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.     

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.     

Modern carbonate microbialites from an asbestos open pit pond, Yukon, Canada

Microbialites were discovered in an open pit pond at an abandoned asbestos mine near Clinton Creek, Yukon, Canada. These microbialites are extremely young and presumably began forming soon after the mine closed in 1978. Detailed characterization of the periphyton and microbialites using light and scanning electron microscopy was coupled with mineralogical and isotopic analyses to investigate the mechanisms by which these microbialites formed. The microbialites are columnar in form (cm scale), have an internal spherulitic fabric (mm scale), and are mostly made of aragonite, which is supersaturated in the subsaline pond water. Initial precipitation is seen as acicular aragonite crystals nucleating onto microbial biomass and detrital particles. Continued precipitation entombs benthic diatoms (e.g. Brachysira vitrea), filamentous algae (e.g. Oedogonium sp.), dinoflagellates, and cyanobacteria. The presence of phototrophs at spherulite centers strongly suggests that these microbes play an important initial role in aragonite precipitation. Substantial growth of individual spherulites occurs abiotically through periodic precipitation of aragonite that forms concentric laminations around spherulite centers while pauses in spherulite growth allow for colonization by microbes. Aragonite associated with biomass (δ(13)C = -4.6‰ VPDB) showed a (13)C-enrichment of 0.8‰ relative to aragonite exhibiting no biomass (δ(13)C = -5.4‰ VPDB), which suggests a modest removal of isotopically light dissolved inorganic carbon by phototrophs. The combination of a low sedimentation rate, high calcification rate, and low microbial growth rate appears to result in the formation of these microbialites. The formation of microbialites at an historic mine site demonstrates that an anthropogenically constructed environment can foster microbial carbonate formation.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Prokaryotic and eukaryotic community structure in field and cultured microbialites from the alkaline Lake Alchichica (Mexico)

The geomicrobiology of crater lake microbialites remains largely unknown despite their evolutionary interest due to their resemblance to some Archaean analogs in the dominance of in situ carbonate precipitation over accretion. Here, we studied the diversity of archaea, bacteria and protists in microbialites of the alkaline Lake Alchichica from both field samples collected along a depth gradient (0-14 m depth) and long-term-maintained laboratory aquaria. Using small subunit (SSU) rRNA gene libraries and fingerprinting methods, we detected a wide diversity of bacteria and protists contrasting with a minor fraction of archaea. Oxygenic photosynthesizers were dominated by cyanobacteria, green algae and diatoms. Cyanobacterial diversity varied with depth, Oscillatoriales dominating shallow and intermediate microbialites and Pleurocapsales the deepest samples. The early-branching Gloeobacterales represented significant proportions in aquaria microbialites. Anoxygenic photosynthesizers were also diverse, comprising members of Alphaproteobacteria and Chloroflexi. Although photosynthetic microorganisms dominated in biomass, heterotrophic lineages were more diverse. We detected members of up to 21 bacterial phyla or candidate divisions, including lineages possibly involved in microbialite formation, such as sulfate-reducing Deltaproteobacteria but also Firmicutes and very diverse taxa likely able to degrade complex polymeric substances, such as Planctomycetales, Bacteroidetes and Verrucomicrobia. Heterotrophic eukaryotes were dominated by Fungi (including members of the basal Rozellida or Cryptomycota), Choanoflagellida, Nucleariida, Amoebozoa, Alveolata and Stramenopiles. The diversity and relative abundance of many eukaryotic lineages suggest an unforeseen role for protists in microbialite ecology. Many lineages from lake microbialites were successfully maintained in aquaria. Interestingly, the diversity detected in aquarium microbialites was higher than in field samples, possibly due to more stable and favorable laboratory conditions. The maintenance of highly diverse natural microbialites in laboratory aquaria holds promise to study the role of different metabolisms in the formation of these structures under controlled conditions.     

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

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

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.     

