Abstract
Lipid biomarkers of seep carbonates and sediments retrieved from the Dongsha area, Shenhu, Site F and Haima in the South China Sea (SCS) over the last two decades were studied. Biomarker inventories, microbial consortia, seepage dynamics, and biogeochemical processes of anaerobic oxidation of methane (AOM), aerobic oxidation of methane (AeOM), and oxidation of non-methane hydrocarbons, were reconstructed. Authigenic carbonates contained varying contents of 13C-depleted archaeal and bacterial biomarkers, reflecting their formation as a result of AOM under varying conditions. Except for the typical isoprenoids found in various cold seeps worldwide, 3,7,11,15-tetramethyl hexadecan-1,3-diol and two novel sn2-/sn3-O-hydroxyphytanyl glycerol monoethers with notable 13C-depletion were observed in authigenic carbonates obtained from Haima, which are most likely hydrolysis products of archaea-specific diethers. Furthermore, molecular fossils, compound-specific δ13C values, and mineralogies, were used to trace dominant microbial consortia, seepage activities, and environmental conditions in the cold seep ecosystems of the SCS. In this chapter, the archaeal and bacterial lipid biomarker geochemistry of methane seeps is systematically introduced. AOM, AeOM, oxidation of non-methane hydrocarbons, oil degradation, and the diagenetic fate of glycerol ethers, are further summarized.
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11.1 Introduction
Marine sediments are the largest methane reservoir on Earth, with greenhouse methane in the form of gas hydrates, free gas, and dissolved methane in pore-water (Wallmann et al. 2012; Joye 2020). At active seep sites, it is common that gas and fluids are transported from deep reservoirs into shallow surface sediments (Wu et al. 2011, 2022; Jin et al. 2022; Wang et al. 2022). However, most of the rising methane is consumed by methanotrophic microbial communities before it can reach the hydrosphere (Reeburgh 2007; Boetius and Wenzhöfer 2013). Methane is primarily consumed via anaerobic oxidation of methane (AOM) and aerobic oxidation of methane (AeOM) (Boetius and Wenzhöfer 2013). Sulfate-driven AOM in cold seep ecosystems is mainly performed by consortia of anaerobic methane-oxidizing archaea (ANME) and sulfate-reducing bacteria (SRB; Boetius et al. 2000), whereas AeOM is performed by aerobic microbes and animal–microbe symbioses (Boetius and Wenzhöfer 2013).
To date, three types of microbial consortia have been demonstrated to mediate sulfate-driven AOM: ANME-1/Desulfosarcina/Desulfococcus (DSS), ANME-2/DSS, and ANME-3/Desulfobulbus spp. (DBB) (Hinrichs et al. 2000; Orphan et al. 2001; Knittel et al. 2003, 2005; Losekann et al. 2007). These archaeal assemblages are closely or loosely associated with their sulfate-reducing partners, varying with the methane and sulfate concentrations, temperature, and oxygen availability (Elvert et al. 2005; Knittel et al. 2005; Nauhaus et al. 2005). Lipid biomarkers and their compound-specific carbon stable isotope compositions are useful tools to monitor the distribution and functioning of microorganisms in seep environments (Birgel et al. 2006; Niemann and Elvert 2008; Stadnitskaia et al. 2008a, b; Blumenberg et al. 2015). This is because different ANME/SRB consortia biosynthesize different lipid biomarkers and produce distinct carbon differences from methane to biomarkers (Niemann and Elvert 2008). For example, ANME-2/DSS consortia are commonly characterized by large amounts of 2,6,11,15-tetramethylhexadecane (crocetane), high sn2-hydroxyarchaeol/archaeol ratios (>1.1), and low ai-C15:0/i-C15:0-fatty acid ratios, whereas high glycerol dibiphytanyl glycerol tetraethers (GDGTs) contents, low sn2-hydroxyarchaeol/archaeol ratios (<0.8), small amounts or absence of crocetane, and high ai-C15:0/i-C15:0-fatty acid ratios, are indicators of ANME-1/DSS (Niemann and Elvert 2008). Except for specific biomarkers, the maxima in rates of AOM and sulfate reduction are commonly correlated with the highest biomarker abundance and the lowest carbon isotopic signatures (Elvert et al. 2005).
The investigation of cold seeps in the SCS started in 2004 (Chen et al. 2005; Suess 2005). Since then, authigenic carbonates, sediments at active seep sites, and seep-dwelling bivalves have been collected from more than 40 seep sites, covering a wide range of water depths on both the northern and southern continental slopes of the SCS (Chen et al. 2005; Han et al. 2008; Liang et al. 2017). Lipid biomarkers of seep carbonates retrieved from the SCS were first reported in 2008 (Birgel et al. 2008; Yu et al. 2008). Based on the molecular fossils of seep carbonates, fluid sources, microbial consortia, and seepage dynamics were constrained (Guan et al. 2013, 2016a, b, 2018; Ge et al. 2015). At the Haima cold seeps, 13C-depleted isoprenoidal 3,7,11,15-tetramethylhexadecan-1,3-diol and two novel isoprenoids, namely, sn2-/sn3-O-hydroxyphytanyl glycerol monoethers, were found as hydrolysis products of isoprenoid diethers in authigenic carbonates (Xu et al. 2022a, b). Furthermore, lipid biomarker patterns combined with the compound-specific carbon stable isotopes of active seep sediments were used to uncover AOM, AeOM and oil degradation processes (Guan et al. 2022). In the seep ecosystems of the SCS, lipid biomarkers and compound-specific carbon stable isotope compositions have been studied in the Site F, northeast Dongsha, southwest Dongsha, Shenhu, and Haima cold seep ecosystems, revealing AOM, sulfate reduction, and oil biodegradation processes (Birgel and Peckmann 2008; Yu et al. 2008; Ge et al. 2010, 2011, 2015; Guan et al. 2013, 2014, 2016a, b, 2018, 2022; Lai et al. 2021; Li et al. 2021). In this chapter, we reviewed current lipid biomarker studies of seep ecosystems in the SCS. Fluid sources, consortia assemblages, and seepage dynamics during carbonate precipitation are summarized in detail.
