A coral-microbialite patch reef from the late jurassic (florigemma-Bank, Oxfordian) of NW Germany (Süntel mountains)

Summary Anin situ Oxfordian patch reef from the Süntel hills (florigemma-Bank, Korallenoolith, NW-Germany) is described. It is composed of an autochthonous reef core overlain by a ‘parautochthonous’ biostrome. The exposed reefal area amounts to about 20 m in lateral and up to 4 m in vertical direction. Nearly all major marine reefal fossil associations from the Tethyal realm are present. In the reef core two facies can be distinguished: (1)Thamnasteria dendroidea thicket facies and (2) thrombolite facies. The first facies is composed of a thin branched autochthonous coral thicket mainly constructed ofTh. dendroidea colonies with only a minor portion ofStylosmilia. Frequently, theTh. dendroidea branches laterally coalesce bridge-like forming a delicate initial framework which was subsequently reinforced by thick microbial coatings, that make up approximately 80% of the rock volume. This facies is an excellent example for microbialite binding in reefal architecture. Additionally, several generations of micromorphic and partly cryptic encrusting organisms settled on theTh. dendroidea branches and microbialite crusts. They successively overgrow each other and fill the space between the coral branches in the thicket forming a characteristic community replacement sequence. Initial colonization of theThamnasteria dendroidea took place on an oncoidic/bioclastic hardground. During this early phase of reefal development, microbialites also played an important role in stabilizing and binding the reef body. The thrombolite facies (2) occupying nearly the same volume of the reef body as facies type (1) consists of a thrombolitic microbialitic limestone which fills the interstice between the coral colonies. It shows a considerably lower faunal diversity than theTh. dendroidea facies. Numerous cavities are interspersed in the thrombolithe and are almost completely filled with dolomitized allomicrite. In contrast, microbialite and allomicrite adjacent to the reef core rarely reveal any dolomitized areas. Above the reef core, mostly toppledSolenopora jurassica thalli occur together with a few massiveIsastrea colonies forming a parautochthnous biostrome. They are inhabited by a low diverse assemblage of encrusting organisms. Microbialites are only rarely present in this biostromal unit. The patch reef is developed within a lagoonal limemud facies both separated by a sharp interface. In contrast, continuous facies transition exists between theSolenopora biostrome and adjacent deposits which are characterized by micritic to pelmicritic limestone sometimes with lenses of oncoids. Debris derived from the patch reef is only sporadically intercalated in the reef surrounding lagoonal sediments. Gastropods, bivalves, and dasycladalean algae dominate the lagoonal biota. Up-section following theSolenopora biostrome nerinean gastropods become the most abundant species amounting to a ‘Nerinea-bed’. This horizon moderately elevates above the patch reef indicating, that is arose above the surrounding sea floor forming a relief. The patch reef established on a secondary hardground probably released by a minor transgression and a nondepositional regime. It grew up on a well-illuminated sea floor only a few meters below sea level. Only a low background sedimentation rate and modest water circulation are assumed during reefal growth. These features characterize an open marine lagoon. A subsequent shallowing upwards trend caused emergence of the early lithifiedflorigemma-Bank sediments. In the following erosional phase the reef core,Solenopora biostrome and ‘Nerinea-bed’ were sharply cut. Paleokarst phenomena (karst solution of the rocks, selective leaching of the aragonitic corals) truncate the surface of theflorigemma-Bank. Released by a transgressive sea level, the paleokarst surface is densely inhabited by marine boring and encrusting organisms (oysters, serpulids). Karst cavities are filled with an oncoid-bearing bioclastic limestone with a large portion of siliciclastics. Theflorigemma-Bank is overlain by the reddish bioclastic sandstone of the ‘Zwischenfl?zregion’.     

