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.     

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.     

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.     

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 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.     

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.     

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.     

Isotopic biosignatures in carbonate‐rich,cyanobacteria‐dominated microbial mats of the Cariboo Plateau,B.C.

Photosynthetic activity in carbonate‐rich benthic microbial mats located in saline, alkaline lakes on the Cariboo Plateau, B.C. resulted in pCO₂ below equilibrium and δ¹³C_DIC values up to +6.0‰ above predicted carbon dioxide (CO₂) equilibrium values, representing a biosignature of photosynthesis. Mat‐associated δ¹³C_carb values ranged from ~4 to 8‰ within any individual lake, with observations of both enrichments (up to 3.8‰) and depletions (up to 11.6‰) relative to the concurrent dissolved inorganic carbon (DIC). Seasonal and annual variations in δ¹³C values reflected the balance between photosynthetic ¹³C‐enrichment and heterotrophic inputs of ¹³C‐depleted DIC. Mat microelectrode profiles identified oxic zones where δ¹³C_carb was within 0.2‰ of surface DIC overlying anoxic zones associated with sulphate reduction where δ¹³C_carb was depleted by up to 5‰ relative to surface DIC reflecting inputs of ¹³C‐depleted DIC. δ¹³C values of sulphate reducing bacteria biomarker phospholipid fatty acids (PLFA) were depleted relative to the bulk organic matter by ~4‰, consistent with heterotrophic synthesis, while the majority of PLFA had larger offsets consistent with autotrophy. Mean δ¹³C_org values ranged from ?18.7 ± 0.1 to ?25.3 ± 1.0‰ with mean Δ¹³C_inorg‐org values ranging from 21.1 to 24.2‰, consistent with non‐CO₂‐limited photosynthesis, suggesting that Precambrian δ¹³C_org values of ~?26‰ do not necessitate higher atmospheric CO₂ concentrations. Rather, it is likely that the high DIC and carbonate content of these systems provide a non‐limiting carbon source allowing for expression of large photosynthetic offsets, in contrast to the smaller offsets observed in saline, organic‐rich and hot spring microbial mats.     

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.     

Microbially influenced lacustrine carbonates: A comparison of Late Quaternary Lahontan tufa and modern thrombolite from Fayetteville Green Lake,NY

Carbonate microbialites in lakes can serve as valuable indicators of past environments, so long as the biogenicity and depositional setting of the microbialite can be accurately determined. Late Pleistocene to Early Holocene frondose draping tufa deposits from Winnemucca Dry Lake (Nevada, USA), a subbasin of pluvial Lake Lahontan, were examined in outcrop, petrographically, and geochemically to determine whether microbially induced precipitation is a dominant control on deposition. These observations were compared to modern, actively accumulating microbialites from Fayetteville Green Lake (New York, USA) using similar methods. In addition, preserved microbial DNA was extracted from the Lahontan tufa and sequenced to provide a more complete picture of the microbial communities. Tufas are texturally and geochemically similar to modern thrombolitic microbialites from Fayetteville Green Lake, and the stable isotopic composition of organic C, N, inorganic C, and O supports deposition associated with a lacustrine microbial mat environment dominated by photosynthetic processes. DNA extraction and sequencing indicate that photosynthetic microbial builders were present during tufa deposition, primarily Chloroflexi and Proteobacteria with minor abundances of Cyanobacteria and Acidobacteria. Based on the sequencing results, the depositional environment of the tufas can be constrained to the photic zone of the lake, contrasting with some previous interpretations that put tufa formation in deeper waters. Additionally, the presence of a number of mesothermophilic phyla, including Deinococcus–Thermus, indicates that thermal groundwater may have played a role in tufa deposition at sites not previously associated with groundwater influx. The interpretation of frondose tufas as microbially influenced deposits provides new context to interpretations of lake level and past environments in the Lahontan lake basins.     

Carbonate facies‐specific stable isotope data record climate,hydrology, and microbial communities in Great Salt Lake,UT

Organic and inorganic stable isotopes of lacustrine carbonate sediments are commonly used in reconstructions of ancient terrestrial ecosystems and environments. Microbial activity and local hydrological inputs can alter porewater chemistry (e.g., pH, alkalinity) and isotopic composition (e.g., δ¹⁸O_water, δ¹³C_DIC), which in turn has the potential to impact the stable isotopic compositions recorded and preserved in lithified carbonate. The fingerprint these syngenetic processes have on lacustrine carbonate facies is yet unknown, however, and thus, reconstructions based on stable isotopes may misinterpret diagenetic records as broader climate signals. Here, we characterize geochemical and stable isotopic variability of carbonate minerals, organic matter, and water within one modern lake that has known microbial influences (e.g., microbial mats and microbialite carbonate) and combine these data with the context provided by 16S rRNA amplicon sequencing community profiles. Specifically, we measure oxygen, carbon, and clumped isotopic compositions of carbonate sediments (δ¹⁸O_carb, δ¹³C_carb, ?₄₇), as well as carbon isotopic compositions of bulk organic matter (δ¹³C_org) and dissolved inorganic carbon (DIC; δ¹³C_DIC) of lake and porewater in Great Salt Lake, Utah from five sites and three seasons. We find that facies equivalent to ooid grainstones provide time‐averaged records of lake chemistry that reflect minimal alteration by microbial activity, whereas microbialite, intraclasts, and carbonate mud show greater alteration by local microbial influence and hydrology. Further, we find at least one occurrence of ?₄₇ isotopic disequilibrium likely driven by local microbial metabolism during authigenic carbonate precipitation. The remainder of the carbonate materials (primarily ooids, grain coatings, mud, and intraclasts) yield clumped isotope temperatures (T(?₄₇)), δ¹⁸O_carb, and calculated δ¹⁸O_water in isotopic equilibrium with ambient water and temperature at the time and site of carbonate precipitation. Our findings suggest that it is possible and necessary to leverage diverse carbonate facies across one sedimentary horizon to reconstruct regional hydroclimate and evaporation–precipitation balance, as well as identify microbially mediated carbonate formation.     

