4), although these might not necessarily exist in all bathyarchaeotal subgroups (Fig. They include Euryarchaeota, and members of the DPANN and Asgard archaea. 2017KZDXM071), and the Science and Technology Innovation Committee of Shenzhen (Grant No. The reconstructed bathyarchaeotal genomes (except for Subgroup-15) also encode proteins with the ability to import extracellular carbohydrates. Genomic characterization and metabolic potentials of Bathyarchaeota. The presence and relative abundance of bathyarchaeotal rRNA can then be estimated based on the hybridization intensity (Stahletal.1988; Kuboetal.2012). Bathyarchaeota are believed to have roles in the carbon cycle in marine systems. The wide availability of buried organic matter in the marine subsurface would favor the heterotrophic feeding of Bathyarchaeota. Bathyarchaeota dominate 16S rRNA clone libraries of transcribed RNA constructed for the Peru Margin ODP site 1229 (Parkesetal.2005; Biddleetal.2006) and the upper 35 m of the subsurface sediments at the Peru Margin ODP site 1227 (Inagakietal.2006; Sorensen and Teske 2006). WebArchaea (/ r k i / ar-KEE-; singular archaeon / r k i n /) is a domain of single-celled organisms.These microorganisms lack cell nuclei and are therefore prokaryotes.Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebacteria kingdom), but this term has fallen out of use.. Archaeal cells have The phylogenetic affiliation of sequences found in peat suggest that members of the thus-far-uncultivated group Candidatus Bathyarchaeota (representing a fourth phylum) may be involved in methane cycling, either anaerobic oxidation of methane and/or methanogenesis, as at least a few organisms within this group contain the essential Because of the high diversity of Bathyarchaeota and various independent analyses of samples from diverse environments, the nomenclature for this archaeal group in previous reports was very complex. (A) Phylogenetic tree of ribosomal proteins obtained from currently available bathyarchaeotal genomes (from GenBank, 29 November 2017 updated). Individual metagenome assemblies facts about bathyarchaeota The Bathyarchaeota formerly known as the Miscellaneous Crenarchaeotal Group is an evolutionarily diverse group of microorganisms found in a wide These indicative subgroups are the dominant ones in the environment, as evaluated by relatively abundant fraction of Bathyarchaeota in corresponding archaeal communities (on average 44% among all studies). Because of the wide distribution of this lipid in many other archaea, it cannot be used for the detection of Bathyarchaeota and its carbon stable isotopic composition cannot be used for metabolic property deductions. The identification of key genes of the MCR complex (mcrA, mcrB and mcrG), and the presence of hdrABC and mcvhADG responsible for the cycling of coenzyme M (CoM) and coenzyme B (CoB), suggest their role in the methanogenesis machinery that mediates the CoM-S-S-CoB cycling, similar to Euryarchaeota methanogens (Evansetal.2015). Furthermore, genes encoding ATP sulfurylase, for the reduction of sulfate to adenosine 5-phosphosulfate, and adenylyl-sulfate reductase, for the reduction of adenosine 5-phosphosulfate to sulfite, were identified in a metagenomic assembly of Bathyarchaeota TCS49 genome from the Thuwal cold seep brine pool of the Red Sea; this suggests that specific bathyarchaeotal members might harbor a dissimilatory sulfate reduction pathway, indicating the existence of additional potential metabolic capacities of Bathyarchaeota (Zhangetal.2016). Consequently, CO2 appears to be the only electron acceptor mediating AOM, like in a reverse acetoclastic methanogenesis (Hallametal.2004; Wangetal.2014). Community, Distribution, and Ecological Roles of Estuarine Archaea Devereux R, Mosher JJ, Vishnivetskaya TA et al. their relatively high abundance in the global marine subsurface ecosystem (Kuboetal.2012; Lloydetal.2013), they are also metabolically active in the subsurface sediments across geological time scales. High-throughput sequencing of the archaeal communities and the analysis of the relationship between the distribution pattern of bathyarchaeotal subgroups and the physicochemical parameters of study sites revealed that sediment depth and sulfate concentration were important environmental factors that shape the distribution of bathyarchaeotal subgroups; Subgroup-8 was shown to be predominantly distributed in the reducing and deeper sediment layers, while Subgroup-10 was preferentially distributed in the relatively more oxidizing and shallow sediment layers (Yuetal.2017). Subgroup-5 is divided into Subgroups-5a and -5b, each with intragroup similarity >90% according to a