11.2 Lipid Biomarker Inventories
11.2.1 Archaeal Biomarkers
The biomarker inventories of seep carbonates retrieved from NE Dongsha, Shenhu, Site F, and Haima revealed diagnostic lipids of specific groups of ANME/SRB (Fig. 11.1; Birgel et al. 2008; Yu et al. 2008; Ge et al. 2010, 2015; Guan et al. 2013, 2016a, b, 2018). The biomarker patterns and stable carbon isotope compositions were found to vary among individual study sites and geographic areas. Among these data, 13C-depleted isoprenoid hydrocarbons, crocetane, 2,6,10,15,19-pentamethyleicosane (PMI), and squalane, are common (Fig. 11.2; Birgel et al. 2008; Yu et al. 2008; Ge et al. 2011, 2015; Guan et al. 2013, 2016a, b, 2018). Crocetane is particularly abundant at ANME-2-dominated methane seeps, whereas it is only present in minor amounts or absent in ANME-1-dominated environments (Guan et al. 2013, 2016a, b; Ge et al. 2015). 13C-depleted PMI-acid specific to methanotrophic archaea was found in an authigenic carbonate from NE Dongsha (Guan et al. 2013). Monocyclic diphytanyl glycerol diethers (monocyclic MDGD), possibly derived from methanotrophic archaea, were observed in carbonate pipes in the Shenhu area (Ge et al. 2011). Other isoprenoids, including 13C-depleted phytanyl-glycerolmonoethers sn2-/sn3-O-phytanyl glycerol ethers, phytanol, and phytanoic acid, are common in authigenic carbonates and sediments (Guan et al. 2013, 2016a, b, 2018, 2022). The GDGT patterns are characterized by the dominance of GDGTs with 0–2 cyclopentane moieties (Guan et al. 2016a).
Biphytanic diacids, including acyclic biphytanic diacid, monocyclic biphytanic diacid, and bicyclic biphytanic diacid, were found in seep carbonates from NE Dongsha (the Jiulong methane reef; Birgel et al. 2008). Except for the most widespread isoprenoids above, isoprenoidal 3,7,11,15-tetramethylhexadecan-1,3-diol and two novel sn2-/sn3-O-hydroxyphytanyl glycerol monoethers hydroxylated at C-3 of the phytanyl moieties were observed in seep carbonates from the Haima seep (Xu et al. 2022a, b). Most archaeal-specific isoprenoids are strongly depleted in 13C with δ13C values ranging from −140‰ to −84‰ (Fig. 11.3; Birgel et al. 2008; Yu et al. 2008; Ge et al. 2011, 2015; Guan et al. 2013, 2016a, b, 2018, 2022; Li et al. 2021).
11.2.2 Sulfate-Reducing Bacterial Biomarkers
Seep carbonates and seep sediments from the SCS are known to contain various bacterial lipids (Fig. 11.4). The otherwise uncommon fatty acids C16:1ω5 and cycC17:0ω5,6 were found in high abundances and were assigned to the sulfate-reducing partners of ANME-2 (Elvert et al. 2003; Blumenberg et al. 2004, 2005). The presence of ai-/i-C15:0-fatty acids with ratios lower than 2 is an indicator of ANME-2/DSS-dominated seeps (Niemann and Elvert 2008; Guan et al. 2013). However, the ai-/i-C15:0-fatty acid ratio does not always discriminate the sulfate-reducing partners of ANME since their contents and δ13C values can be modified by SRBs involved in oil degradation (Guan et al. 2018, 2022; Krake et al. 2022). Non-isoprenoidal monoglycerol ethers (MAGEs), ascribed to DSS associated with ANME-2 and DBB associated with ANME-3 (Niemann and Elvert 2008), were only abundant in Haima seep carbonates (Xu et al. 2022a, b). SRB-specific fatty acids were less 13C-depleted than those originating from archaea because they can assimilate dissolved inorganic carbon (DIC) as a carbon source (Wegener et al. 2008). The SRB-specific biomarkers yielded δ13C values from −11133‰ to −111‰ in seep carbonates of the SCS (Fig. 11.3; Guan et al. 2013, 2016a, b, 2018, 2022). In addition, various DAGEs with extremely low δ13C values were found with high contents in authigenic carbonates from Site F, NE Dongsha, and Haima (Fig. 11.5; Guan et al. 2013, 2016a; Xu et al. 2022a, b).
11.2.3 Aerobic Methanotrophic Biomarkers
Another striking feature of the seep carbonates originating from the SCS is the significant amounts of hopanoids and steroids with moderate to significant 13C-depletions (Fig. 11.5; Guan et al. 2016a, b, 2018, 2022). These hopanoids generally include tetrahymanol, diploptene, diplopterol, hopanoic acids (e.g., 17α(H)-21β(H)-bishomo-hopanoic acid and 17β(H)-21β(H)-bishomo-hopanoic acid), and hopanols (e.g., 17α(H)-21β(H)-bishomo-hopanol and 17β(H)-21β(h)-bishomo-hopanol), reflecting a characteristic suitably agreeing with patterns found at other seep sites (Fig. 11.6; Birgel et al. 2008, 2011; Himmler et al. 2015; Natalicchio et al. 2015). However, bacteriohopanepolyols (BHPs) and their related geohopanoids are known to be produced by both aerobic and anaerobic bacteria (Talbot et al. 2001; Sinninghe Damste et al. 2004; Blumenberg et al. 2006; Rush et al. 2016). Therefore, diplopterol, diploptene, hopanols, hopanoic acids, and their possible diagenetic products generally cannot be used alone as reliable indicators of past aerobic methanotrophic bacteria at seeps (Birgel and Peckmann 2008). Generally, 13C-depleted 3β-methylated hopanoids, 4-methylated steroids, and BHPs aminotetrol/aminotriol and further related BHPs are diagnostic biomarkers of aerobic methanotrophic bacteria (Talbot et al. 2001; Birgel and Peckmann 2008; Birgel et al. 2011; Himmler et al. 2015; Natalicchio et al. 2015; Rush et al. 2016). In the SCS, hopanoids were observed in seep carbonates and sediments originating from NE Dongsha, Shenhu, Site F, and Haima, yielding δ13C values ranging from −31‰ to −101‰ (Fig. 11.5; Guan et al. 2013, 2014, 2016a, b, 2018, 2022). Regarding the Haima cold seeps, 4-methylated steroid 4α-methylcholesta-2,8(14),24-triene and two tetracyclic hopanoid ketones, C30-3β-methyl tetracyclic-ketone and C34-3β-methyl tetracyclic-ketone, were found in seep sediments (Guan et al. 2022). In this cold seep ecosystem, both 4-methylated steroid and tetracyclic hopanoid ketones were attributed to Type I and/or X methanotrophs, with precursors most likely including 4α-methylcholesta-8(14),24-dien-3β-ol and 3-methylated hopanoids, respectively (Guan et al. 2022). These hopanoids along with aerobic methanotrophic bacteria-specific biomarkers are indicators of aerobic methanotrophic bacteria at seeps.
