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1.
  • Akam, Sajjad A., et al. (författare)
  • Methane-derived authigenic carbonates – A case for a globally relevant marine carbonate factory
  • 2023
  • Ingår i: Earth-Science Reviews. - 0012-8252 .- 1872-6828. ; 243
  • Tidskriftsartikel (refereegranskat)abstract
    • Precipitation of methane-derived authigenic carbonates (MDAC) is an integral part of marine methane production and consumption, but MDAC's relative significance to the global marine carbon cycle is not well understood. Here we provide a synthesis and perspective to highlight MDAC from a global marine carbon biogeochemistry viewpoint. MDAC formation is a result and archive of carbon‑sulfur (C S) coupling in the shallow sulfatic zone and carbon‑silicon (C Si) coupling in deeper methanic sediments. MDAC constitute a carbon sequestration of 3.93 Tmol C yr−1 (range 2.34–5.8 Tmol C yr−1) in the modern ocean and are the third-largest carbon burial mechanism in marine sediments. This burial compares to 29% (11–57%) organic carbon and 10% (6–23%) skeletal carbonate carbon burial along continental margins. MDAC formation is also an important sink for benthic alkalinity and, thereby, a potential contributor to bottom water acidification. Our understanding of the impact of MDAC on global biogeochemical cycles has evolved over the past five decades from what was traditionally considered a passive carbon sequestration mechanism in a seep-oasis setting to what is now considered a dynamic carbonate factory expanding from deep sediments to bottom waters—a factory that has been operational since the Precambrian. We present a strong case for the need to improve regional scale quantification of MDAC accumulation rates and associated carbonate biogeochemical parameters, leading to their incorporation in present and paleo‑carbon budgets in the next phase of MDAC exploration.
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3.
  • Foster, William J., et al. (författare)
  • Suppressed competitive exclusion enabled the proliferation of Permian/Triassic boundary microbialites
  • 2020
  • Ingår i: Depositional Record. - : Wiley. - 2055-4877. ; 6:1, s. 62-74
  • Tidskriftsartikel (refereegranskat)abstract
    • During the earliest Triassic microbial mats flourished in the photic zones of marginal seas, generating widespread microbialites. It has been suggested that anoxic conditions in shallow marine environments, linked to the end-Permian mass extinction, limited mat-inhibiting metazoans allowing for this microbialite expansion. The presence of a diverse suite of proxies indicating oxygenated shallow sea-water conditions (metazoan fossils, biomarkers and redox proxies) from microbialite successions have, however, challenged the inference of anoxic conditions. Here, the distribution and faunal composition of Griesbachian microbialites from China, Iran, Turkey, Armenia, Slovenia and Hungary are investigated to determine the factors that allowed microbialite-forming microbial mats to flourish following the end-Permian crisis. The results presented here show that Neotethyan microbial buildups record a unique faunal association due to the presence of keratose sponges, while the Palaeotethyan buildups have a higher proportion of molluscs and the foraminifera Earlandia. The distribution of the faunal components within the microbial fabrics suggests that, except for the keratose sponges and some microconchids, most of the metazoans were transported into the microbial framework via wave currents. The presence of both microbialites and metazoan associations were limited to oxygenated settings, suggesting that a factor other than anoxia resulted in a relaxation of ecological constraints following the mass extinction event. It is inferred that the end-Permian mass extinction event decreased the diversity and abundance of metazoans to the point of significantly reducing competition, allowing photosynthesis-based microbial mats to flourish in shallow water settings and resulting in the formation of widespread microbialites.
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4.
