity further declined following a peak in the Mesoproterozoic. Notwithstanding contention, grazing and bioturbation effects of metazoans have been hypothesized as the dominant driver of modern microbialite scarcity. However, this metazoan–microbialite exclusion has not been fully explored in the few extant microbialites. Here we provide further evidence showing that living marine layered microbialites (stromatolites) coexist with a persistent assemblage of benthic macro-invertebrates, as has previously been demonstrated in some thrombolitic (clotted) microbialites. Surprisingly, these metazoans have active habits, such as burrowing, which should be expected to disrupt the layered matrix. As other studies have shown, through a network of burrows, metazoans can exploit local diurnal oxygen refugia within microbialites as well as escape predation. Our results, therefore, add novel evidence in support of the hypotheses that geologically, metazoans are not always incompatible with stromatolites, while ecologically, microbialites may act as micro-refugia for modern metazoans and historically have performed a similar inferred role in past ecosystems.     

Lacustrine Nostoc (Nostocales) and associated microbiome generate a new type of modern clotted microbialite

Microbialites are mineral formations formed by microbial communities that are often dominated by cyanobacteria. Carbonate microbialites, known from Proterozoic times through the present, are recognized for sequestering globally significant amounts of inorganic carbon. Recent ecological work has focused on microbial communities dominated by cyanobacteria that produce microbial mats and laminate microbialites (stromatolites). However, the taxonomic composition and functions of microbial communities that generate distinctive clotted microbialites (thrombolites) are less well understood. Here, microscopy and deep shotgun sequencing were used to characterize the microbiome (microbial taxa and their genomes) associated with a single cyanobacterial host linked by 16S sequences to Nostoc commune Vaucher ex Bornet & Flahault, which dominates abundant littoral clotted microbialites in shallow, subpolar, freshwater Laguna Larga in southern Chile. Microscopy and energy‐dispersive X‐ray spectroscopy suggested the hypothesis that adherent hollow carbonate spheres typical of the clotted microbialite begin development on the rigid curved outer surfaces of the Nostoc balls. A surface biofilm included >50 nonoxygenic bacterial genera (taxa other than Nostoc) that indicate diverse ecological functions. The Laguna Larga Nostoc microbiome included the sulfate reducers Desulfomicrobium and Sulfospirillum and genes encoding all known proteins specific to sulfate reduction, a process known to facilitate carbonate deposition by increasing pH. Sequences indicating presence of nostocalean and other types of nifH, nostocalean sulfide:ferredoxin oxidoreductase (indicating anoxygenic photosynthesis), and biosynthetic pathways for the secondary products scytonemin, mycosporine, and microviridin toxin were identified. These results allow comparisons with microbiota and microbiomes of other algae and illuminate biogeochemical roles of ancient microbialites.     

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.     

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.     

Modern cryptic microbialite/metazoan facies from Lizard Island (Great Barrier Reef,Australia) formation and concepts

Summary From shallow water caves of fringing reefs related to continental islands of the Lizard Island Section thrombolitic micritic microbialites were observed. The microbialites exhibit always a light decreasing facies succession. The succession starts with a coralgal community and ends with light independent microbial biofilms and benthos (coralline sponges). The sessile mineralized benthos community is constructed of crustose foraminifera, serpulids, thecidean brachiopods, bryozoans, and coralline sponges. The observed benthic community is very similar to those one observed in cryptic habitates of Aptian and Albian reefs of northern Spain. For longtime studies of the microbialite formation and growth rates of coralline sponges the specimens were stained in vivo, within their natural habitat with histochemical fluorochromes and nonfluorescent agents. Main results are a very slow growth of the microbialite and associated sponges (50–100 μm/y). Only few calcifying microbes are participators during microbialite formation. Calcifying acidic organic macromolecules are mainly responsible for microbialite formation by cementing detritical material. Fe/Mn-bacterial biofilms are responsible for strong corrosion of the microbialite. Beside the corrosive activity of the Fe/Mn-bacterial biofilms boring sponges (Aka, Cliona) are the main destructors. Geochemically the observed microbialites are composed of mainly high-Mg calcites and exhibit high positive δ¹³C (+3 to +4) values.     