11.3 Authigenic Carbonate Formations Constrained by Lipid Biomarkers
11.3.1 Environmental Conditions Inferred from Biomarker Patterns
Microbial activity, diversity and related geochemical processes in cold seep ecosystems are of great concern because the upward ascending methane is mainly consumed by anaerobic methanotrophs in consortia with sulfate-reducing bacteria (Boetius et al. 2000). In the cold seep ecosystems of the SCS, microbial diversity has been reported at the NE Dongsha, Site F, Shenhu, and Haima cold seeps. Methanotrophic archaea (ANME-1, ANME-2, and ANME-3) and sulfate-reducing partners (Desulfosarcina/Desulfococcus) were observed, and their abundance varied with the depth and site (Niu et al. 2017; Cui et al. 2019; Zhuang et al. 2019). Processes of methanogenesis, AOM, and sulfate reduction could be archived by microbial activities at active seep sites (Boetius et al. 2000; Blumenberg et al. 2004). However, these advantages of microbial tools disappear during rock periods. Instead, lipid biomarkers are preferred because of their stable carbon skeleton and the carried information on their precursors (Aloisi et al. 2000; Peckmann et al. 2004, 2009; Birgel et al. 2008; Stadnitskaia et al. 2008a, b).
The lipid biomarker inventories of seep carbonates from NE Dongsha, Shenhu, Site F, and Haima revealed diagnostic lipids of specific groups of ANMEs and their sulfate-reducing partners (Fig. 11.1; Birgel et al. 2008; Yu et al. 2008; Guan et al. 2013, 2016a, b, 2018; Ge et al. 2015; Xu et al. 2022b). The biomarker patterns and carbon stable isotope compositions were found to vary among individual study sites and among different geographic areas (Figs. 11.3 and 11.5). Lipid biomarkers of ANMEs and SRBs originating from NE Dongsha and Site F carbonates commonly yielded lower δ13C values than those originating from Haima carbonates, reflecting differences in fluid compositions and methane types (Fig. 11.2; Guan et al. 2013, 2016a, 2018). NE Dongsha and Site F are located in the convergence zone of the active and passive margins, and seeps appear to be largely controlled by strike-slip faults (Liu et al. 2008, 2015; Chen et al. 2017; Feng et al. 2018). Methane at both sites indicates biogenic sources and is released from gas hydrate dissociation with δ13C values ranging from −72.3‰ to −69.4‰ (Zhuang et al. 2016). Regarding the Haima cold seeps, hydrate-bound methane is a mixture of biogenic and thermogenic gas, with most δ13C values ranging from −72.3‰ to −48.4‰ (Fang et al. 2019; Ye et al. 2019; Wei et al. 2019, 2021; Lai et al. 2022a, b). Potential methane types during carbonate precipitation could be inferred according to specific microbial consortia and δ13C values of archaea-specific isoprenoids because the δ13C differences from methane to archaeal-specific lipids reach approximately −30‰ and −50‰ for ANME-1 and ANME-2, respectively (Niemann and Elvert 2008; Birgel et al. 2011; Himmler et al. 2015). The δ13C variability of lipid biomarkers originating from NE Dongsha, Site F, and Haima carbonates reflected the patterns of their parent methane and carbon fractionations from methane to lipids.
The application of molecular fossils in combination with their compound-specific isotope signatures is an efficient tool to reconstruct seepage intensities. At seep sites, the sulfate methane transition zone (SMTZ) is largely controlled by the seepage intensity (Borowski et al. 1996), which tends to occur shallower within sediments at sites with a high methane flux than that at sites with a relatively low methane flux (Luff and Wallmann 2003). The relatively high sulfate concentration and porosity on or near the seafloor surface favor the precipitation of aragonite over high-Mg-calcite (Fig. 11.7; Greinert et al. 2001; Haas et al. 2010). In regard to microbial consortia, ANME-2 is better adapted to higher methane concentrations and lower temperatures and is less sensitive to oxygen than ANME-1, whereas ANME-1 can tolerate moderate and lower methane concentrations (Elvert et al. 2005; Knittel et al. 2005; Nauhaus et al. 2005). In the SCS, including the Dongsha area, Site F, Shenhu, and Haima cold seeps, aragonite- and calcite-dominated carbonates are commonly formed at sites predominated by ANME-2 and ANME-1, respectively, indicating varying environmental conditions and seepage intensities (Guan et al. 2013, 2016a, b, 2018; Ge et al. 2015). Therefore, lipid biomarkers, combined with mineralogy, petrography, and carbon stable and oxygen isotope compositions, are effective indicators to reconstruct the formation conditions of seep-related carbonates.
11.3.2 Sulfate Reductions Coupled to AOM and Organoclastic Oxidation
At marine seep sites, both AOM and oxidation of non-methane hydrocarbons can fuel sulfate reduction (Joye et al. 2004; Kallmeyer and Boetius 2004; Bowles et al. 2011; Kleindienst and Knittel 2020). Joye et al. (2004) found that sulfate-reducing rates generally exceeded AOM rates and concluded that the majority of sulfate reduction at Gulf of Mexico hydrocarbon seep sites was likely fueled by the oxidation of other organic matter. At oil-related cold seeps, hydrocarbons other than methane are the most likely reductants. Among sulfate reducers, DSS of Deltaproteobacteria are known as the primary non-methane hydrocarbon degraders at methane seeps (Kniemeyer et al. 2007; Kleindienst et al. 2012, 2014; Jaekel et al. 2013). DSS involved in sulfate-driven AOM lives in consortia with ANME, using CO2 as a carbon source and yielding extreme 13C-depletion of DSS-derived fatty acids (Wegener et al. 2008, 2015). In contrast, all known DSS capable of non-methane hydrocarbon degradation are free-living organisms, coupling substrate oxidization and sulfate reduction in a single-cell process (Kleindienst et al. 2012, 2014; Jaekel et al. 2013; Petro et al. 2019). These DSS were demonstrated to assimilate carbon from short-chain hydrocarbons in enrichment cultures (Jaekel et al. 2013; Kleindienst et al. 2014).