  • Friesenbichler, Evelyn, et al. (författare)
  • Sponge-microbial build-ups from the lowermost Triassic Chanakhchi section in southern Armenia: Microfacies and stable carbon isotopes
  • 2018
  • Ingår i: Palaeogeography, Palaeoclimatology, Palaeoecology. - : Elsevier BV. - 0031-0182. ; 490, s. 653-672
  • Tidskriftsartikel (refereegranskat)abstract
    • The end-Permian mass extinction was the most severe biotic crisis in Earth's history. In its direct aftermathmicrobial communities colonized some of the space left vacant after the severe decline of skeletal metazoans.The Permian-Triassic boundary microbialites were peculiarly abundant on low-latitude shallow-marine carbonateshelves of central Tethyan continents. Armenia features particularly well preserved and diverse basalTriassic sponge-microbial build-ups (BTSMBs), which were not studied in detail to date. Here, the Chanakhchisection in southern Armenia is described petrographically and by means of stable isotope analyses. TheArmenian BTSMBs formed in a distally open marine setting on a pelagic carbonate ramp in the course of twophases of microbial growth during the Induan (Lower Triassic). The BTSMBs are represented by predominantlythrombolitic but also dendrolitic and digitate stromatolite biostromes and mounds that vary in height between5 cm to 12 m. The digitate stromatolites are associated with calcium carbonate crystal fans (CCFs). Microfaciesanalyses revealed that the BTSMBs exhibit a number of different growth forms and internal fabrics. The formationof CCFs was apparently not devoid of biological influence and took place above the sediment surface. Theabundance of sponges in the BTSMBs reveals that ecologically complex metazoan-microbial reefs have beenpresent already early after the end-Permian mass extinction. However, the formation of biostromes and moundsdid not depend on sponges or other metazoans. BTSMBs that formed during the second microbial growth phaserevealed similar δ13C-values like the surrounding sediment. In contrast, the δ13Cmicrobialite and δ13Csediment valuesfrom the BTSMBs and CCFs of the first growth phase show a difference of up to +2.3‰, suggesting a significantinfluence of photoautotrophy during microbially induced carbonate precipitation.
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5.
  • Guan, Hongxiang, et al. (författare)
  • Lipid Biomarker Patterns Reflect Nutritional Strategies of Seep-Dwelling Bivalves From the South China Sea
  • 2022
  • Ingår i: Frontiers in Marine Science. - Lausanne : Frontiers Media. - 2296-7745. ; 9
  • Tidskriftsartikel (refereegranskat)abstract
    • Invertebrates living at methane seeps such as mussels and clams gain nutrition through symbiosis with chemosynthetic, chiefly methanotrophic and thiotrophic bacteria. Lipid biomarkers, including their compound-specific carbon stable isotope compositions, extracted from the host tissues are predestined for deciphering the various sources of diets and the associations among varying environments, endosymbionts, and hosts. Here, we investigated lipid inventories of soft tissues of two bathymodiolin mussel species hosting aerobic methanotrophic bacteria (Gigantidas platifrons from Site F and Gigantidas haimaensis from Haima seeps), one bathymodiolin mussel with thiotrophic bacteria (Bathymodiolus aduloides from Haima seeps), and one vesicomyid clam (Archivesica marissinica from Haima seeps) from the South China Sea. The gills of mussels hosting methanotrophic symbionts were found to contain high amounts of lipids of aerobic methanotrophic bacteria, such as the 4,4-dimethyl lanosterol, and other 4-methyl sterols, and the type I methanotroph-specific monounsaturated fatty acids (MUFAs) C16:1ω9 and C16:1ω8. Production of methyl-sterols is favored over fatty acids at low oxygen concentrations, as demonstrated in culture experiments with Methylococcus capsulatus. Since lesser fatty acids and abundant sterols are found in G. haimaensis compared to G. platifrons, G. haimaensis apparently lived at very low oxygen levels. Extremely high levels of MUFAs C16:1ω7 and C18:1ω7 were found in gill tissue of both B. aduloides and the vesicomyid clam A. marissinica. Given the absence of ω8 fatty acids, both B. aduloides and the vesicomyid clam contain thiotrophic bacteria only. The occurrence of 13C-enriched 24-methylenecholesterol in B. aduloides indicates that the animal complemented its diet by filter-feeding (ca. 3% of the total sterol inventory) on photosynthetically derived carbon, whereas the majority of sterols are pointing to a diet relying on endosymbionts. Different types of 4-methyl sterols were observed between the thiotroph-containing mussel and methanotroph-containing mussels, suggesting different biosynthetic steps are present from lanosterol to cholesterol between animal hosts and aerobic methanotrophs. Among the four bivalve species, specific lipid biomarker patterns diagnostic for either the symbionts or the hosts yielded similar δ13C values in each species, indicating that the host obtained its nutrition either directly from the symbionts or derived at least most of its carbon in this way. The information derived from lipid biomarkers of bivalves and their corresponding symbionts in modern environments is vital to interpret data from the rock record, where most other methods to study microbial community composition are not applicable.
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6.
