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.     

Mineralized microbialites as archives of environmental evolution,Laguna Negra,Catamarca Province,Argentina

Environmental fluctuations are recorded in a variety of sedimentary archives of lacustrine depositional systems. Geochemical signals recovered from bottom sediments in closed‐basin lakes are among the most sensitive paleoenvironmental indicators and are commonly used in reconstructing lake evolution. Microbialites (i.e., organosedimentary deposits accreted through microbial trapping and binding of detrital sediment or in situ mineral precipitation on organics [Palaios, 2, 1987, 241]), however, have been largely overlooked as paleoenvironmental repositories. Here, we investigate concentrically laminated mineralized microbialites from Laguna Negra, a high‐altitude (4,100 m above sea level) hypersaline, closed‐basin lake in northwestern Argentina, and explore the potential for recovery of environmental signals from these unique sedimentary archives. Spatial heterogeneity in hydrological regime helps define zones inside Laguna Negra, each with their own morphologically distinct microbialite type. Most notably, platey microbialites (in Zone 3A) are precipitated by evaporative concentration processes, while discoidal oncolites (in Zone 3C) are interpreted result from fluid mixing and biologically mediated nucleation. This spatial heterogeneity is reflected in petrographically distinct carbonate fabrics: micritic, botryoidal, and isopachous. Fabric type is interpreted to reflect a combination of physical and biological influences during mineralization, and paired C‐isotope measurement of carbonate and organic matter supports ecological differences as a dominant control on C‐isotopic evolution between zones. Laminae of Laguna Negra microbialites preserve a range of δ¹³C_carb from +5.75‰ to +18.25‰ and δ¹⁸O_carb from ?2.04‰ to +9.28‰. Temporal trends of lower carbon and oxygen isotopic compositions suggest that the influence of CO₂ degassing associated with evaporation has decreased over time. Combined, these results indicate that microbialite archives can provide data that aid in interpretation of both lake paleohydrology and paleoenvironmental change.     

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.     

Characterization of bacterial diversity associated with microbial mats,gypsum evaporites and carbonate microbialites in thalassic wetlands: Tebenquiche and La Brava,Salar de Atacama,Chile

In this paper, we report the presence of sedimentary microbial ecosystems in wetlands of the Salar de Atacama. These laminated systems, which bind, trap and precipitate mineral include: microbial mats at Laguna Tebenquiche and Laguna La Brava, gypsum domes at Tebenquiche and carbonate microbialites at La Brava. Microbial diversity and key biogeochemical characteristics of both lakes (La Brava and Tebenquiche) and their various microbial ecosystems (non-lithifying mats, flat and domal microbialites) were determined. The composition and abundance of minerals ranged from trapped and bound halite in organic-rich non-lithifying mats to aragonite-dominated lithified flat microbialites and gypsum in lithified domal structures. Pyrosequencing of the V4 region of the 16s rDNA gene showed that Proteobacteria comprised a major phylum in all of the microbial ecosystems studied, with a marked lower abundance in the non-lithifying mats. A higher proportion of Bacteroidetes was present in Tebenquiche sediments compared to La Brava samples. The concentration of pigments, particularly that of Chlorophyll a, was higher in the Tebenquiche than in La Brava. Pigments typically associated with anoxygenic phototrophic bacteria were present in lower amounts. Organic-rich, non-lithifying microbial mats frequently formed snake-like, bulbous structures due to gas accumulation underneath the mat. We hypothesize that the lithified microbialites might have developed from these snake-like microbial mats following mineral precipitation in the surface layer, producing domes with endoevaporitic communities in Tebenquiche and carbonate platforms in La Brava. Whereas the potential role of microbes in carbonate platforms is well established, the contribution of endoevaporitic microbes to formation of gypsum domes needs further investigation.     

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 diversity and biomarker analysis of modern freshwater microbialites from Laguna Bacalar,Mexico

Laguna Bacalar is a sulfate‐rich freshwater lake on the Yucatan Peninsula that hosts large microbialites. High sulfate concentrations distinguish Laguna Bacalar from other freshwater microbialite sites such as Pavilion Lake and Alchichica, Mexico, as well as from other aqueous features on the Yucatan Peninsula. While cyanobacterial populations have been described here previously, this study offers a more complete characterization of the microbial populations and corresponding biogeochemical cycling using a three‐pronged geobiological approach of microscopy, high‐throughput DNA sequencing, and lipid biomarker analyses. We identify and compare diverse microbial communities of Alphaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria that vary with location along a bank‐to‐bank transect across the lake, within microbialites, and within a neighboring mangrove root agglomeration. In particular, sulfate‐reducing bacteria are extremely common and diverse, constituting 7%–19% of phylogenetic diversity within the microbialites, and are hypothesized to significantly influence carbonate precipitation. In contrast, Cyanobacteria account for less than 1% of phylogenetic diversity. The distribution of lipid biomarkers reflects these changes in microbial ecology, providing meaningful biosignatures for the microbes in this system. Polysaturated short‐chain fatty acids characteristic of cyanobacteria account for <3% of total abundance in Laguna Bacalar microbialites. By contrast, even short‐chain and monounsaturated short‐chain fatty acids attributable to both Cyanobacteria and many other organisms including types of Alphaproteobacteria and Gammaproteobacteria constitute 43%–69% and 17%–25%, respectively, of total abundance in microbialites. While cyanobacteria are the largest and most visible microbes within these microbialites and dominate the mangrove root agglomeration, it is clear that their smaller, metabolically diverse associates are responsible for significant biogeochemical cycling in this microbialite system.     