maximum-likelihood estimation. The Archaebacteria kingdom is divided into three S. butanivorans forms a distinct cluster with those of Bathyarchaeota origin, separately from other methanogens and methanotrophs (Laso-Prezetal.2016). The Miscellaneous Crenarchaeotal Group (MCG) archaea were firstly detected from a hot spring (Barnsetal.1996) and later proposed with a name in a study surveying 16S rRNA gene sequences from marine subsurface sediments (Inagakietal.2003). In addition to the global distribution, expanding prokaryotic community investigations of deep ocean drilling sediments revealed that members of Bathyarchaeota occupy considerable fractions of the archaeal communities (Teske 2006). Open reading frames encoded by the three fosmid clones comprised genes related to lipid biosynthesis, energy metabolism and resistance to oxidants. Background Bathyarchaeota, a newly proposed archaeal phylum, is considered as an important driver of the global carbon cycle. In this process, methane is not assimilated by Bathyarchaeota but serves as an energy source. The archaeal community structure, including Bathyarchaeota, is not correlated with a general geochemical categorization, but with the depth and sulfate concentration, subsequently linking to the redox potential, age and the (increasing) degree of organic matter recalcitrance. The phylum Bathyarchaeota, which has high species and functional diversity, is abundant and widespread in marine sediments. In one study, small amounts of stable isotope-labeled substrates, including glucose, acetate and CO2, were introduced multiple times into slurries from different biogeochemical depths of tidal sediments from the Severn estuary (UK) to better reflect the in situ environmental conditions (Websteretal.2010). A new phylum name for this group was proposed, i.e. 3) (Lloydetal.2013; Evansetal.2015; Lazaretal.2015; Heetal.2016; Lazaretal.2016; Lever 2016). Several pre-/non-enriched sediment cultures afforded preliminary evidence for the trophic properties and metabolic capacities of Bathyarchaeota. However, due to the great diversity of them, there is limited genomic information that accurately encompasses the metabolic potential of the entire archaeal phylum. Subgroup-5 thrives in the euxinic bottom water layer, characterized as anoxic and sulfide-rich, with accumulated inorganic and organic reduced compounds; Subgroup-6 is a group of generalists that are adapted to both planktonic and sediment habitats with a wide range of sulfidic conditions. To compare the coverage and specificity of analysis using the qPCR primer pairs MCG242dF/MCG678R and MCG528F/MCG732R for freshwater and marine sediment samples, amplicons obtained with these two primer pairs were analyzed and community structures compared (Filloletal.2015). 3C). The in silico tests revealed that primers MCG528, MCG493, MCG528 and MCG732 cover 87, 79, 44 and 27% of sequences of Subgroups-1 to -12 on average, respectively. Based on the above, it is proposed that Bathyarchaeota might mediate the AOM without assimilating the carbon in methane. Introduction. Furthermore, the lack of genes for ATPases and membrane-bound electron transport enzymes in the two genomic bins (BA1 and BA2) and the presence of the ion pumping, energy-converting hydrogenase complex (only in BA1), which might allow solute transportation independently of energy-generation mechanisms, suggest that the soluble substrate transportation is solely responsible for energy conservation (Evansetal.2015). This will have a profound impact not only on deciphering the metabolic properties of Bathyarchaeota, by using butanetriol dibiphytanyl glycerol tetraethers as biomarkers to trace carbon acquisition by isotopic labeling, but also by representing their pivotal contribution, associated with their global abundance, to biogeochemical carbon cycling on a large ecological scale. Bathyarchaeia occurrence in rich methane sediments Furthermore, both BA1 and BA2 lack ATP-synthase, indicating that they are restricted to substrate-level phosphorylation for energy, which was first found in methanogenic archaea (Evansetal.2015). Td stands for dissociation temperature for RNA slot-bolt. WebBathyarchaeota are abundant in sediments, and they may involve in sedimentary organic matter degradation, acetogenesis, and, potentially, methane metabolism, based on genomics. Fryetal. Recent data point to the global occurrence of Bathyarchaeota and their potential impact on global carbon transformation, highlighting their important role as a group of global generalists participating in carbon cycling, similar to euryarchaeotal methanogens and Thaumarchaeota. Further, based on genomic inferences, Evansetal. All assigned