On the other hand, because of the bidirectionality of these enzymes in methane metabolism (Boetiusetal.2000; Knittel and Boetius 2009), it is still possible that some members of Bathyarchaeota are involved in anaerobic methane oxidation. The metabolic properties are also considerably diverse based on genomic analysis (Fig. Four major heterotrophic pathways centralized on the acetyl-CoA generation are summarized below, reflecting the core metabolism of fermentation and acetogenesis (Fig. Among these are Subgroups-1 and -8 with high IndVal values in marine sediments, and Subgroups-5 and -11 with high IndVal values in fresh sediments (Filloletal.2016). Lineage (full): cellular organisms; Archaea; TACK group. 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. They are able to use a variety of substrates, including (i) detrital proteins, (ii) polymeric carbohydrates, (iii) fatty acids/aromatic compound, (iv) methane (or short alkane) and methylated compounds, and/or (v) potentially other organic matter to generate acetyl-CoA, subsequently using it to obtain energy or assimilate it in biosynthetic processes. 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. However, due to the great diversity of them, there is limited genomic information that accurately encompasses the metabolic potential of the entire archaeal phylum. It is well known that isoprenoid glycerol dialkyl glycerol tetraether lipids are specifically synthesized by archaea. Phylogenetic analysis of the Pta and Ack coding sequences in He et al.s study revealed that these genes form a monophyletic clade and are different from all other know sequences, indicating that they evolved independently of the currently known bacterial counterparts (Heetal.2016). Rossel PE, Lipp JS, Fredricks HF et al. (2016), it appears that these microbes rely on the acetyl-CoA synthetase (Acd) to generate acetate (Heetal.2016). This approach revealed that the separation of subgroups according to saline and anoxic levels could explain 13% of the phylogenetic lineage variance. In some flange subsamples, Bathyarchaeota were even more dominant than ANME; however, compared with the well-studied metabolism of ANME, the exact function of Bathyarchaeota in that ecological setting remains unknown. 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). 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). 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. In a recent study, Bathyarchaeota and ANME were shown to predominate on the flange of a hydrothermal chimney wall in the Soria Moria Vent field, where the local energy condition favors anaerobic methane oxidizers (Dahleetal.2015). Until now, This review is supported by the National Natural Science Foundation of China (grant numbers 31622002, 41506163, 31600093, 41525011, 91428308), the State Key R&D project of China (grant number 2016YFA0601102), the Key Project of Department of Education of Guangdong Province (No. Several sets of PCR primers and probes have been developed to detect and quantify Bathyarchaeota in natural community (Table 1). Acetyl-CoA might be involved in acetate generation in a fermentative pathway; however, genomic evidence suggests that Subgroup-1 cells might rely on both fermentative and respiratory metabolism (a simple respiratory metabolism based on a membrane-bound hydrogenase). Hence, the primer pair MCG242dF and MCG678R was developed based on a collection of bathyarchaeotal sequences of freshwater origin (Filloletal.2016). Moreover, the carbonyl branch of the WoodLjungdahl pathway might reduce CO2 into acetyl-CoA. The ability to use a wide range of substrates for energy conservation and biosynthesis, rather than a single reductive acetyl-CoA pathway, enhances the survival of Bathyarchaeota in energy-limited environments (Lazaretal.2016). Genes responsible for the dissimilatory nitrite reduction to ammonium (nirB and nrfD) were identified in Subgroups-1, -17 (formally Subgroup-7/17), -6 and -15, respectively, suggesting the potential existence of a respiratory pathway involving nitrite reduction (Lazaretal.2016). Third, only limited reports on the distribution patterns of bathyarchaeotal subgroups and the associated environmental factors are available. 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). Sousa FL, Neukirchen S, Allen JF 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). 1) (Heetal.2016; Lazaretal.2016). WebEtymology: Gr. The branching order of Subgroups-13 to -17 was unstable when analyzed by different tree-construction methods, and they were presented as multifurcated branches. For example, Bathyarchaeota dominates the archaeal community within Louisiana continental shelf (LCS) surface sediment, in both hypoxic and oxic covering water conditions in two distinct seasons (Devereuxetal.2015). A new phylum name for this group was proposed, i.e. (iii) The relatively small 13C signature of the archaeal intact polar lipids in comparison with the archaeal biomass suggests that the C isotopic fractionation during lipid biosynthesis is different from that of typical methylotrophic methanogens (Summons, Franzmann and Nichols 1998). The uptake and breakdown of polymeric hydrocarbons is facilitated by extracellular hydrolases; Bathyarchaeota also acquired the EmbdenMeyerhof Parnas/EntnerDoudoroff glycolysis and gluconeogenesis pathway for the core hydrocarbon utilization metabolism. Bathyarchaeota was the most dominant archaeal taxa with 108 nodes and 501 edges in the network. Based on the physiological and genomic evidence, acetyl-coenzyme A-centralized heterotrophic pathways of energy conservation have been proposed to function in Bathyarchaeota; these microbes are able to anaerobically utilize (i) detrital proteins, (ii) polymeric carbohydrates, (iii) fatty acids/aromatic compounds, (iv) methane (or short chain alkane) and methylated compounds, and/or (v) potentially other organic matter. The first two separation nodes representing the hypersaline, saline and fresh environments accounted for 9.1% of the total phylogenetic lineage variance. Individual metagenome assemblies Institute for Advanced Study, Shenzhen University, Shenzhen 518060, People's Republic of China, Laboratory