Although the δ13C values of methane and oil-related hydrocarbons overlap to a certain extent (−110‰ to −15‰ and higher than −35‰, respectively; Milkov and Etiope (2018)), methane in the SCS yielded notably lower carbon stable isotopes (lower than −48‰) than oil-related hydrocarbons (Zhuang et al. 2016; Yang et al. 2018; Fang et al. 2019; Wei et al. 2019, 2021; Ye et al. 2019; Lai et al. 2022a, b; Liang et al. 2022). Consequently, the lipid biomarkers of DSS involved in AOM and oxidation of non-methane hydrocarbons are expected to be different. Guan et al. (2018) analyzed seep carbonates originating from the Haima cold seeps and found that i-/ai-C15:0-fatty acids were accompanied by less negative δ13C values, suggesting that the input of DSS was involved in the oxidation of non-methane hydrocarbons. Similar observations were also found for sediments obtained from the Haima cold seeps (Guan et al. 2022). These conclusions were supported by lipid biomarker comparisons between methane-seep carbonates and oil-seep carbonates (Krake et al. 2022). The authors found that oil-seep-related carbonates not only yielded a high abundance of DAGEs with more diverse alkyl chains but also heavier δ13C values on average than those of methane-seep carbonates (Krake et al. 2022).
Unresolved complex mixtures (UCMs) are an additional indicator of the oxidation of non-methane hydrocarbons. UCMs are commonly reported in seep-related carbonates or sediments, especially oil seepages (Sassen et al. 1993; Zhang et al. 2003; Naehr et al. 2009; Birgel et al. 2011; Feng et al. 2014, 2018; Li et al. 2021). UCMs contain naphthenes and other related compounds that cannot be completely resolved by conventional GC analysis (Zhang et al. 2003; Naehr et al. 2009; Birgel et al. 2011). At the Haima cold seeps, UCMs occurred in hydrocarbon fractions of both seep carbonates and sediments, with partially altered hydrocarbons (Guan et al. 2018, 2022). Since n-alkanes were not completely altered, these oils in the sediments were attributed to light to middle biodegradation (Peters and Moldowan 1993). Furthermore, the heavier δ13CTIC values of sediments retrieved from the Haima seeps were found to be correlated with a high total organic carbon content, which again agreed with non-methane hydrocarbons supplied to sulfate-reducing bacteria (DSS) and resulted in relatively heavier δ13C values of DSS-derived fatty acids (Guan et al. 2022).
11.3.3 Constraints on the Hydrolysis Process
In lipid biomarker analysis of authigenic carbonates originating from the Haima cold seep, 3,7,11,15-tetramethylhexadecan-1,3-diol and two novel sn2-/sn3-O-hydroxyphytanyl glycerol monoethers hydroxylated at C-3 of the phytanyl moieties were identified. Both methanogens and methanotrophic archaea were found to produce hydroxydiethers (Sprott et al. 1990, 1992, 1993; Rossel et al. 2008, 2011). Among methanotrophic archaea, ANME-1 contains abundant GDGTs and small amounts of sn2-/sn3-hydroxyarchaeols, whereas ANME-2 and ANME-3 are dominated by phosphate-based polar derivatives of di-hydroxyarchaeol, sn2-/sn3-hydroxyarchaeols and archaeol (Rossel et al. 2008, 2011). Regarding these Haima seep carbonates, the abundant crocetane and high ratios of sn2-hydroxyarchaeol relative to archaeol (>1.1) indicated the dominance of ANME-2 or ANME-3 (Xu et al. 2022b).
Generally, understanding the composition of molecular fossils, biological precursors and past diagenetic evolution process constitutes a key step in applying a fossil lipid indicator (Briggs and Summons 2014). The 13C-depleted short-chain isoprenoids in cold seep ecosystems are putative degradation products of archaeol and sn2-/sn3-hydroxyarchaeols, and can be attributed to ANMEs (Thiel et al. 2001a, 2018; Oba et al. 2006). According to Koga et al. (1993) and Oba et al. (2006), all hydroxyarchaeols were converted into monophytanyl glycerol ethers in the hydrolysis process with chloroform/methanol/concentrated HCl. Based on the sequential degradation pathway of hydroxyarchaeols, sn2-/sn3-O-hydroxyphytanyl glycerol monoethers are likely hydrolysis products of di-hydroxyarchaeol rather than sn2-/sn3-hydroxyarchaeols, whereas isoprenoidal 3,7,11,15-tetramethylhexadecan-1,3-diol is most likely a hydrolysis product of sn2-/sn3-O-hydroxyarchaeols, di-hydroxyarchaeol, and sn2-/sn3-O-hydroxyphytanyl glycerol monoethers (Fig. 11.8; Xu et al. 2022b). Except for these isoprenoid ethers, a similar relationship was found for MAGEs, DAGEs and corresponding non-isoprenoid short-chain alcohols in the Haima seep carbonates. The MAGEs, DAGEs, and non-isoprenoid short-chain alcohols yielded similar δ13C values, chain (side-chain) lengths and unsaturation patterns, indicating that non-isoprenoid alcohols might be either biosynthetic intermediates or degradation products of MAGEs and DAGEs (Xu et al. 2022b).
11.4 Summary and Outlook
Authigenic carbonates sampled from the SCS contained varying contents of 13C-depleted archaeal and bacterial biomarkers, indicating their formation as a result of anaerobic oxidation of methane (AOM) via anaerobic methane-oxidizing archaea (ANMEs) and sulfate-reducing bacteria (SRBs). Based on various biomarker patterns, methane was predominantly oxidized by ANME-1/DSS and ANME-2/DSS, varying from site to site. Generally, ANME-2/DSS-specific crocetane, n-C16:1ω5, cyc-C17:0ω5,6-fatty acids, high sn2-hydroxyarchaeol/archaea ratios and low anteiso-C15:0/iso-C15:0-fatty acid (ai-/i-C15:0) ratios were found in aragonite-dominated carbonates, suggesting carbonates formed in seep ecosystems with high seepage intensities. Carbonates characterized by GDGTs, small amounts of crocetane, low sn2-hydroxyarchaeol/archaea ratios and high ai-C15:0/i-C15:0-fatty acid ratios were dominated by high-magnesium calcite, reflecting their formation at a greater depth with a relatively low seepage intensity. The oxidation of non-methane hydrocarbons fueled by sulfate reduction was demonstrated by abundant i-/ai-C15:0 fatty acids with relatively higher δ13C values. Distinct biomarker patterns and respective geochemical processes at cold seeps may be correlated with the microbial community succession sequence during seepage evolution because the different growth rates of aerobic methanotrophs, SRBs, and ANMEs could be recorded in an emerging cold seep ecosystem. Moreover, the ether hydrolysis products isoprenoidal 3,7,11,15-tetramethylhexadecan-1,3-diol and sn2-/sn3-O-hydroxyphytanyl glycerol monoethers were observed, reflecting a diagenetic process from isoprenoid dieters to isoprenoid monoethers and isoprenoid alcohols.