subgroups have minimum intra-group >90%, and are clustered into one clade with previously reported anchor sequences (Kuboetal.2012). Here we provide several lines of converging evidence suggesting the bathyarchaeotal group Bathy-8 is able to grow with lignin as an energy source and WebHost. It is known that a methane microbiome can be established in methane seeps sites; however, they are still poorly characterised. Proteins or polypeptides are first degraded by extracellular peptidases, with the resultant amino acids and oligopeptides imported into the cell, where they would be finally metabolized into acetyl-CoA via the peptide-degradation pathway. The possibility of the replacement of the AOM function of ANME by Bathyarchaeota was also suggested by a microbial community composition in a study of the microbial colonization within an artificial micro-niche, basaltic glass imposed by hydrothermal conditions (Callacetal.2013). Thaumarchaeota MG-I was present in the 12C-DNA library in the corresponding zone but was not detected in the 13C-DNA library, suggesting that these microbes are not able to use 13C-acetate (Websteretal.2010). Members of the Bathyarchaeota, formerly known as the Miscellaneous Crenarchaeota Group (MCG), are widely distributed in various environments such as freshwater lake, marine, and estuarine sediments [ 18, 19, 20, 21 ]. It has been proposed that the deduced last common ancestor was most likely a saline-adapted organism, and the evolutionary progression occurred most likely in the saline-to-freshwater direction, with few environmental transitional events. Diversity, metabolism and cultivation of archaea Methanogenesis and acetogenesis are considered to be the two most fundamental and ancient microbial biochemical energy conservation processes, and they both employ the WoodLjungdahl pathway for CO2 reduction and ATP generation (Weissetal.2016). This is the first ever genomic evidence for homoacetogenesis, the ability to solely utilize CO2 and H2 to generate acetate, in an archaeal genome and of distinct archaeal phylogenetic origin other than that of Bacteria (Heetal.2016). The primer pair MCG242dF/MCG528R may potentially be used for the determination of the bathyarchaeotal community abundance, with relatively high subgroup coverage and specificity in silico; however, experimental tests are needed to confirm this. Callac N, Rommevaux-Jestin C, Rouxel O et al. With respect to its function, the protein might be responsible for photosynthesis in archaea; this suggests that photosynthesis may have evolved before the divergence of the bacteria and archaea domains (Mengetal.2009; Lietal.2012). the most persistent detrital matter in marine sediments (Lomsteinetal.2012; Lloydetal.2013). Methanogens and acetogenic Clostridia are the most frequent basal-branching archaea and bacteria, respectively, in phylogenetic reconstructions reflecting the descendants of the last universal common ancestor; gene categories proposed for the last universal common ancestor also point to the acetogenic and methanogenic roots, reflecting its autotrophic lifestyle as H2-dependent and N2-fixing, utilizing the WoodLjungdahl pathway and originating from a hydrothermal environmental setting (Weissetal.2016). This would be supported by a coupled AOM and syntrophic SRB metabolism, with methane consumed by Bathyarchaeota through reverse acetoclastic methanogenesis with the production of acetate, which is readily oxidized by sulfate in SRB. Two highly abundant MCR variants were detected in Ca. Furthermore, another study demonstrated that the archaeal communities of the sulfatemethane transition zone at diffusion-controlled sediments of Aarhus Bay (Denmark) contain considerable amounts of Bathyarchaeota; the overall archaeal community structure did not change greatly during the experimentits diversity was lower after 6 months of incubation under heterotrophic conditions, with periodic modest sulfate and acetate additions (Websteretal.2011). Jacquemet A, Barbeau J, Lemiegre L et al. Members of Bathyarchaeota are able to use CO2 and H2 from natural sources and fermentation products to fuel acetogenesis (Heetal.2016; Martinetal.2016). They also acquired some subunits of coenzyme F420 hydrogenase; this enzyme generates reduced ferredoxin, with hydrogen as the electron donor, as an alternative to MvhADG in many Methanomicrobiales (Thaueretal.2008; Lazaretal.2016; Sousaetal.2016). Surprisingly, these genes fall closely to the Bathyarchaeota mcr genes. WebGiven the wide environmental and phylogenetic diversity of Bathyarchaeota, additional genomes are required to understand the metabolic capabilities of this understudied Recently, two more bathyarchaeotal