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.     

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.     

14C in Methane and DIC in the Deep Terrestrial Subsurface: Implications for Microbial Methanogenesis

A comparison between the ¹⁴C content of the methane and dissolved inorganic carbon (DIC) in deep, terrestrial subsurface systems was used to assess the timing of microbial methanogenesis contributing to gases in fracture water samples from three mines in the Witwatersrand Basin, South Africa. The results demonstrated that the majority of methane was produced over geologic timescales. In four of the samples, the methane contained no significant radiocarbon, indicating that the estimated 90% microbial methane in these samples was produced in the geologic past by indigenous microbial communities. In two samples from different mines, methane Δ¹⁴C levels indicated a primarily ancient origin for the microbial methane with the potential for more recent contributions from ongoing indigenous microbial activities constrained to between 0 and 40%, and 0 and 24%, respectively. Microbiological evidence for methanogenic archaea was observed in both of these samples. One sample had a Δ¹⁴C CH₄ that was higher than the corresponding DIC, indicating an extreme decoupling between these species and raising concerns over the representative quality of this sample. The variations in the Δ¹⁴C of DIC and CH₄ between and within mines demonstrate the need for a thorough assessment of each sample to obtain an accurate understanding of the role and timing of microbiological gas production in these complex, heterogeneous, terrestrial subsurface systems. The approach detailed here introduces timing as a new and widely applicable signature for the recognition of a major geochemical marker of indigenous life in the deep subsurface.     

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.     

Bacterial diversity and carbonate precipitation in the giant microbialites from the highly alkaline Lake Van, Turkey

Lake Van harbors the largest known microbialites on Earth. The surface of these huge carbonate pinnacles is covered by coccoid cyanobacteria whereas their central axis is occupied by a channel through which neutral, relatively Ca-enriched, groundwater flows into highly alkaline (pH ~9.7) Ca-poor lake water. Previous microscopy observations showed the presence of aragonite globules composed by rounded nanostructures of uncertain origin that resemble similar bodies found in some meteorites. Here, we have carried out fine-scale mineralogical and microbial diversity analyses from surface and internal microbialite samples. Electron transmission microscopy revealed that the nanostructures correspond to rounded aragonite nanoprecipitates. A progressive mineralization of cells by the deposition of nanoprecipitates on their surface was observed from external towards internal microbialite areas. Molecular diversity studies based on 16S rDNA amplification revealed the presence of bacterial lineages affiliated to the Alpha-, Beta- and Gammaproteobacteria, the Cyanobacteria, the Cytophaga-Flexibacter-Bacteroides (CFB) group, the Actinobacteria and the Firmicutes. Cyanobacteria and CFB members were only detected in surface layers. The most abundant and diverse lineages were the Firmicutes (low GC Gram positives). To the exclusion of cyanobacteria, the closest cultivated members to the Lake Van phylotypes were most frequently alkaliphilic and/or heterotrophic bacteria able to degrade complex organics. These heterotrophic bacteria may play a crucial role in the formation of Lake Van microbialites by locally promoting carbonate precipitation.     

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.     

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:

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.     

Underwater sinkhole sediments sequester Lake Huron’s carbon

Lake Huron’s submerged sinkhole habitats are impacted by high-conductivity groundwater that allows photosynthetic cyanobacterial mats to form over thick, carbon-rich sediments. To better understand nutrient cycling in these habitats, we measured the stable isotopic content of carbon and nitrogen in organic and inorganic carbon pools in Middle Island sinkhole, a ~23 m deep feature influenced by both groundwater and overlying lake water. Two distinct sources of dissolved CO₂ (DIC) were available to primary producers. Lake water DIC (δ ¹³C = ?0.1 ‰) differed by +5.9 ‰ from groundwater DIC (δ ¹³C = ?6.0 ‰). Organic carbon fixed by primary producers reflected the two DIC sources. Phytoplankton utilizing lake water DIC were more enriched in ¹³C (δ ¹³C = ?22.2 to ?23.2 ‰) than mat cyanobacteria utilizing groundwater DIC (δ ¹³C = ?26.3 to ?30.0 ‰). Sinkhole sediments displayed an isotopic signature (δ ¹³C = ?23.1 ‰) more similar to sedimenting phytoplankton than the cyanobacterial mat. Corroborated by sediment C/N ratios, these data suggest that the carbon deposited in sinkhole sediments originates primarily from planktonic rather than benthic sources. ²¹⁰Pb/¹³⁷Cs radiodating suggests rapid sediment accumulation and sub-bottom imaging indicated a massive deposit of organic carbon beneath the sediment surface. We conclude that submerged sinkholes may therefore act as nutrient sinks within the larger lake ecosystem.