Although diagnostic biomarkers could be used to discriminate ANME-1/DSS and ANME-2/DSS consortia, it is unclear what factors control their ability to respond to changing seepage intensities and environments. Since cold seeps comprise a substantial carbon pool worldwide, it is critical to increase the efficiency of hydrocarbon biofilters. In addition to the well-advanced identification of sulfate-AOM, AeOM, and oxidation of non-methane hydrocarbons with biomarkers as outlined above, future studies combining culturing, molecular microbiology, and lipid biomarkers may provide further insight into (1) the modes and variability of hydrocarbon consumption possibly involving different electron acceptors and (2) the allochthonous and autochthonous contributions of seep carbonates to biomarker inventories.
References
Aloisi G, Pierre C, Rouchy JM et al (2000) Methane–related authigenic carbonates of eastern Mediterranean Sea mud volcanoes and their possible relation to gas hydrate destabilisation. Earth Planet Sci Lett 184(1):321–338
Birgel D, Elvert M, Han XQ et al (2008) 13C–depleted biphytanic diacids as tracers of past anaerobic oxidation of methane. Org Geochem 39(1):152–156
Birgel D, Peckmann J (2008) Aerobic methanotrophy at ancient marine methane seeps: a synthesis. Org Geochem 39(12):1659–1667
Birgel D, Feng D, Roberts HH et al (2011) Changing redox conditions at cold seeps as revealed by authigenic carbonates from Alaminos Canyon, Northern Gulf of Mexico. Chem Geol 285(1–4):82–96
Birgel D, Thiel V, Hinrichs K-U et al (2006) Lipid biomarker patterns of methane–seep microbialites from the Mesozoic convergent margin of California. Org Geochem 37(10):1289–1302
Blumenberg M, Krüger M, Nauhaus K et al (2006) Biosynthesis of hopanoids by sulfate–reducing bacteria (genus Desulfovibrio). Environ Microbiol 8(7):1220–1227
Blumenberg M, Seifert R, Nauhaus K et al (2005) In vitro study of lipid biosynthesis in an anaerobically methane-oxidizing microbial mat. Appl Environ Microbiol 71(8):4345–4351
Blumenberg M, Seifert R, Reitner J et al (2004) Membrane lipid patterns typify distinct anaerobic methanotrophic consortia. Proc Natl Acad Sci USA 101(30):11111–11116
Blumenberg M, Walliser E, Taviani M et al (2015) Authigenic carbonate formation and its impact on the biomarker inventory at hydrocarbon seeps e a case study from the Holocene Black Sea and the Plio-Pleistocene Northern Apennines (Italy). Mar Pet Geol 66:532–541
Boetius A, Ravenschlag K, Schubert CJ et al (2000) A marine microbial consortium apparently mediating anaerobic oxidation of methane. Nature 407(6804):623–626
Boetius A, Wenzhöfer F (2013) Seafloor oxygen consumption fuelled by methane from cold seeps. Nat Geosci 6(9):725–734
Borowski WS, Paul CK, Ussler WU (1996) Marine pore–water sulfate profles indicate in situ methane flux from underlying gas hydrate. Geology 24(7):655–658
Bowles MW, Samarkin VA, Bowles KM et al (2011) Weak coupling between sulfate reduction and the anaerobic oxidation of methane in methane–rich seafloor sediments during ex situ incubation. Geochim Cosmochim Acta 75(2):500–519
Briggs DEG, Summons RE (2014) Ancient biomolecules: their origins, fossilization, and role in revealing the history of life. BioEssays 36(5):482–490
Chen DF, Huang YY, Yuan XL et al (2005) Seep carbonates and preserved methane oxidizing archaea and sulfate reducing bacteria fossils suggest recent gas venting on the seafloor in the Northeastern South China Sea. Mar Pet Geol 22(5):613–621
Chen WH, Huang CY, Yan Y et al (2017) Stratigraphy and provenance of forearc sequences in the Lichi Melange, Coastal Range: geological records of the active Taiwan arc–continent collision. J Geophys Res-Solid Earth 122(9):7408–7436
Chevalier N, Bouloubassi I, Stadnitskaia A et al (2010) Distributions and carbon isotopic compositions of lipid biomarkers in authigenic carbonate crusts from the Nordic Margin (Norwegian Sea). Org Geochem 41(9):885–890
Chevalier N, Bouloubassi I, Birgel D et al (2013) Microbial methane turnover at Marmara Sea cold seeps: a combined 16S rRNA and lipid biomarker investigation. Geobiology 11(1):55–71
Cui HP, Su X, Chen F et al (2019) Microbial diversity of two cold seep systems in gas hydrate–bearing sediments in the South China Sea. Mar Environ Res 144:230–239
Elvert M, Boetius A, Knittel K et al (2003) Characterization of specific membrane fatty acids as chemotaxonomic markers for sulfate–reducing bacteria involved in anaerobic oxidation of methane. Geomicrobiol J 20(4):403–419
Elvert M, Hopmans EC, Treude T et al (2005) Spatial variations of methanotrophic consortia at cold methane seeps: implications from a high–resolution molecular and isotopic approach. Geobiology 3(3):195–209
Fang Y, Wei J, Lu H et al (2019) Chemical and structural characteristics of gas hydrates from the Haima cold seeps in the Qiongdongnan Basin of the South China Sea. J Asian Earth Sci 182:103924
Feng D, Birgel D, Peckmann J et al (2014) Time integrated variation of sources of fluids and seepage dynamics archived in authigenic carbonates from Gulf of Mexico Gas Hydrate Seafloor Observatory. Chem Geol 385:129–139
Feng D, Qiu J-W, Hu Y et al (2018) Cold seep systems in the South China sea: an overview. J Asian Earth Sci 168:3–16
Ge L, Jiang SY, Blumenberg M et al (2015) Lipid biomarkers and their specific carbon isotopic compositions of cold seep carbonates from the South China sea. Mar Pet Geol 66:501–510