Molecular indicators of microbial diversity in oolitic sands of Highborne Cay,Bahamas

Microbialites (stromatolites and thrombolites) are mineralized mat structures formed via the complex interactions of diverse microbial‐mat communities. At Highborne Cay, in the Bahamas, the carbonate component of these features is mostly comprised of ooids. These are small, spherical to ellipsoidal grains characterized by concentric layers of calcium carbonate and organic matter and these sand‐sized particles are incorporated with the aid of extra‐cellular polymeric substances (EPS), into the matrix of laminated stromatolites and clotted thrombolite mats. Here, we present a comparison of the bacterial diversity within oolitic sand samples and bacterial diversity previously reported in thrombolitic and stromatolitic mats of Highborne Cay based on analysis of clone libraries of small subunit ribosomal RNA gene fragments and lipid biomarkers. The 16S‐rRNA data indicate that the overall bacterial diversity within ooids is comparable to that found within thrombolites and stromatolites of Highborne Cay, and this significant overlap in taxonomic groups suggests that ooid sands may be a source for much of the bacterial diversity found in the local microbialites. Cyanobacteria were the most diverse taxonomic group detected, followed by Alphaproteobacteria, Gammaproteobacteria, Planctomyces, Deltaproteobacteria, and several other groups also found in mat structures. The distributions of intact polar lipids, the fatty acids derived from them, and bacteriohopanepolyols provide broad general support for the bacterial diversity identified through analysis of nucleic acid clone libraries.     

Comparative Characterization of the Microbial Diversities of an Artificial Microbialite Model and a Natural Stromatolite

Microbialites are organosedimentary structures that result from the trapping, binding, and lithification of sediments by microbial mat communities. In this study we developed a model artificial microbialite system derived from natural stromatolites, a type of microbialite, collected from Exuma Sound, Bahamas. We demonstrated that the morphology of the artificial microbialite was consistent with that of the natural system in that there was a multilayer community with a pronounced biofilm on the surface, a concentrated layer of filamentous cyanobacteria in the top 5 mm, and a lithified layer of fused oolitic sand grains in the subsurface. The fused grain layer was comprised predominantly of the calcium carbonate polymorph aragonite, which corresponded to the composition of the Bahamian stromatolites. The microbial diversity of the artificial microbialites and that of natural stromatolites were also compared using automated ribosomal intergenic spacer analysis (ARISA) and 16S rRNA gene sequencing. The ARISA profiling indicated that the Shannon indices of the two communities were comparable and that the overall diversity was not significantly lower in the artificial microbialite model. Bacterial clone libraries generated from each of the three artificial microbialite layers and natural stromatolites indicated that the cyanobacterial and crust layers most closely resembled the ecotypes detected in the natural stromatolites and were dominated by Proteobacteria and Cyanobacteria. We propose that such model artificial microbialites can serve as experimental analogues for natural stromatolites.     

Palaeoecology of microconchids from microbialites near the Permian–Triassic boundary in South China

Abundant isolated specimens of microconchid tubes have been extracted from a microbialite deposit near the Permian–Triassic boundary (PTB) in the Dajiang section, southern Guizhou Province, South China. They are assignable to Microconchus aff. utahensis, M. aff. aberrans and Helicoconchus aff. elongatus, all of which possess micro‐lamellar tube walls. Quantitative analysis of bulk samples indicates that most microconchids occur in the upper part of the PTB microbialite deposit and show substrate selectivity for bioclastic grainstone–packstones. In contrast, very few microconchids were found in the rocks bearing well‐developed microbialite structures. Both stratigraphical and substrate preferences indicate proliferation of microconchids coincided with an ebb of microbialite development. Microconchids therefore only proliferated in local niches in which microbial activities were not very active within the PTB microbialite ecosystem. The presence of abundant microconchids further strengthens the impression that PTB microbialite metazoans are much more diverse than previously thought. The end‐Permian mass extinction is calibrated to the base of microbialite deposit in South China. Thus, abundant microbialite metazoans, such as ostracods, lingulid brachiopods, microgastropods and microconchids, together with the considerable, temporarily surviving faunas reported from non‐microbialite PTB sections in South China, indicate that metazoans diversified immediately after the first episode of the end‐Permian mass extinction, supporting the scenario that marine ecosystems underwent episodic collapses during the devastating biocrisis over the Permian–Triassic transition.     