‘Stromatolites’ built by sponges and microbes – a new type of Phanerozoic bioconstruction

Two ‘stromatolites’ from Carboniferous and Triassic carbonates previously regarded as microbial bioconstructions are analysed and reinterpreted as sponge‐microbial build‐ups. The automicritic aggregations in these build‐ups are similar to the previously reported fossils of keratose demosponges in showing moulded anastomosing filamentous structures. All the studied columnar or domal constructions were formed in turbulent water with high sedimentation rate. The Carboniferous build‐ups were constructed in the shallow subtidal zone of an open shelf or a ramp. The laminations within the stromatolite‐like columns are composed of alternating dark micritic laminae of sponge fossils and pale laminae of neomorphic microspars. The accretion of these columns is probably related to the repeated cycles of sponge growth, rapid lithification after burial, re‐exposure and erosion, and settlement of new generations. The Triassic rocks are presumed to have been precipitated in a slightly evaporitic environment based on lithological features. They show a transition from planar laminae, which were formed under the influence of microbial mats, to stromatolitic columnar or domal build‐ups, which are dominated by stacked micritic clumps of probable sponge fossils. The sponge–microbe alternation may have been controlled by variation of salinity. Comparable with a recent study, this work shows that sponge‐related bioconstructions can be morphologically similar to microbialites in the level of mega‐ and mesostructures.     

Rare earth and yttrium elements (REY) patterns of mesostructures of Miaolingian (Cambrian) thrombolites at Jiulongshan,Shandong Province,China

The relationship between the thrombolitic mesostructures and their depositional environments is still poorly understood due to inconsistent results by sedimentary investigation. Rare earth elements plus yttrium (REY) in ancient microbialites have been extensively applied to paleoenvironmental studies owing to their fractionation in different depositional environments. In order to investigate the environmental controls on thrombolitic mesostructures, we present the REY concentrations and patterns of four types of mesostructures of the Miaolingian (Cambrian) thrombolites in the Changhia Formation at the Jiulongshan section, Shandong Province, China. The REY compositions of those thrombolites show two distinctive groups: (1) light REY depleted patterns with negative Ce anomalies in spotted (SM) and layered mesostructures (LM) of thrombolites; and (2) flat patterns with weak Ce anomalies in dendritic (DM) and meshed mesostructures (MM) of thrombolites. Controlling factors analysis reveals that terrigenous detritus inputs have stronger influence on REY in SM and LM. In contrast, early diagenetic porewaters from underlying sediments have more serious impacts on REY concentrations and patterns in DM and MM. Our results clearly indicate that SM and LM were formed under oxic marine settings with minor terrigenous inputs, whereas DM and MM formed under suboxic marine settings suffered from early diagenetic porewater from underlying sediments. This new geochemical evidence suggests that thrombolitic mesostructures were strongly influenced by paleoenvironment, and REY of thrombolites with controlling factors analysis can be utilized as effective proxies for paleoenvironments.     

Potential role for microbial ureolysis in the rapid formation of carbonate tufa mounds

Modern carbonate tufa towers in the alkaline (~pH 9.5) Big Soda Lake (BSL), Nevada, exhibit rapid precipitation rates (exceeding 3 cm/year) and host diverse microbial communities. Geochemical indicators reveal that carbonate precipitation is, in part, promoted by the mixing of calcium-rich groundwater and carbonate-rich lake water, such that a microbial role for carbonate precipitation is unknown. Here, we characterize the BSL microbial communities and evaluate their potential effects on carbonate precipitation that may influence fast carbonate precipitation rates of the active tufa mounds of BSL. Small subunit rRNA gene surveys indicate a diverse microbial community living endolithically, in interior voids, and on tufa surfaces. Metagenomic DNA sequencing shows that genes associated with metabolisms that are capable of increasing carbonate saturation (e.g., photosynthesis, ureolysis, and bicarbonate transport) are abundant. Enzyme activity assays revealed that urease and carbonic anhydrase, two microbial enzymes that promote carbonate precipitation, are active in situ in BSL tufa biofilms, and urease also increased calcium carbonate precipitation rates in laboratory incubation analyses. We propose that, although BSL tufas form partially as a result of water mixing, tufa-inhabiting microbiota promote rapid carbonate authigenesis via ureolysis, and potentially via bicarbonate dehydration and CO₂ outgassing by carbonic anhydrase. Microbially induced calcium carbonate precipitation in BSL tufas may generate signatures preserved in the carbonate microfabric, such as stromatolitic layers, which could serve as models for developing potential biosignatures on Earth and elsewhere.     