Ge L, Jiang S, Swennen R et al (2010) Chemical environment of cold seep carbonate formation on the northern continental slope of South China sea: evidence from trace and rare earth element geochemistry. Mar Geol 277(1–4):21–30
Ge L, Jiang S, Yang T et al (2011) Glycerol ether biomarkers and the carbon isotopic compositions in a cold seep carbonate chimney from the Shenhu area, northern South China Sea. Chin Sci Bull 56(16):1700–1707
Greinert J, Bohrmann G, Suess E (2001) Gas hydrate associated carbonates and methane–venting at Hydrate Ridge: classification, distribution and origin of authigenic lithologies. In: Paull CK, Dillon WP (eds) Natural gas hydrates: occurrence, distribution, and detection. American Geophysical Union, Washington, DC, USA, pp 91–113
Guan H, Feng D, Wu N et al (2016a) Methane seepage intensities traced by biomarker patterns in authigenic carbonates from the South China sea. Org Geochem 91:109–119
Guan H, Zhang M, Mao S et al (2016b) Methane seepage in the Shenhu area of the northern South China sea: constraints from carbonate chimneys. Geo-Mar Lett 36(3):175–186
Guan H, Liu L, Hu Y et al (2022) Rising bottom–water temperatures induced methane release during the middle Holocene in the Okinawa Trough. East China Sea. Chem Geol 590:120707
Guan H, Sun Y, Mao S et al (2014) Molecular and stable carbon isotopic compositions of hopanoids in seep carbonates from the South China Sea continental slope. J Asian Earth Sci 92:254–261
Guan H, Sun Y, Zhu X et al (2013) Factors controlling the types of microbial consortia in cold–seep environments: a molecular and isotopic investigation of authigenic carbonates from the South China Sea. Chem Geol 354:55–64
Guan H, Xu L, Wang Q et al (2019) Lipid Biomarkers and Their Stable Carbon Isotopes in Ancient Seep Carbonates from SW Taiwan. China. Acta Geol Sin-Engl Ed 93(1):167–174
Guan H, Birgel D, Peckmann J et al (2018) Lipid biomarker patterns of authigenic carbonates reveal fluid composition and seepage intensity at Haima cold seeps, South China Sea. J Asian Earth Sci 168:163–172
Haas A, Peckmann J, Elvert M et al (2010) Patterns of carbonate authigenesis at the Kouilou pockmarks on the Congo deep–sea fan. Mar Geol 268(1–4):129–136
Han X, Suess E, Huang Y et al (2008) Jiulong methane reef: Microbial mediation of seep carbonates in the South China Sea. Mar Geol 249(3–4):243–256
Himmler T, Birgel D, Bayon G et al (2015) Formation of seep carbonates along the Makran convergent margin, northern Arabian Sea and a molecular and isotopic approach to constrain the carbon isotopic composition of parent methane. Chem Geol 415:102–117
Hinrichs K–U, Boetius A (2002) The anaerobic oxidation of methane: new insights in microbial ecology and biogeochemistry. In: Wefer G, Billett D, Hebbeln D, Jørgensen BB, Schlüter M, van Weering, T.C.E. (Eds)Ocean Margin Systems. Springer, pp 457–477
Hinrichs K-U, Summons RE, Orphan V et al (2000) Molecular and isotopic analysis of anaerobic methane–oxidizing communities in marine sediments. Org Geochem 31(12):1685–1701
Jaekel U, Musat N, Adam B et al (2013) Anaerobic degradation of propane and butane by sulfate–reducing bacteria enriched from marine hydrocarbon cold seeps. ISME J 7(5):885–895
Jin M, Feng D, Huang KJ et al (2022) Magnesium Isotopes in Pore Water of Active Methane Seeps of the South China Sea. Front Mar Sci 9:858860
Joye SB (2020) The geology and biogeochemistry of hydrocarbon seeps. Annu Rev Earth Planet Sci 48:205–231
Joye SB, Boetius A, Orcutt BN et al (2004) The anaerobic oxidation of methane and sulfate reduction in sediments from Gulf of Mexico cold seeps. Chem Geol 205(3–4):219–238
Kallmeyer J, Boetius A (2004) Effects of temperature and pressure on sulfate reduction and anaerobic oxidation of methane in hydrothermal sediments of Guaymas Basin. Appl Environ Microbiol 70(2):1231–1233
Kleindienst S, Herbst F-A, Stagars M et al (2014) Diverse sulfatereducing bacteria of the Desulfosarcina/Desulfococcus clade are the key alkane degraders at marine seeps. ISME J 8(10):2029–2044
Kleindienst S, Knittel K (2020) Anaerobic hydrocarbon–degrading sulfate–reducing bacteria at marine gas and oil seeps. In: Teske A, Carvalho V (eds) Marine Hydrocarbon Seeps: Microbiology and Biogeochemistry of a Global Marine Habitat, Springer Oceanography. Springer International Publishing, Cham, pp 21–41
Kleindienst S, Ramette A, Amann R et al (2012) Distribution and in situ abundance of sulfate–reducing bacteria in diverse marine hydrocarbon seep sediments. Environ Microbiol 14(10):2689–2710
Kniemeyer O, Musat F, Sievert SM et al (2007) Anaerobic oxidation of short–chain hydrocarbons by marine sulphate–reducing bacteria. Nature 449(7164):898–901
Knittel K, Boetius A, Lemke A et al (2003) Activity, distribution, and diversity of sulfate reducers and other bacteria in sediments above gas hydrate (Cascadia margin, Oregon). Geomicrobiol J 20(4):269–294
Knittel K, Loekann T, Boetius A et al (2005) Diversity and distribution of methanotrophic archaea at cold seeps. Appl Environ Microbiol 71(1):467–479
Koga Y, Nishihara M, Morii H et al (1993) Ether polar lipids of methanogenic bacteria: Structures, comparative aspects, and biosyntheses. Microbiol Rev 57(1):164–182
Krake N, Birgel D, Smrzka D et al (2022) Molecular and isotopic signatures of oil–driven bacterial sulfate reduction at seeps in the southern Gulf of Mexico. Chem Geol 595:120797
Lai H, Fang Y, Kuang Z et al (2021) Geochemistry, origin and accumulation of natural gas hydrates in the Qiongdongnan Basin, South China Sea: Implications from site GMGS5–W08. Mar Pet Geol 123:104774