Carbonate fabrics in the modern microbialites of Pavilion Lake: two suites of microfabrics that reflect variation in microbial community morphology,growth habit,and lithification

Modern microbialites in Pavilion Lake, BC, provide an analog for ancient non‐stromatolitic microbialites that formed from in situ mineralization. Because Pavilion microbialites are mineralizing under the influence of microbial communities, they provide insights into how biological processes influence microbialite microfabrics and mesostructures. Hemispherical nodules and micrite–microbial crusts are two mesostructures within Pavilion microbialites that are directly associated with photosynthetic communities. Both filamentous cyanobacteria in hemispherical nodules and branching filamentous green algae in micrite–microbial crusts were associated with calcite precipitation at microbialite surfaces and with characteristic microfabrics in the lithified microbialite. Hemispherical nodules formed at microbialite surfaces when calcite precipitated around filamentous cyanobacteria with a radial growth habit. The radial filament pattern was preserved within the microbialite to varying degrees. Some subsurface nodules contained well‐defined filaments, whereas others contained only dispersed organic inclusions. Variation in filament preservation is interpreted to reflect differences in timing and amount of carbonate precipitation relative to heterotrophic decay, with more defined filaments reflecting greater lithification prior to degradation than more diffuse filaments. Micrite–microbial crusts produce the second suite of microfabrics and form in association with filamentous green algae oriented perpendicular to the microbialite surface. Some crusts include calcified filaments, whereas others contained voids that reflect the filamentous community in shape, size, and distribution. Pavilion microbialites demonstrate that microfabric variation can reflect differences in lithification processes and microbial metabolisms as well as microbial community morphology and organization. Even when the morphology of individual filaments or cells is not well preserved, the microbial growth habit can be captured in mesoscale microbialite structures. These results suggest that when petrographic preservation is extremely good, ancient microbialite growth structures and microfabrics can be interpreted in the context of variation in community organization, community composition, and lithification history. Even in the absence of distinct microbial microfabrics, mesostructures can capture microbial community morphology.     

Metagenomic and stable isotopic analyses of modern freshwater microbialites in Cuatro Ciénegas, Mexico

Ancient biologically mediated sedimentary carbonate deposits, including stromatolites and other microbialites, provide insight into environmental conditions on early Earth. The primary limitation to interpreting these records is our lack of understanding regarding microbial processes and the preservation of geochemical signatures in contemporary microbialite systems. Using a combination of metagenomic sequencing and isotopic analyses, this study describes the identity, metabolic potential and chemical processes of microbial communities from living microbialites from Cuatro Ciénegas, Mexico. Metagenomic sequencing revealed a diverse, redox-dependent microbial community associated with the microbialites. The microbialite community is distinct from other marine and freshwater microbial communities, and demonstrates extensive environmental adaptation. The microbialite metagenomes contain a large number of genes involved in the production of exopolymeric substances and the formation of biofilms, creating a complex, spatially structured environment. In addition to the spatial complexity of the biofilm, microbial activity is tightly controlled by sensory and regulatory systems, which allow for coordination of autotrophic and heterotrophic processes. Isotopic measurements of the intracrystalline organic matter demonstrate the importance of heterotrophic respiration of photoautotrophic biomass in the precipitation of calcium carbonate. The genomic and stable isotopic data presented here significantly enhance our evolving knowledge of contemporary biomineralization processes, and are directly applicable to studies of ancient microbialites.     

Evidence of oxygenic phototrophy in ancient phosphatic stromatolites from the Paleoproterozoic Vindhyan and Aravalli Supergroups,India