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.     

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.     

Prokaryote populations of extant microbialites along a depth gradient in Pavilion Lake,British Columbia,Canada

Pavilion Lake in British Columbia, Canada, is home to modern‐day microbialites that are actively growing at multiple depths within the lake. While microbialite morphology changes with depth and previous isotopic investigations suggested a biological role in the formation of these carbonate structures, little is known about their microbial communities. Microbialite samples acquired through the Pavilion Lake Research Project (PLRP) were first investigated for phototrophic populations using Cyanobacteria‐specific primers and 16S rRNA gene cloning. These data were expounded on by high‐throughput tagged sequencing analyses of the general bacteria population. These molecular analyses show that the microbial communities of Pavilion Lake microbialites are diverse compared to non‐lithifying microbial mats also found in the lake. Phototrophs and heterotrophs were detected, including species from the recently described Chloroacidobacteria genus, a photoheterotroph that has not been previously observed in microbialite systems. Phototrophs were shown as the most influential contributors to community differences above and below 25 meters, and corresponding shifts in heterotrophic populations were observed at this interface as well. The isotopic composition of carbonate also mirrored this shift in community states. Comparisons to previous studies indicated this population shift may be a consequence of changes in lake chemistry at this depth. Microbial community composition did not correlate with changing microbialite morphology with depth, suggesting something other than community changes may be a key to observed variations in microbialite structure.     

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.     

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.     

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:

Microbialite response to an anthropogenic salinity gradient in Great Salt Lake,Utah

A railroad causeway across Great Salt Lake, Utah (GSL), has restricted water flow since its construction in 1959, resulting in a more saline North Arm (NA; 24%–31% salinity) and a less saline South Arm (SA; 11%–14% salinity). Here, we characterized microbial carbonates collected from the SA and the NA to evaluate the effect of increased salinity on community composition and abundance and to determine whether the communities present in the NA are still actively precipitating carbonate or if they are remnant features from prior to causeway construction. SSU rRNA gene abundances associated with the NA microbialite were three orders of magnitude lower than those associated with the SA microbialite, indicating that the latter community is more productive. SSU rRNA gene sequencing and functional gene microarray analyses indicated that SA and NA microbialite communities are distinct. In particular, abundant sequences affiliated with photoautotrophic taxa including cyanobacteria and diatoms that may drive carbonate precipitation and thus still actively form microbialites were identified in the SA microbialite; sequences affiliated with photoautotrophic taxa were in low abundance in the NA microbialite. SA and NA microbialites comprise smooth prismatic aragonite crystals. However, the SA microbialite also contained micritic aragonite, which can be formed as a result of biological activity. Collectively, these observations suggest that NA microbialites are likely to be remnant features from prior to causeway construction and indicate a strong decrease in the ability of NA microbialite communities to actively precipitate carbonate minerals. Moreover, the results suggest a role for cyanobacteria and diatoms in carbonate precipitation and microbialite formation in the SA of GSL.     

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.     

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.     

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.     

Microbialite concretions in a dolostone crust at the Permian–Triassic boundary of the Xishan section in Jiangsu Province,South China

Carbonate concretions with structures and fossil groups associated with microbialite developed in a dolostone crust at the Permian–Triassic boundary of the Xishan section in Jiangsu Province, South China. These structures include clotted fabrics and laminated carbonate needles, as well as abundant carbonate crystal fans. Fossil groups associated with microbialite include microconchids, small gastropods, and small foraminifers. These fabrics and fossils suggest that the concretions are carbonate microbialite blocks developed in the dolostone crust. On the basis of the analysis of the microfabrics and the fossil groups together with a comparison to modern analogues, we attribute the formation of the micritic patches in the microbialite concretions to the calcification of cyanobacterial mats via carbonate nanoparticles and we attribute the carbonate crystal fans to the direct recrystallization of micritic carbonates. The sparitic patches were interpreted as either the direct recrystallization of micritic carbonates or the precipitation of carbonate spars in the inter-/intra-spaces of metazoan shells together with the recrystallization of these shells. The similarities to modern stromatolites, both in morphology and in internal texture, suggest that the laminated carbonate needles are stromatolite laminae built by filamentous cyanobacteria. The preservation of these microbialite microfabrics indicates that early lithification by carbonate precipitation was widespread and intense following the end-Permian boundary events. The weak development of microbialites as small concretions may be attributed to the deeper water depth and the lower water energy in the Xishan area during the earliest Triassic.     

The Girvanella-like remains from Messinian marine deposits (Sardinia,Italy): Lagerstätten paradigm for microbial biota?