Irradiance Regulation of Photosynthesis and Respiration in Modern Marine Microbialites Built by Benthic Cyanobacteria in a Tropical Lagoon (New Caledonia)

Microbialites are organosedimentary deposits that have built up as a result of the growth and binding of detrital sediment by a benthic microbial community. This study focuses on microbialites built by monospecific populations of cyanobacteria in the south-west lagoon of New Caledonia, where they have been observed down to 20–25 m depth. The aim was to study their photosynthetic and respiratory responses to various light intensities. The Phormidium sp. TK1 microbialite was collected at 19 m depth and the P. crosbyanum (Tilden) microbialite was collected at 0.5 and 13 m depth. Phormidium sp. TK1 showed all the characteristic features of a low-light adapted species. The initial slope of the Photosynthesis versus Irradiance curve for this microbialite was close to the maximum quantum yield indicating an efficient light absorption and utilization at low light. The photosynthesis maximum was located 0.2–0.4 mm below the surface and did not shift with changing light intensity. Respiration rates were low and not enhanced by light; photoinhibition was observed at higher light intensities. In Phormidium crosbyanum (Tilden) microbialites, the photosynthesis maximum shifted downward to lower depths with increasing light, probably as a result of phototactic migration of cyanobacterial filaments, and light-enhanced respiration was observed at light intensities above light saturation. The photosynthetic para- meters measured in P. crosbyanum indicate that P. crosbyanum is capable of photo-acclimation at high light intensities. The gross productivity of the different microbialites was comparable to values measured in cyanobacterial stromatolites observed in other shallow environments. However, the microbialites studied here were characterized by a lower respiration / production ratio which indicates a higher growth efficiency.     

Photosynthetic efficiency and rate of CO2 assimilation by Arthrospira (Spirulina) platensis continuously cultivated in a tubular photobioreactor

Similar to other photosynthetic microorganisms, the cyanobacterium Arthrospira platensis can be used to produce pigments, single cell proteins, fatty acids (which can be used for bioenergy), food and feed supplements, and biofixation of CO₂. Cultivation in a specifically designed tubular photobioreactor is suitable for photosynthetic biomass production, because the cultivation area can be reduced by distributing the microbial cells vertically, thus avoiding loss of ammonia and CO₂. The aim of this study was to investigate the influence of light intensity and dilution rate on the photosynthetic efficiency and CO₂ assimilation efficiency of A. platensis cultured in a tubular photobioreactor in a continuous process. Urea was used as a nitrogen source and CO₂ as carbon source and for pH control. Steady‐state conditions were achieved in most of the runs, indicating that continuous cultivation of this cyanobacterium in a tubular photobioreactor could be an interesting alternative for the large‐scale fixation of CO₂ to mitigate the greenhouse effect while producing high protein content biomass.     

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.     

Stromatolitic knobs in Storr's Lake (San Salvador,Bahamas): a model system for formation and alteration of laminae

The initial lamination in young, metabolically active Scytonema knobs developing in Storr's Lake (Bahamas) results from the iterative succession of two different stages of microbial growth at the top of this microbialite. Stage 1 is dominated by vertically oriented cyanobacterial filaments and is characterized by a high porosity of the fabric. Stage 2 shows a higher microbial density with the filaments oriented horizontally and with higher carbonate content. The more developed, dense microbial community associated with Stage 2 of the Scytonema knobs rapidly degrades extracellular organic matter (EOM) and coupled to this, precipitates carbonate. The initial nucleation forms high‐Mg calcite nanospheroids that progressively replace the EOM. No precipitation is observed within the thick sheath of the Scytonema filaments, possibly because of strong cross‐linking of calcium and EOM (forming EOM‐Ca‐EOM complexes), which renders Ca unavailable for carbonate nucleation (inhibition process). Eventually, organominerals precipitate and form an initial lamina through physicochemical and microbial processes, including high rates of photosynthetic activity that lead to ¹³C‐enriched DIC available for initial nucleation. As this lamina moves downward by the iterative production of new laminae at the top of the microbialite, increased heterotrophic activity further alters the initial mineral product at depth. Although some rare relic preservation of ‘Stage 1–Stage 2’ laminae in subfossil knobs exists, the very fine primary lamination is considerably altered and almost completely lost when the knobs develop into larger and more complex morphologies due to the increased accommodation space and related physicochemical and/or biological alteration. Despite considerable differences in microstructure, the emerging ecological model of community succession leading to laminae formation described here for the Scytonema knobs can be applied to the formation of coarse‐grained, open marine stromatolites. Therefore, both fine‐ and coarse‐grained extant stromatolites can be used as model systems to understand the formation of microbialites in the fossil record.     

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.     

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.     

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.