fosmid clones were screened from estuarine mangrove sediments (Mengetal.2014). However, the ecological knowledge of Bathyarchaeota is limited in peatland ecosystems. As suggested by the classification of uncultured archaea based on nearly full-length 16S rRNA gene sequences, the bathyarchaeotal sequence boundary falls into the minimum sequence identity range of phylum level (74.9579.9%), and each subgroup generally falls into the median sequence identity range of family and order levels (91.6592.9% and 88.2590.1%, respectively) (Yarzaetal.2014). In a recent study exploring the stratified distribution of archaeal groups in a tropical water column, the analysis of archaeal 16S rRNA community distribution was combined with isoprenoid glycerol dialkyl glycerol tetraether lipid abundance information to reveal that glycerol dibiphytanyl glycerol tetraether lacking the cyclopentane rings [GDGT(0)] likely originated from the Bathyarchaeota-enriched layer in the water column (Bucklesetal.2013). (Kuboetal.2012), and the outgroup sequences of Crenarchaeota, YNPFFA group and Korarchaeota were added. The group was termed miscellaneous because of its occurrence in diverse habitats; it is not only abundant in marine sediments but is also widely distributed in terrestrial, freshwater, hot spring, hydrothermal, etc., environments (Kuboetal.2012). Vanwonterghem I, Evans PN, Parks DH et al. Thus, this systematic nomenclature based on clear monophyletic or phylogenetically stable subgroups not only facilitates further sequence assignment, but also provides useful information for understanding the evolutionary separation of specific lineages subjected to natural selection (Filloletal.2016). According to the meta-analysis of archaeal sequences available in the ARB SILVA database (Kuboetal.2012), Bathyarchaeota was further recognized as a group of global generalists dwelling in various environments, including marine sediments, hydrothermal vents, tidal flat and estuary sediments, hypersaline sediments, terrestrial subsurface, biomats, limnic water and sediments, underground aquifers, hot springs, soils, municipal wastewaters, animal digestive tract, etc. Lineage (full): cellular organisms; Archaea; TACK group. Combined with the aforementioned specific heterotrophic metabolic potentials of members within bathyarchaeotal subgroups and their occurrence in sediment layers of distinct biogeochemical properties (Lazaretal.2015), it was proposed that the acquisition of diverse physiological capacities by Bathyarchaeota is driven by adaptation to specific habitats rather than there being a common metabolic capacity. Peptidases targeting d-amino acids, which are highly enriched in the peptidoglycan of bacterial cell walls, are encoded as well, indicating that Bathyarchaeota may have acquired the capacity to degrade recalcitrant components of bacterial cell walls, i.e. On the other hand, the subgroups MCG-18 and MCG-19 were also named in Fillol et al.s research (Filloletal.2016). Specific lipids, exclusively synthesized by certain archaea, can serve as a supplementary biomarker for tracing the existence and abundance of targeted archaeal groups; their isotopic composition can be used to indicate specific carbon acquisition pathways (Schouten, Hopmans and Damste 2013). Subgroup-15 was recently found to be enriched in 13C-labeled DNA after a 3-month incubation experiment using sulfate-reducing sediments from Aarhus Bay, but was not present in the corresponding total DNA library or in a control incubation sample (i.e. Subgroup-6 persists in such suboxic, sulfide-depleted shallow sediment layers, while Subgroups-1, -5 and -8 preferentially occur in deeper, more reducing subsurface layers (Lazaretal.2015). Characteristics of the Bathyarchaeota community in The versatile metabolic properties of Bathyarchaeota, including acetogenesis, methane cycling, potential photosynthesis, and dissimilatory nitrite and sulfate reduction, etc., indicate that their ecological and phylogenetic characteristics are quite diverse, and given their basal phylogenetic position at the root of archaea, the evolutionary paths of those capabilities are also of great meaning for understanding the evolution of early life (Evansetal.2015; Heetal.2016; Lazaretal.2016; Zhangetal.2016). Following the four treatments, the viable bathyarchaeotal communities mainly comprised Subgroups-4 and -8, thus indicating that these two subgroups could tolerate the initial aerobic conditions (Gagenetal.2013). The available genomic evidence of various known and unknown methyltransferases harbored by BA1 and BA2 suggests the existence of a methylated compound utilization pathway, with the methyl group being ultimately reduced