Lai H, Qiu H, Liang J et al (2022a) Geochemical Characteristics and Gas–to–Gas Correlation of Two Leakage–type Gas Hydrate Accumulations in the Western Qiongdongnan Basin, South China Sea. Acta Geol Sin – Engl Ed 96(2):680–690
Lai H, Qiu H, Kuang Z et al (2022b) Integrated signatures of secondary microbial gas within gas hydrate reservoirs: A case study in the Shenhu area, northern South China Sea. Mar Pet Geol 136:105486
Li Y, Fang YX, Zhou QZ et al (2021) Geochemical insights into contribution of petroleum hydrocarbons to the formation of hydrates in the Taixinan Basin, the South China Sea. Geosci Front 12(6):100974
Liang Q, Xiao X, Zhao J et al (2022) Geochemistry and sources of hydrate-bound gas in the shenhu area, northern south China sea: Insights from drilling and gas hydrate production tests. J Pet Sci Eng 208:109459
Liang Q, Hu Y, Feng D et al (2017) Authigenic carbonates from newly discovered active cold seeps on the northwestern slope of the South China Sea: Constraints on fluid sources, formation environments, and seepage dynamics. Deep-Sea Research Part I-Oceanogr Res Pap 124:31–41
Liu C, Meng Q, He X et al (2015) Characterization of natural gas hydrate recovered from Pearl River Mouth basin in South China Sea. Mar Pet Geol 61:14–21
Liu C, Morita S, Liao YH et al (2008) High resolution seismic images of the Formosa ridge off southwestern Taiwan where “hydrothermal” chemosynthetic community is present at a cold seep site. In: Proceedings of the 6th International Conference on Gas Hydrates (ICGH 2008), Vancouver, British Columbia, July, pp 6–10. doi: https://doi.org/10.14288/1.0041106
Losekann T, Knittel K, Nadalig T et al (2007) Diversity and abundance of aerobic and anaerobic methane oxidizers at the Haakon Mosby mud volcano. Barents Sea. Appl Environ Microbiol 73(10):3348–3362
Luff R, Wallmann K (2003) Fluid flow, methane fluxes, carbonate precipitation and biogeochemical turnover in gas hydrate–bearing sediments at Hydrate Ridge, Cascadia Margin: numerical modeling and mass balances. Geochim Cosmochim Acta 67(18):3403–3421
Milkov AV, Etiope G (2018) Revised genetic diagrams for natural gases based on a global dataset of >20,000 samples. Org Geochem 125:109–120
Naehr TH, Birgel D, Bohrmann G et al (2009) Biogeochemical controls on authigenic carbonate formation at the Chapopote asphalt volcano. Bay of Campeche. Chem Geol 266(3–4):390–402
Natalicchio M, Peckmann J, Birgel D et al (2015) Seep deposits from northern Istria, Croatia: a frst glimpse into the Eocene seep fauna of the Tethys region. Geol Mag 152(3):444–459
Nauhaus K, Treude T, Boetius A et al (2005) Environmental regulation of the anaerobic oxidation of methane: a comparison of ANME–I and ANME–II communities. Environ Microbiol 7(1):98–106
Niemann H, Elvert M (2008) Diagnostic lipid biomarker and stable carbon isotope signatures of microbial communities mediating the anaerobic oxidation of methane with sulphate. Org Geochem 39(12):1668–1677
Niu M, Fan X, Zhuang G et al (2017) Methane–metabolizing microbial communities in sediments of the Haima cold seep area, northwest slope of the South China Sea. FEMS Microbiol Ecol 93(9): fix101
Oba M, Sakata S, Tsunogai U (2006) Polar and neutral isopranyl glycerol ether lipids as biomarkers of archaea in near–surface sediments from the Nankai Trough. Org Geochem 37(12):1643–1654
Orphan VJ, House CH, Hinrichs K-U et al (2001) Methane–consuming archaea revealed by directly coupled isotopic and phylogenetic analysis. Science 293(5529):484–487
Peckmann J, Birgel D, Kiel S (2009) Molecular fossils reveal fluid composition and flow intensity at a Cretaceous seep. Geology 37(9):847–850
Peckmann J, Thiel V, Reitner J et al (2004) A microbial mat of a large sulfur bacterium preserved in a Miocene methane–seep limestone. Geomicrobiol J 21(4):247–255
Peters KE, Moldowan JM (1993) The Biomarker Guide: Interpreting Molecular Fossils in Petroleum and Ancient Sediments. Prentice Hall, Englewood Cliffs, NJ. ISBN 0-13-086752-7
Petro C, Jochum LM, Schreiber L et al (2019) Single–cell amplifed genomes of two uncultivated members of the deltaproteobacterial SEEP–SRB1 clade, isolated from marine sediment. Mar Genom 46:66–69
Reeburgh WS (2007) Oceanic methane biogeochemistry. Chem Rev 107(2):486–513
Rossel PE, Elvert M, Ramette A et al (2011) Factors controlling the distribution of anaerobic methanotrophic communities in marine environments: evidence from intact polar membrane lipids. Geochim Cosmochim Acta 75(1):164–184
Rossel PE, Lipp JS, Fredricks HF et al (2008) Intact polar lipids of anaerobic methanotrophic archaea and associated bacteria. Org Geochem 39(8):992–999
Rush D, Osborne KA, Birgel D et al (2016) The bacteriohopanepolyol inventory of novel aerobic methane oxidizing bacteria reveals new biomarker signatures of aerobic methanotrophy in marine systems. PLoS ONE 11(11):e0165635
Sassen R, Roberts HH, Aharon P et al (1993) Chemosynthetic bacterial mats at cold hydrocarbon seeps, Gulf of Mexico continental slope. Org Geochem 20(1):77–89
Sinninghe Damsté JS, Rijpstra WIC, Schouten S et al (2004) The occurrence of hopanoids in planctomycetes: implications for the sedimentary biomarker record. Org Geochem 35(5):561–566
Suess E (2005) RV SONNE cruise report SO 177, Sino–German cooperative project, South China Sea Continental Margin: Geological methane budget and environmental effects of methane emissions and gas hydrates. IFM–GEOMAR Reports. doi: https://doi.org/10.3289/ifm-geomar_rep_4_2005