  • Heindel, Katrin, et al. (författare)
  • The formation of microbial-metazoan bioherms and biostromes following the latest Permian mass extinction
  • 2018
  • Ingår i: Gondwana Research. - : Elsevier BV. - 1342-937X. ; 61, s. 187-202
  • Tidskriftsartikel (refereegranskat)abstract
    • After the latest Permian mass extinction event, microbial mats filled the ecological niche previously occupied by metazoan reefs, resulting in widespread microbialites. This study focuses on the lipid biomarker (molecular fossil) and invertebrate fossil records from Neotethyan platform margin sections to understand microbial-metazoan bioherm formation. Here, we find that early Griesbachian thrombolitic and stromatolitic microbialites from Çürük Dag (Turkey) and Kuh e Surmeh (Iran) contain abundant lipid biomarkers, representing input from cyanobacteria, anoxygenic phototrophic bacteria, sulfate-reducing bacteria, and halophilic archaea. The biomarker inventory suggests that the microbialites were constructed by cyanobacteria-dominated microbial mats. Biomarkers of halophilic archaea are interpreted to reflect input from the water column, suggesting that the Neotethys experienced at least episodically hypersaline conditions. We also demonstrate that bacteria, possible keratose sponges (up to 50% of the carbonate is represented by the possible sponges), and microconchids lived synergistically to form microbial-metazoan bioherms in the immediate aftermath of the extinction along the western margin of the Neotethys. Abundant fossils of oxygen-dependent invertebrates (i.e. microconchids, bivalves, gastropods, brachiopods, and ostracods) and foraminifers were also found within these bioherms. The presence of invertebrates in conjunction with abundant molecular fossils of cyanobacteria indicates an oxygenated water column. Even though the presence of the biomarker isorenieratane in microbialites may considered as evidence for euxinic conditions in the water column, its absence in the background sediments rather points to a source organism belonging to the mat community. The new finding of bioherms built in part by metazoans suggests that reef ecosystems underwent a major turnover across the extinction event, and shortens the ‘metazoan reef gap’ to just the uppermost Changhsingian. During the Early Triassic, therefore, reefal ecosystems were able to recover in oxygenated settings since the earliest Griesbachian, albeit in an impoverished state.
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7.
  • Kiel, Steffen, et al. (författare)
  • A thyasirid-dominated methane-seep deposit from Montañita, southwestern Ecuador, from the Oligocene-Miocene boundary
  • 2021
  • Ingår i: Palaeogeography, Palaeoclimatology, Palaeoecology. - Amsterdam : Elsevier. - 0031-0182 .- 1872-616X. ; 575
  • Tidskriftsartikel (refereegranskat)abstract
    • A mass occurrence of the thyasirid bivalve Thyasira montanita in a limestone bed, exposed at Punta Montañita on the northern side of the Santa Elena peninsula in southeastern Ecuador, is here identified as an ancient methane-seep deposit. The massive to nodular limestone shows carbonate phases and microfabrics typical of seep limestones, such as banded and botryoidal cements. Its δ13 Ccarbonate values, as low as −52.6‰, suggest a biogenic methane source. Furthermore, the carbonate microfabrics and biomarkers typical of ANME-2 archaea indicate high seepage rates. Tubular fossils previously interpreted as worm tubes show cone-shaped projections and an internal structure consistent with that of crustacean cuticle and are here interpreted as crustacean fragments. Seep deposits with mass occurrences of thyasirid bivalves have so far only been reported from the Cretaceous, making the Montañita seep deposit the first such example from the Cenozoic. The faunal and biomarker inventories of this deposit indicate deposition on an open shelf; the oysters on the upper surface of the limestone are likely secondary encrusters that arrived only after the emergence of this deposit in shallower water.
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8.