to CH3-H4MPT and integrated into the methyl-branch of the WoodLjungdahl pathway (Evansetal.2015). Obtaining direct physiological evidence for the generation or oxidization of methane by Bathyarchaeota in the future is also important. For us, phenotypical and genotypical information on subgroups whose existing patterns have only been sporadically reported still remains elusive and more explicit investigations are lacking. The use of MCG242dF resulted in an adequate coverage of almost all subgroups with 0/1 nucleotide mismatches, except for Subgroups-10 and -17, which showed low coverage efficiency with no nucleotide mismatches. These results have not only demonstrated multiple and important ecological functions of this archaeal phylum, but also paved the way for a detailed understanding of the evolution and metabolism of archaea as such. Hlne A, Mylne H, Christine D et al. We also highlighted the unique genomic features and potential adaptation strategies of estuarine archaea, pointing out major unknowns in the field and scope for future research. Stahl DA, Flesher B, Mansfield HR et al. More recently, Heetal. 1) (Heetal.2016; Lazaretal.2016). Furthermore, analysis of clone libraries retrieved after 13C-DNA amplification combined with matched terminal fragment length polymorphism peaks suggested that the heterotrophic bathyarchaeotal community possibly comprised Subgroups-6 and -8 (Seyler, McGuinness and Kerkhof 2014). The assignment of bathyarchaeotal subgroups was made based on either having been formerly defined or being monophyletic, using both distance and maximum-likelihood estimations (Kuboetal.2012). The novel Bathyarchaeota lineage possesses an incomplete methanogenesis pathway lacking the methyl co-enzyme M reductase complex and encodes a non-canonical acetogenic pathway potentially coupling methylotrophy to acetogenesis via the methyl branch of Wood-Ljundahl pathway. 2012 ). Four major heterotrophic pathways centralized on the acetyl-CoA generation are summarized below, reflecting the core metabolism of fermentation and acetogenesis (Fig. Buckles LK, Villanueva L, Weijers JWH et al. Community, Distribution, and Ecological Roles The subgroups MCG-18, -19 and -20 were firstly named in Lazar et al.s study, but only MCG-19 was represented in the phylogenetic tree (Lazaretal.2015). Members of the archaeal phylum Bathyarchaeota are widespread and abundant in the energy-deficient marine subsurface sediments. Kellermann MY, Wegener G, Elvert M et al. This approach revealed that the separation of subgroups according to saline and anoxic levels could explain 13% of the phylogenetic lineage variance. However, because of the high intragroup diversity and potential heterogeneous metabolic properties and adaptive strategies within the bathyarchaeotal subgroups, investigation into the subgroup distribution patterns at a fine-sorted phylotype level was recommended. Hallam SJ, Putnam N, Preston CM et al. Physiological incubation experiments with stable isotopic probing demonstrated that members of Bathyarchaeota are able to assimilate a wide variety of the tested 13C-organic compounds, including acetate, glycine, urea, simple biopolymers (extracted algal lipids) and complex biopolymers (ISOGRO) (Websteretal.2010; Seyler, McGuinness and Kerkhof 2014). Together with evidence of few phylogenetic changes throughout the incubation, it was suggested that the microbial community detected by stable isotopic probing could serve well in reflecting the metabolically active components. (C) The metabolic properties of 24 bathyarchaeotal genomes. The knowledge of their physiological and genomic properties, as well as their adaptive strategies in various eco-niches, is nonetheless still rudimentary. Summary. Bathyarchaeota, a recently proposed archaeal phylum, is globally distributed and highly abundant in anoxic sediments. This group of lipids has not been found in natural environments or microorganism enrichments dominated by methanotrophic archaea before (Rosseletal.2008; Kellermannetal.2012), nor have they been detected after re-analyzing lipid extracts from the above two studies using the same method in the study (Meadoretal.2015). (2015) presumed the syntrophy between Bathyarchaeota and sulfate-reducing bacteria (SRB) toward anaerobic oxidation of methane (AOM) (Evansetal.2015). Amend JP, McCollom TM, Hentscher M et al. n. Bathyarchaeota Gender: neuter To avoid the confusion, Subgroups-18 and -19 were named to be consistent with subgroups MCG-18 and MCG-19 as proposed in two previous reports (respectively Lazaretal.2015; Filloletal.2016), while Subgroup-20 was renamed to replace the subgroup MCG-19 in Fillol et al.s tree (Filloletal.2016). Combined