Fossil microbiotas are rare in the early rock record, limiting the type of ecological information extractable from ancient microbialites. In the absence of body fossils, emphasis may instead be given to microbially derived features, such as microbialite growth patterns, microbial mat morphologies, and the presence of fossilized gas bubbles in lithified mats. The metabolic affinity of micro‐organisms associated with phosphatization may reveal important clues to the nature and accretion of apatite‐rich microbialites. Stromatolites from the 1.6 Ga Chitrakoot Formation (Semri Group, Vindhyan Supergroup) in central India contain abundant fossilized bubbles interspersed within fine‐grained in situ‐precipitated apatite mats with average δ¹³C_org indicative of carbon fixation by the Calvin cycle. In addition, the mats hold a synsedimentary fossil biota characteristic of cyanobacterial and rhodophyte morphotypes. Phosphatic oncoid cone‐like stromatolites from the Paleoproterozoic Aravalli Supergroup (Jhamarkotra Formation) comprise abundant mineralized bubbles enmeshed within tufted filamentous mat fabrics. Construction of these tufts is considered to be the result of filamentous bacteria gliding within microbial mats, and as fossilized bubbles within pristine mat laminae can be used as a proxy for oxygenic phototrophy, this provides a strong indication for cyanobacterial activity in the Aravalli mounds. We suggest that the activity of oxygenic phototrophs may have been significant for the formation of apatite in both Vindhyan and Aravalli stromatolites, mainly by concentrating phosphate and creating steep diurnal redox gradients within mat pore spaces, promoting apatite precipitation. The presence in the Indian stromatolites of alternating apatite‐carbonate lamina may result from local variations in pH and oxygen levels caused by photosynthesis–respiration in the mats. Altogether, this study presents new insights into the ecology of ancient phosphatic stromatolites and warrants further exploration into the role of oxygen‐producing biotas in the formation of Paleoproterozoic shallow‐basin phosphorites.     

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.     

Reconstruction of limnology and microbialite formation conditions from carbonate clumped isotope thermometry

Quantitative tools for deciphering the environment of microbialite formation are relatively limited. For example, the oxygen isotope carbonate‐water geothermometer requires assumptions about the isotopic composition of the water of formation. We explored the utility of using ‘clumped’ isotope thermometry as a tool to study the temperatures of microbialite formation. We studied microbialites recovered from water depths of 10–55 m in Pavilion Lake, and 10–25 m in Kelly Lake, spanning the thermocline in both lakes. We determined the temperature of carbonate growth and the ¹⁸O/¹⁶O ratio of the waters that microbialites grew in. Results were then compared to current limnological data from the lakes to reconstruct the history of microbialite formation. Modern microbialites collected at shallow depths (11.7 m) in both lakes yield clumped isotope‐based temperatures of formation that are within error of summer water temperatures, suggesting that clumped isotope analyses may be used to reconstruct past climates and to probe the environments in which microbialites formed. The deepest microbialites (21.7–55 m) were recovered from below the present‐day thermoclines in both lakes and yield radioisotope ages indicating they primarily formed earlier in the Holocene. During this time, pollen data and our reconstructed water ¹⁸O/¹⁶O ratios indicate a period of aridity, with lower lake levels. At present, there is a close association between both photosynthetic and heterotrophic communities, and carbonate precipitation/microbialite formation, with biosignatures of photosynthetic influences on carbonate detected in microbialites from the photic zone and above the thermocline (i.e., depths of generally <20 m). Given the deeper microbialites are receiving <1% of photosynthetically active radiation (PAR), it is likely these microbialites primarily formed when lower lake levels resulted in microbialites being located higher in the photic zone, in warm surface waters.     

Patterns of metal distribution in hypersaline microbialites during early diagenesis: Implications for the fossil record

The use of metals as biosignatures in the fossil stromatolite record requires understanding of the processes controlling the initial metal(loid) incorporation and diagenetic preservation in living microbialites. Here, we report the distribution of metals and the organic fraction within the lithifying microbialite of the hypersaline Big Pond Lake (Bahamas). Using synchrotron‐based X‐ray microfluorescence, confocal, and biphoton microscopies at different scales (cm–μm) in combination with traditional geochemical analyses, we show that the initial cation sorption at the surface of an active microbialite is governed by passive binding to the organic matrix, resulting in a homogeneous metal distribution. During early diagenesis, the metabolic activity in deeper microbialite layers slows down and the distribution of the metals becomes progressively heterogeneous, resulting from remobilization and concentration as metal(loid)‐enriched sulfides, which are aligned with the lamination of the microbialite. In addition, we were able to identify globules containing significant Mn, Cu, Zn, and As enrichments potentially produced through microbial activity. The similarity of the metal(loid) distributions observed in the Big Pond microbialite to those observed in the Archean stromatolites of Tumbiana provides the foundation for a conceptual model of the evolution of the metal distribution through initial growth, early diagenesis, and fossilization of a microbialite, with a potential application to the fossil record.     