  • Kiel, S., et al. (författare)
  • Brachiopods in early Mesozoic cryptic habitats: Continuous colonization, rapid adaptation, and wide geographic distribution
  • 2021
  • Ingår i: Palaeogeography, Palaeoclimatology, Palaeoecology. - Amsterdam : Elsevier. - 0031-0182 .- 1872-616X. ; 583, s. 110668-110668
  • Tidskriftsartikel (refereegranskat)abstract
    • Late Triassic and early Jurassic dikes and fissures in the Dachstein Limestone in the Northern Calcareous Alps harbor mass occurrences of the rhynchonellide brachiopods Sulcirostra juvavica and Halorella amphitoma. To test recent hypotheses about their paleoecology, we characterized these habitats using petrography, carbon stable isotopes, and trace element patterns, and found no evidence for hydrocarbon seepage or hydrothermal venting. Thus the brachiopods lived under normal-marine conditions, in darkness and absence of local photosynthetic primary production, hence relying on the supply of limited and presumably small-sized food washed into the dikes and fissures. Because Halorella and Sulcirostra occur in dikes and fissures since the beginning of their stratigraphic ranges, these rhynchonellides are not relics of formerly widely distributed taxa, but instead are two genera that rapidly adapted to these habitats. Both Halorella and Sulcirostra occur also in late Triassic and early Jurassic deep-water settings such as deep-marine sills and hydrocarbon seeps, indicating that close phylogenetic relationships between submarine cave faunas and deep-sea faunas, as seen today, existed also in the early Mesozoic, albeit among very different taxa. Another analogy to the modern cave fauna is the wide but disjunct geographic distribution of Sulcirostra and Halorella, both found throughout the Tethys and Panthalassa oceans. Our findings support the view that submarine cave habitats were continuously colonized by new taxa throughout Earth’s history.
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9.
  • Kiel, Steffen, et al. (författare)
  • Fossiliferous methane-seep deposits from the Cenozoic Talara Basin in northern Peru
  • 2020
  • Ingår i: Lethaia. - : Scandinavian University Press / Universitetsforlaget AS. - 0024-1164 .- 1502-3931. ; 53:2, s. 166-182
  • Tidskriftsartikel (refereegranskat)abstract
    • Thirteen fossiliferous limestone deposits from Cenozoic strata in the Talara Basin in northern Peru are identified as ancient methane-seep deposits. Planktonic foraminifera and the existing stratigraphical framework of the Talara Basin indicate an early Oligocene, or possibly late Eocene, age of these deposits. They are found in three distinct areas - Belen, Cerro La Salina and Cerros El Pelado - and differ in their petrography, stable isotope signatures, and lipid biomarker and macrofaunal contents. At Belen, the carbon stable isotope signature of the carbonate and the abundance of n-alkanes indicates the possibility of oil seepage in addition to methane seepage; for Belen and Cerro La Salina the high abundance of the biomarker crocetane indicates a dominance of anaerobic methane-oxidizing archaea of the ANME-2 group, whereas the rather small combined crocetane/phytane peak of a Cerros El Pelado limestone agrees with mixed ANME-1/ANME-2 input. The macrofauna consists mainly of molluscs; the Cerro La Salina sites include mostly infaunal thyasirid and lucinid bivalves and only few vesicomyid bivalves; gastropods include Provanna antiqua, the limpet Pyropelta and several vetigastropods. The Belen site is dominated by the elongate vesicomyid bivalve Pleurophopsis lithophagoides. The most common bivalve at the Cerros El Pelado sites is an undetermined, possible vesicomyid, and a smooth provannid gastropod. Biogeographically the faunas are most similar to those of the northwestern United States, as indicated by two joint species; similarities on the genus level (Conchocele, Lucinoma, Pleurophopsis, Provanna, Colus) exist also with Japan and the Caribbean region.
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10.
  • Kiel, Steffen, et al. (författare)
  • Late Triassic mollusk-dominated hydrocarbon-seep deposits from Turkey
  • 2017
  • Ingår i: Geology. - Boulder : Geological Society of America. - 0091-7613 .- 1943-2682. ; 44, s. 751-754
  • Tidskriftsartikel (refereegranskat)abstract
    • Deep-sea hydrothermal vents and hydrocarbon seeps host unique ecosystems relying on geochemical energy rather than photosynthesis. Whereas the fossil and evolutionary history of these ecosystems is increasingly well known from the Cretaceous onward, their earlier history remains poorly understood and brachiopods are considered to have played a dominant role during the Paleozoic and Mesozoic. Here we report five new hydrocarbon-seep deposits from the Upper Triassic Kasımlar shales in southern Turkey. The pyritiferous seep limestones predominantly consist of 13C-depleted micrite with δ13C values as low as −10.4‰, and contain only sparse 13C-depleted rim cement (δ13C as low as −12.0‰), interpreted to result from the recrystallization of banded and botryoidal crystal aggregates of fibrous cement. The geologic ages of the studied seep deposits were determined as late Carnian and early Norian using conodonts. The associated fauna is dominated by modiomorphid and anomalodesmatan bivalves, and also includes a diversity of gastropods and the dimerelloid brachiopod Halorella. These faunal assemblages allow a comparison between seep faunas from the two major Triassic ocean basins—the present assemblages being from Tethys, and the only previously known examples being from eastern Panthalassa—and indicate that a cosmopolitan, seep-restricted fauna as in the present-day oceans has existed since the Late Triassic. With almost 20 species, the seep fauna of the Kasımlar shales approaches the diversity of Cretaceous to present-day seep faunas, further emphasizing the ecological similarity of seep faunas since the early Mesozoic. Our findings also highlight that brachiopods and bivalves had a more complex history of coexistence at seeps than currently appreciated.