The lower Messinian marine sediments of the Capo San Marco Formation in the Sinis area (Sardinia, Italy) contain extensive carbonate buildups mainly made of microbialites. These microbialites exhibit general thrombolitic fabric and occur in meso-macroscopic scale as dominant cauliflower-like structures, digitations and encrusting rings. All the microbialites are here associated with serpulid tubes and bryozoan colonies. Examination of thin sections from microbialite samples reveal the presence of dense flexuous, not ramified and erect tubular micritic structures, with an external diameter ranging from 30 to 40 μm, all characters being very close to those of the Girvanella-type filaments. Although all microbialites show quite similar structural aspects, only two levels contain clearly visible networks of such filaments. The associated marine biota is diverse (cemented, borers, burrowers) related to the available biotopes (hard substrates, fine grained sediment, cavities…). The general scarcity of microbial remains in Messinian microbialites points out to the problem of taphonomic processes allowing a good preservation of microbial structures. The concept of Lagerstätten could well be extended to the preservation of microscopic organisms in the carbonate material. The discovery of Girvanella-like filaments demonstrates the implication of cyanobacterial organisms in the construction processes of the Messinian thrombolitic buildups. Furthermore, it is the first time that Girvanella-like microbialites are documented from Upper Miocene marine rocks.     

Photosynthesis versus Exopolymer Degradation in the Formation of Microbialites on the Atoll of Kiritimati,Republic of Kiribati,Central Pacific

Aragonitic microbialites, characterized by a reticulate fabric, were discovered beneath lacustrine microbial mats on the atoll of Kiritimati, Republic of Kiribati, Central Pacific. The microbial mats, with cyanobacteria as major primary producers, grow in evaporated seawater modified by calcium carbonate and gypsum precipitation and calcium influx via surface and/or groundwaters. Despite the high aragonite supersaturation and a high photosynthetic activity, only minor aragonite precipitates are observed in the top parts of the microbial mats. Instead, major aragonite precipitation takes place in lower mat parts at the transition to the anoxic zone. The prokaryotic community shows a high number of phylotypes closely related to halotolerant taxa and/or taxa with preference to oligotrophic habitats. Soil- and plant- inhabiting bacteria underline a potential surface or subsurface influx from terrestrial areas, while chitinase-producing representatives coincide with the occurrence of insect remains in the mats. Strikingly, many of the clones have their closest relatives in microorganisms either involved in methane production or consumption of methane or methyl compounds. Methanogens, represented by the methylotrophic genus Methanohalophilus, appear to be one of the dominant organisms in anaerobic mat parts. All this points to a significant role of methane and methyl components in the carbon cycle of the mats. Nonetheless, thin sections and physicochemical gradients through the mats, as well as the ¹²C-depleted carbon isotope signatures of carbonates indicate that spherulitic components of the microbialites initiate in the photosynthesis-dominated orange mat top layer, and further grow in the green and purple layer below. Therefore, these spherulites are considered as product of an extraordinary high photosynthesis effect simultaneous to a high inhibition by pristine exopolymers. Then, successive heterotrophic bacterial activity leads to a condensation of the exopolymer framework, and finally to the formation of crevice-like zones of partly degraded exopolymers. Here initiation of horizontal aragonite layers and vertical aragonite sheets of the microbialite occurs, which are considered as a product of high photosynthesis at decreasing degree of inhibition. Finally, at low supersaturation and almost lack of inhibition, syntaxial growth of aragonite crystals at lamellae surfaces leads to thin fibrous aragonite veneers. While sulfate reduction, methylotrophy, methanogenesis and ammonification play an important role in element cycling of the mat, there is currently no evidence for a crucial role of them in CaCO₃ precipitation. Instead, photosynthesis and exopolymer degradation sufficiently explain the observed pattern and fabric of microbialite formation.     

Legacies of recent environmental change in the benthic communities of Lake Joyce, a perennially ice-covered Antarctic lake

Many Antarctic lakes provide habitat for extensive microbial mats that respond on various timescales to environmental change. Lake Joyce contains calcifying microbialites and provides a natural laboratory to constrain how environmental changes influence microbialite development. In Lake Joyce, depth-specific distributions of calcitic microbialites, organic carbon, photosynthetic pigments and photosynthetic potential cannot be explained by current growth conditions, but are a legacy of a 7-m lake level rise between 1973 and 2009. In the well-illuminated margins of the lake, photosynthetically active benthic communities colonised surfaces submerged for just a few years. However, observed increases in accumulated organic material with depth from 5 to 20 m (2-40 mg ash-free dry weight cm(-2)) and the presence of decimetre-scale calcite microbialites at 20-22 m depth, apparently related to in situ photosynthetic growth, are inconsistent with the current distributions of irradiance, photosynthetic pigments and mat photosynthetic potential (as revealed by pulse-amplitude-modulated fluorometry). The microbialites appeared photosynthetically active in 1986 and 1997, but were outside the depth zone where significant phototrophic growth was possible and were weakly photosynthetically competent in 2009. These complex microbial structures have persisted after growth has ceased, demonstrating how fluctuating environmental conditions and the hysteresis between environmental change, biological response and microbialite development can be important factors to consider when interpreting modern, and by inference ancient, microbially mediated structures.