Sprott GD (1992) Structures of archaebacterial membrane lipids. J Bioenerg Biomembr 24(6):555–566
Sprott GD, Dicaire CJ, Choquet CG et al (1993) Hydroxydiether lipid structures in Methanosarcina spp. and Methanococcus voltae. Appl Environ Microbiol 59 (3):912–914
Sprott GD, Ekiel I, Dicaire C (1990) Novel, acid–labile, hydroxydiether lipid cores in methanogenic bacteria. J Bioll Chem 265(23):13735–13740
Stadnitskaia A, Bouloubassi I, Elvert M et al (2008a) Extended hydroxyarchaeol, a novel lipid biomarker for anaerobic methanotrophy in cold seepage habitats. Org Geochem 39(8):1007–1014
Stadnitskaia A, Nadezhkin D, Abbas B et al (2008b) Carbonate formation by anaerobic oxidation of methane: evidence from lipid biomarker and fossil 16S rDNA. Geochim Cosmochim Acta 72(7):1824–1836
Talbot HM, Watson DF, Murrell JC et al (2001) Analysis of intact bacteriohopanepolyols from methanotrophic bacteria by reversed–phase highperformance liquid chromatography–atmospheric pressure chemical ionization mass spectrometry. J Chromatogr A 921(2):175–185
Thiel V (2018) Methane carbon cycling in the past: Insights from hydrocarbon and lipid biomarkers. In: Wilkes H (ed) Hydrocarbons, oils and lipids: diversity, origin, chemistry and fate. Switzerland, Springer International Publishing, Cham, pp 1–30
Thiel V, Peckmann J, Richnow HH et al (2001a) Molecular signals for anaerobic methane oxidation in Black Sea seep carbonates and a microbial mat. Mar Chem 73(2):97–112
Thiel V, Peckmann J, Schmale O et al (2001b) A new straight–chain hydrocarbon biomarker associated with anaerobic methane cycling. Org Geochem 32(8):1019–1023
Wallmann K, Pinero E, Burwicz E et al (2012) The Global Inventory of Methane Hydrate in Marine Sediments: A Theoretical Approach. Energies 5(7):2449–2498
Wang X, Guan H, Qiu JW et al (2022) Macro–ecology of cold seeps in the South China Sea. Geosystems and Geoenvironment 1(3):100081
Wegener G, Krukenberg V, Riedel D et al (2015) Intercellular wiring enables electron transfer between methanotrophic archaea and bacteria. Nature 526(7574):587–590
Wegener G, Niemann H, Elvert M et al (2008) Assimilation of methane and inorganic carbon by microbial communities mediating the anaerobic oxidation of methane. Environ Microbiol 10(9):2287–2298
Wei J, Liang J, Lu J et al (2019) Characteristics and dynamics of gas hydrate systems in the northwestern South China Sea-Results of the fifth gas hydrate drilling expedition. Mar Pet Geol 110:287–298
Wei J, Wu T, Zhu L et al (2021) Mixed gas sources induced co–existence of sI and sII gas hydrates in the Qiongdongnan Basin. South China Sea. Mar Pet Geol 128:105024
Wu N, Xu C, Li A et al (2022) Oceanic carbon cycle in a symbiotic zone between hydrothermal vents and cold seeps in the Okinawa Trough. Geosystems and Geoenvironment 1(3):100059
Wu N, Zhang H, Yang S et al (2011) Gas hydrate system of Shenhu area, northern South China Sea: Geochemical results. Journal of Geological Research 2011:370298
Xu LF, Guan HX, Liu L et al (2022a) Determining the double–bond positions of monounsaturated compounds in the alcohol fraction in seep carbonate. J Chromatogr A 1672:463009
Xu L, Guan H, Su Z et al (2022b) Diagenetic fate of glycerol ethers revealed by two novel isoprenoid hydroxyphytanyl glycerol monoethers and non–isoprenoid alkyl glycerol ethers. Org Geochem 163:104344
Yang K, Chu F, Zhu Z et al (2018) Formation of methane–derived carbonates during the last glacial period on the northern slope of the South China Sea. J Asian Earth Sci 168:173–185
Ye J, Wei J, Liang J et al (2019) Complex gas hydrate system in a gas chimney, South China Sea. Mar Pet Geol 104:29–39
Yu X, Han X, Li H et al (2008) Biomarkers and carbon isotope composition of anaerobic oxidation of methane in sediments and carbonates of northeastern part of Dongsha, South China Sea. Acta Oceanol Sin 30:77–84
Zhang CL, Li YL, Ye Q et al (2003) Carbon isotope signatures of fatty acids in Geobacter metallireducens and Shewanella algae. Chem Geol 195(1–4):17–28
Zhang W, Liang J, Wei J et al (2020) Geological and geophysical features of and controls on occurrence and accumulation of gas hydrates in the frst offshore gas–hydrate production test region in the Shenhu area, Northern South China Sea. Mar Pet Geol 114:104191
Zhuang C, Chen F, Cheng S et al (2016) Light carbon isotope events of foraminifera attributed to methane release from gas hydrates on the continental slope, northeastern South China Sea. Sci. China-Earth Sci 59(10):1981–1995
Zhuang G, Xu L, Liang Q et al (2019) Biogeochemistry, microbial activity, and diversity in surface and subsurface deep–sea sediments of South China Sea. Limnol Oceanogr 64(5):2252–2270
Acknowledgements
We sincerely thank the crew and scientists of the ROV Haima, ROPOS and RV SONNE, Haiyang-06, and “Tan Kah Kee” for their professional work to meet the scientific goal during expeditions. S. Gao (GIG, CAS) and S. Liu (GIEC, CAS) are thanked for their help with technical assistance. This research was supported by Laoshan Laboratory (No. LSKJ202203502), the National Natural Science Foundation of China (Grants: 42276053 and 91958105), and the Young Taishan Scholars Program (Grant No. tsqn202211069).
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Guan, H., Liu, L., Wu, N., Li, S. (2023). Biomarker Indicators of Cold Seeps. In: Chen, D., Feng, D. (eds) South China Sea Seeps. Springer, Singapore. https://doi.org/10.1007/978-981-99-1494-4_11
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