Active eukaryotes in microbialites from Highborne Cay,Bahamas, and Hamelin Pool (Shark Bay), Australia

Microbialites are organosedimentary structures that are formed through the interaction of benthic microbial communities and sediments and include mineral precipitation. These lithifying microbial mat structures include stromatolites and thrombolites. Exuma Sound in the Bahamas, and Hamelin Pool in Shark Bay, Western Australia, are two locations where significant stands of modern microbialites exist. Although prokaryotic diversity in these structures is reasonably well documented, little is known about the eukaryotic component of these communities and their potential to influence sedimentary fabrics through grazing, binding and burrowing activities. Accordingly, comparisons of eukaryotic communities in modern stromatolitic and thrombolitic mats can potentially provide insight into the coexistence of both laminated and clotted mat structures in close proximity to one another. Here we examine this possibility by comparing eukaryotic diversity based on Sanger and high-throughput pyrosequencing of small subunit ribosomal RNA (18S rRNA) genes. Analyses were based on total RNA extracts as template to minimize input from inactive or deceased organisms. Results identified diverse eukaryotic communities particularly stramenopiles, Alveolata, Metazoa, Amoebozoa and Rhizaria within different mat types at both locations, as well as abundant and diverse signatures of eukaryotes with <80% sequence similarity to sequences in GenBank. This suggests the presence of significant novel eukaryotic diversity, particularly in hypersaline Hamelin Pool. There was evidence of vertical structuring of protist populations and foraminiferal diversity was highest in bioturbated/clotted thrombolite mats of Highborne Cay.     

Diverse earliest Triassic ostracod fauna of the non‐microbialite‐bearing shallow marine carbonates of the Yangou section,South China

Since diverse ostracod faunas in the immediate aftermath of the latest Permian mass extinction are mainly found within Permian–Triassic boundary microbialites (PTBMs), the idea of an ostracod ‘microbial‐related refuge’ has been proposed. Here, we report a diversified earliest Triassic ostracod fauna from the Yangou section in South China, where no PTBMs were deposited, providing evidence inconsistent with this ‘microbial‐related refuge’ hypothesis. In addition, a significant ostracod extinction is recorded, corresponding with the earliest Triassic mass extinction (ETME). This ETME of ostracods is associated with size increases and a length/height ratio (L/H) decrease, indicating varied evolutionary patterns of shape and size of ostracods through the Permian–Triassic (P‐Tr) extinction events. Although the nature of these biotic changes is somewhat unclear, the temporally varied ‘refuge zone’ scenario provides us with a window to reconstruct the environmental dynamics of ecosystem changes during the P‐Tr transition.     

Microbes and mass extinctions: paleoenvironmental distribution of microbialites during times of biotic crisis

Widespread development of microbialites characterizes the substrate and ecological response during the aftermath of two of the 'big five' mass extinctions of the Phanerozoic. This study reviews the microbial response recorded by macroscopic microbial structures to these events to examine how extinction mechanism may be linked to the style of microbialite development. Two main styles of response are recognized: (i) the expansion of microbialites into environments not previously occupied during the pre-extinction interval and (ii) increases in microbialite abundance and attainment of ecological dominance within environments occupied prior to the extinction. The Late Devonian biotic crisis contributed toward the decimation of platform margin reef taxa and was followed by increases in microbialite abundance in Famennian and earliest Carboniferous platform interior, margin, and slope settings. The end-Permian event records the suppression of infaunal activity and an elimination of metazoan-dominated reefs. The aftermath of this mass extinction is characterized by the expansion of microbialites into new environments including offshore and nearshore ramp, platform interior, and slope settings. The mass extinctions at the end of the Triassic and Cretaceous have not yet been associated with a macroscopic microbial response, although one has been suggested for the end-Ordovician event. The case for microbialites behaving as 'disaster forms' in the aftermath of mass extinctions accurately describes the response following the Late Devonian and end-Permian events, and this may be because each is marked by the reduction of reef communities in addition to a suppression of bioturbation related to the development of shallow-water anoxia.