with the large amount of carbon deposited in the subseafloor (ca 15 1021 g) (Fryetal.2008), the high abundance of MCG archaea in marine sediments (10100% of total archaeal abundance) (Parkesetal.2005; Biddleetal.2006; Fryetal.2008; Kuboetal.2012; Lloydetal.2013) and their heterotrophic properties on detrital proteins, acetate, aromatic compounds and/or other organic substrates (Biddleetal.2006; Websteretal.2010; Websteretal.2011; Lloydetal.2013; Naetal.2015), naturally led to the proposal that this group of archaea may play an important role in global carbon biogeochemical cycling (Kuboetal.2012; Lloydetal.2013; Filloletal.2016; Heetal.2016). Zhichao Zhou, Jie Pan, Fengping Wang, Ji-Dong Gu, Meng Li, Bathyarchaeota: globally distributed metabolic generalists in anoxic environments, FEMS Microbiology Reviews, Volume 42, Issue 5, September 2018, Pages 639655, https://doi.org/10.1093/femsre/fuy023. Genomic fragments of the fosmid clone 75G8 harbor a putative methyl-accepting chemotaxis protein- and 4-carboxymuconolactone decarboxylase-encoding genes, suggesting that this bathyarchaeotal member (Subgroup-8) is able to utilize aromatic compounds. Methanogenic archaea in peatlands However, Lokiarchaeota and most members of the Bathyarchaeota phylum lack the essential methane metabolizing elements, such as CoB or CoM synthase and methyl-CoM reductase, etc., though they use H4MPT as the C1-carrier, which is common in methanogens. Both Bathyarchaeota and the recently identified more basally branched Lokiarchaeota acquired the H4MPT-dependent WoodLjungdahl pathway and the hydrogen-dependent electron bifurcating system MvhADG-HdrABC, viewed as typical for the anaerobic and hydrogen-dependent archaeal lifestyle (Lazaretal.2016; Sousaetal.2016). Although the Pta-Ack pathway has been previously identified in the methanogenic genus Methanosarcina, it was shown that the encoding pta-ack gene pair might be derived from a horizontal transfer of genes of bacterial origin (Fournier and Gogarten 2008). Oxford University Press is a department of the University of Oxford. Bathyarchaeota, formerly known as the Miscellaneous Crenarchaeotal Group, is a phylum of global generalists that are widespread in anoxic sediments, which host relatively high abundance archaeal communities. Bathyarchaeota is of great interest to microbial ecologists for its wide distribution, high abundance, and diversity, as well as its potential ability to degrade detrital organic matter in aquatic environments and drive global elements cycling . the census of energy availability for redox reactions, is used, to some extent, to constrain and predict the distribution of functional groups of chemotrophic microorganisms (Amendetal.2011; LaRowe and Amend 2014). Furthermore, in contrast to the consistent vertical distribution of all archaeal lineages in freshwater sediments with almost no abundance changes, the total abundance of all Bathyarchaeota and the fraction of Subgroup-15 increase along with the depths of sediments, with significantly high abundance within the archaeal community (Liuetal.2014). In the White Oak River estuary, the abundance of Bathyarchaeota decreases with decreasing reductive redox conditions of the sediment (Lazaretal.2015). This review summarizes the recent findings pertaining to the ecological, physiological and genomic aspects of Bathyarchaeota, highlighting the vital role of this phylum in global carbon cycling. Genomic inferences from the two reconstructed bathyarchaeotal genomic bins from the coal-bed methane wells suggest that some Bathyarchaeota are methylotrophic methanogens feeding on a wide variety of methylated compounds, possessing an additional ability to ferment peptides, glucose and fatty acids (Evansetal.2015). The groups of B24 and B25 (Heetal.2016) were added into the tree representing Subgroups-21 and -22, respectively. Low collinear regions were found between bathyarchaeotal and reported archaeal genomic fragments, suggesting that the gene arrangement of Bathyarchaeota is distinct from that of sequenced archaea. For instance, a study into the stratification of the archaeal community from a shallow sediment in the Pearl River Estuary defined bathyarchaeotal subgroups from MCG-A to -F (Jiangetal.2011), including the NT-A3 group, which is predominantly isolated from the hydrate stability zone in the deep subsurface hydrate-bearing marine sediment core in the Nankai Trough (Reedetal.2002); meanwhile, an investigation of archaeal composition in ca 200 m deep sub-seafloor sediment cores at the offshore Peru Margin ODP sites 1228 and 1229 listed Bathyarchaeota subgroups PM-1 to -8 (Websteretal.2006).

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