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  • Müller, Sandro P, et al. (författare)
  • Mass occurrence of echinoids in an Oligocene hydrocarbon-seep limestone from the Olympic Peninsula, Washington State, USA
  • 2023
  • Ingår i: Geological Magazine. - Cambridge : Cambridge University Press. - 0016-7568 .- 1469-5081. ; 160:5, s. 941-954
  • Tidskriftsartikel (refereegranskat)abstract
    • Loose limestone blocks of a newly recognized hydrocarbon-seep deposit from the lower Oligocene Jansen Creek Member of the Makah Formation were collected on a beach terrace close to the mouth of Bullman Creek in Washington State, USA. The limestone consists largely of authigenic carbonate phases, including 13C-depleted fibrous cement forming banded and botryoidal crystal aggregates with δ13C values as low as –23.5 ‰. Lipids extracted from the limestone yielded molecular fossils of anaerobic methanotrophic archaea (ANME), dominated by compounds of an ANME-2/DSS consortium with δ13C values as low as −106 ‰, indicating formation at an ancient methane seep. The fossil inventory of the seep deposit is remarkable, consisting almost solely of echinoid remains, whereas typical seep biota are absent. Varying preservation of the echinoid fossils indicates parautochthonous deposition, corroborated by evidence for high fluid flow at the ancient seep, possibly responsible for displacement of echinoids after death. Although a full taxonomic description of the echinoids cannot be given, almost all fossils were assigned to one taxon of irregular spatangoids, except for a single regular echinoid. Abundance and lifestyle of the irregular spatangoids in the Bullman Creek echinoid seep deposit resemble those of the fossil Tithonia oxfordiana from an upper Jurassic seep deposit in France and extant Sarsiaster griegii from modern seeps in the Gulf of Mexico. The Bullman Creek echinoid deposit probably represents a fossil analogue of the Gulf of Mexico Sarsiaster mass occurrence, indicating that the adaptation of spatangoid echinoids to chemosynthesis-based ecosystems ranges back at least to the earliest Oligocene.
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13.
  • Qu, Yuangao, et al. (författare)
  • Traces of Ancient Life in Oceanic Basalt Preserved as Iron-Mineralized Ultrastructures: Implications for Detecting Extraterrestrial Biosignatures
  • 2023
  • Ingår i: Astrobiology. - : Mary Ann Liebert. - 1531-1074 .- 1557-8070. ; 23:7, s. 769-785
  • Tidskriftsartikel (refereegranskat)abstract
    • Benefiting from their adaptability to extreme environments, subsurface microorganisms have been discovered in sedimentary and igneous rock environments on Earth and have been advocated as candidates in the search for extraterrestrial life. In this article, we study iron-mineralized microstructures in calcite-filled veins within basaltic pillows of the late Ladinian Fernazza group (Middle Triassic, 239 Ma) in Italy. These microstructures represent diverse morphologies, including filaments, globules, nodules, and micro-digitate stromatolites, which are similar to extant iron-oxidizing bacterial communities. In situ analyses including Raman spectroscopy have been used to investigate the morphological, elemental, mineralogical, and bond-vibrational modes of the microstructures. According to the Raman spectral parameters, iron minerals preserve heterogeneous ultrastructures and crystallinities, coinciding with the morphologies and precursor microbial activities. The degree of crystallinity usually represents a microscale gradient decreasing toward previously existing microbial cells, revealing a decline of mineralization due to microbial activities. This study provides an analog of possible rock-dwelling subsurface life on Mars or icy moons and advocates Raman spectroscopy as an efficient tool for in situ analyses. We put forward the concept that ultrastructural characteristics of minerals described by Raman spectral parameters corresponding to microscale morphologies could be employed as carbon-lean biosignatures in future space missions. Key Words: Ultrastructures—Iron minerals—Oceanic basalt—Subsurface biosignatures.