四川江油渔洞子剖面下三叠统飞仙关组底部龙介类的古生态意义

三叠系龙介类化石在欧洲早有报道。在我国四川江油渔洞子剖面下三叠统飞仙关组底部牙形刺Hindeodus pavus带微生物岩中首次发现环节动物龙介类化石Spirorbisphlyctaena,与以蓝菌为主的自养型底栖微生物群落共生,是早三叠世生物复苏的先驱者。     

Photosynthesis,Respiration and Exopolymer Calcium-Binding in Biofilm Calcification (Westerhöfer and Deinschwanger Creek,Germany)

The impact of microbial activity on biofilm calcification in aquatic environments is still a matter of debate, especially in settings where ambient water has high CaCO₃ mineral supersaturation. In this study, biofilms of two CO₂-degassing karst-water creeks in Germany, which attain high calcite supersaturation during their course downstream, were investigated with regard to water chemistry of the biofilm microenvironment. The biofilms mainly consisted of filamentous cyanobacteria (Phormidium morphotype) and heterotrophic bacteria (including sulfate-reducing bacteria), which affect the microenvironment and produce acidic exopolymers. In situ and ex situ microelectrode measurements showed that a strong pH increase, coupled with Ca^{2 +} consumption, occurred in light conditions at the biofilm surface, while the opposite occurred in the dark. Calcite supersaturation at the biofilm surface, calculated from ex situ Ca^{2 +} and CO₃ ^2? microelectrode measurements, showed that photosynthesis resulted in high omega values during illumination, while respiration slightly lowered supersaturation values in the dark, compared to values in the water column. Dissociation calculation demonstrated that the potential amount of Ca^{2 +} binding by exopolymers would be insufficient to explain the Ca^{2 +} loss observed, although Ca^{2 +} complexation to exopolymers might be crucial for calcite nucleation. No spontaneous precipitation occurred on biofilm-free limestone substrates under the same condition, regardless of high supersaturation. These facts indicate that photosynthesis is a crucial mechanism to overcome the kinetic barrier for CaCO₃ precipitation, even in highly supersaturated settings.     

Cambrian Series 3 lithistid sponge–microbial reefs in Shandong Province,North China: reef development after the disappearance of archaeocyaths

The Cambrian Series 3 Zhangxia Formation in Shandong Province, North China, includes small‐scale lithistid sponge–microbial reefs. The lithistid sponges grew on oolitic and bioclastic sediments, which were stabilized by microbial activities. The relative abundances of microbial components (e.g. calcimicrobe Epiphyton and stromatolites) vary among the reefs. However, the microbial components commonly encrusted or bound the lithistid sponges, formed remarkable encrustations on the surfaces of the sponges. Epiphyton especially grew upward and downward. The lithistid sponges thus provided substrates for the attachment and development of microbes, and the microbes played essential roles as consolidators, by encrusting reef‐building sponges. Additionally, the lithistid sponges were prone to degradation via microbial activities and diagenetic processes, and were thus preserved as micritic bodies, showing faint spicular networks or abundant spicules. Such low preservation potential within the reef environment obscured the presence of the sponges and their widespread contribution as reef‐building organisms during the Cambrian. During the prolonged interval after the demise of archaeocyaths, purely microbial reefs, such as stromatolites and thrombolites have been considered to be the principal reef builders, in association with rare lithistid sponge–microbial associations. However, recent findings, including those from Shandong Province and Korea, suggest that the lithistid sponge‐bearing reefs were more extensive during the Epoch 3 to the Furongian than previously thought. These lithistid sponge–microbial reefs were precursors of the sponge–microbial reefs that dominated worldwide in the Early Ordovician.     

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

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

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.     

微生物成因的碳酸盐矿物研究进展

微生物诱导碳酸盐类矿物沉淀是地质微生物学的研究热点之一.微生物主要通过其代谢活动促进细胞周围微环境pH值及水体[CO32-]的升高,最终表现为碳酸盐类矿物饱和指数的增加.此外,微生物及其分泌的胞外聚合物可作为碳酸盐晶核的成核位点,为碳酸盐矿物晶体的生长进一步提供有利条件.微生物成因与纯化学成因的碳酸盐类矿物相比具有不同的矿物特征(如形貌、微量元素含量及碳同位素等).深入了解微生物诱导碳酸钙沉淀的行为对理解地质时期微生物活动及其在二氧化碳的地质封存中的潜在应用具有指导意义.本文综述了微生物诱导碳酸盐矿物沉淀的机理、代谢过程,总结了该领域的最新进展,探讨了生物成因以及化学成因碳酸盐矿物的区别,最后指出了该项研究在微生物岩以及CO2地质封存上的一些可能的拓展方向.