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14.
  • Rubin-Blum, Maxim, et al. (författare)
  • Fueled by methane : deep-sea sponges from asphalt seeps gain their nutrition from methane-oxidizing symbionts
  • 2019
  • Ingår i: The ISME Journal. - : NATURE PUBLISHING GROUP. - 1751-7362 .- 1751-7370. ; 13:5, s. 1209-1225
  • Tidskriftsartikel (refereegranskat)abstract
    • Sponges host a remarkable diversity of microbial symbionts, however, the benefit their microbes provide is rarely understood. Here, we describe two new sponge species from deep-sea asphalt seeps and show that they live in a nutritional symbiosis with methane-oxidizing (MOX) bacteria. Metagenomics and imaging analyses revealed unusually high amounts of MOX symbionts in hosts from a group previously assumed to have low microbial abundances. These symbionts belonged to the Marine Methylotrophic Group 2 Glade. They are host-specific and likely vertically transmitted, based on their presence in sponge embryos and streamlined genomes, which lacked genes typical of related free-living MOX. Moreover, genes known to play a role in host-symbiont interactions, such as those that encode eukaryote-like proteins, were abundant and expressed. Methane assimilation by the symbionts was one of the most highly expressed metabolic pathways in the sponges. Molecular and stable carbon isotope patterns of lipids confirmed that methane-derived carbon was incorporated into the hosts. Our results revealed that two species of sponges, although distantly related, independently established highly specific, nutritional symbioses with two closely related methanotrophs. This convergence in symbiont acquisition underscores the strong selective advantage for these sponges in harboring MOX bacteria in the food-limited deep sea.
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  • Zwicker, Jennifer, et al. (författare)
  • Mineral authigenesis within chemosynthetic microbial mats: Coated grain formation and phosphogenesis at a Cretaceous hydrocarbon seep, New Zealand
  • 2021
  • Ingår i: The Depositional Record. - : John Wiley & Sons. - 2055-4877. ; 7:2, s. 294-310
  • Tidskriftsartikel (refereegranskat)abstract
    •  Marine hydrocarbon seeps are sites of chemosynthetic microbial activity and authigenic carbonate formation. Seep limestones are typified by a range of geochemical signatures of microbial hydrocarbon oxidation, but only few seep deposits reveal mesofabrics that can be regarded as evidence of microbial activity. A Cretaceous methane-seep limestone from Waipiro Bay, New Zealand, exhibits a fabric composed of cryptocrystalline carbonate fluorapatite between carbonate coated grains and spheroidal calcite. To understand the unusual Waipiro deposit, a paragenetic sequence has been derived for coated grains, spheroidal calcite and carbonate fluorapatite using thin section petrography, scanning electron microscopy, Fourier-transform infrared spectroscopy and stable isotope geochemistry. The formation of 13C-depleted coated grains (δ13C values as low as -15.8‰ Vienna-Pee Dee Belemnite) and spheroidal calcite (δ13C values as low as -21.3‰) was favoured by hydrocarbon oxidation. Fibrous banded and botryoidal cement, a typical early diagenetic phase of hydrocarbon-seep deposits, features δ13C values as low as -22.9‰. Coated grains and spheroidal calcite grew by displacive growth in a gel-like medium, probably a microbial mat. Phosphorus is a mobile element and marine pore waters are typically undersaturated with respect to carbonate fluorapatite. Specific conditions are consequently required to retain sufficient concentrations to precipitate carbonate fluorapatite. Possible sources of phosphorus for the formation of the 13C-depleted Waipiro carbonate fluorapatite (mean δ13C value of -15.4‰) include (a) the oxidation of sedimentary organic matter by organoclastic sulphate reduction, (b) the degradation of the microbial mat itself and (c) the active release of polyphosphate by sulphide-oxidizing bacteria. This study suggests that the formation of the Cretaceous Waipiro seep limestone involved an interplay of biogeochemical processes including sulphate-driven anaerobic oxidation of methane, organoclastic sulphate reduction and possibly nitrate-dependent sulphide oxidation. It further demonstrates that  coated grains resembling oncoids are not restricted to shallow water, photosynthesis dependent ecosystems. 
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