of Environmental Microbiology and Toxicology, School of Biological Sciences, The University of Hong Kong, Pokfulam Road, Hong Kong SAR, People's Republic of China. This primer pair shows good specificity toward Bathyarchaeota; it allowed amplification of 10100 times more bathyarchaeotal 16S rRNA gene sequences from the sediment samples from the South China Sea, and the Atlantic and Antarctic Oceans than the MCG242dF/MCG678R primers (Yuetal.2017). Hlne A, Mylne H, Christine D et al. Two highly abundant MCR variants were detected in Ca. Laso-Prez R, Wegener G, Knittel K et al. Membrane lipids are an informative indicator of the distribution and activity of living microbial cells, independently of their culturing (Sturtetal.2004; Jacquemetetal.2009; Lipp, Liu and Hinrichs 2009). Recently, two bathyarchaeotal genome bins (BA1 and BA2) were recovered from the formation waters of coal-bed methane wells within the Surat Basin (Evansetal.2015). Given the substrate specificity of this MCR type in utilizing butane instead of methane, and amino acid divergence of this MCR type from its methane metabolizing related counterparts, it is possible that the MCR clusters in some members of Bathyarchaeota are responsible for butane oxidation instead of methane metabolism (Laso-Prezetal.2016). Stahl DA, Flesher B, Mansfield HR et al. Here we reported the abundance of Bathyarchaeota members across different ecosystems and their correlation with environmental factors by constructing 16S Given the high phylogenetic diversity within the 25 subgroups of Bathyarchaeota, many efforts have been made to understand the key factors that control their distribution and evolution. This method has been used to target the bathyarchaeotal 16S rRNA gene with specific probes, providing information on the active bathyarchaeotal community without culturing (Table 1). This could be explained by the versatile pathways of organic matter assimilation present in the majority of Bathyarchaeota, reflected by inferences from genomic data. However, the global methane cycle should be reconsidered since the previously unrecognized methane metabolic capacity appears to be present within such a widespread and abundant phylum. 4), although these might not necessarily exist in all bathyarchaeotal subgroups (Fig. 3B). BA1 also lacks other genes for energy-conserving complexes, including F420H2 dehydrogenase, energy-converting hydrogenases A and B, Rhodobacter nitrogen fixation complex and V/A-type ATP synthase. These findings expand the metabolic potential of archaea and argue for a revision of the role of archaea in the carbon cycle in marine sediments (Heetal.2016). 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). Consequently, CO2 appears to be the only electron acceptor mediating AOM, like in a reverse acetoclastic methanogenesis (Hallametal.2004; Wangetal.2014). A segregated distribution of bathyarchaeotal subgroups was also observed in the water column and sediments in freshwater karstic lakes (Filloletal.2015). Jacquemet A, Barbeau J, Lemiegre L et al. 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. S. butanivorans forms a distinct cluster with those of Bathyarchaeota origin, separately from other methanogens and methanotrophs (Laso-Prezetal.2016). The wide phylogenetic coverage increases the difficulty of inferring the general metabolic properties across whole lineages. Furthermore, a principal coordinate analysis also clearly separates the bathyarchaeotal community into freshwater and saline sediment groups. Members of Bathyarchaeota are able to use CO2 and H2 from natural sources and fermentation products to fuel acetogenesis (Heetal.2016; Martinetal.2016). 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. 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). 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). Markers for individual pathway/function were scanned against genomes using the HMM and KEGG databases (Anantharamanetal.2016; Kanehisa, Sato and Morishima 2016; Spang, Caceres and Ettema 2017). It also contains typical methane metabolism genes (hdrABC and mvhADG) but lacks hdrE, similar to Methanomassiliicoccales genomes (Evansetal.2015). Evans PN, Parks DH, Chadwick GL et al. Within Bathyarchaeota, the sequences were classified into six subclades according to . 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). The archaeal phylum Bathyarchaeota, which is composed of a large number of diverse lineages, is widespread and abundant in marine sediments. The distinct bathyarchaeotal subgroups diverged to adapt to marine and freshwater environments. Future experiments investigating substrate specificity of these proteins and analyses of the intermediate metabolites will help establish their actual functions. (ii) Similar 13C signatures of the archaeal biomass and total organic carbon suggest that the organic matter assimilation contributes to the bulk of the archaeal biomass; the relatively small 13C signature of the archaeal biomass in comparison with the dissolved inorganic carbon suggests that only a small amount of archaeal biomass is derived from autotrophic CO2 fixation (Biddleetal.2006). Heetal. WebBathyarchaeota dominated the archaeal interaction network with 82% nodes, 96% edges, and 71% keystone species. Some of these Crenarchaeota were able to assimilate all 13C-organic compounds tested, including acetate, glycine, urea, simple biopolymers (extracted algal lipids) and complex biopolymers (ISOGRO), while others were only detected in specific substrates (acetate or urea). Background Bathyarchaeota, a newly proposed archaeal phylum, is considered as an important driver of the global carbon cycle. It has been suggested that Bathyarchaeota is one of the cosmopolitan groups frequently detected in the freshwater and marine sediments (68% of all sediments analyzed), accounting for a large proportion of the sediment microbial communities (average 36 22%) (Filloletal.2016). Given that they are abundant, globally distributed and phylogenetically diverse, continued exploration of new potential bathyarchaeotal subgroups is encouraged. In addition, the catalyzed reporter deposition-fluorescent in situ hybridization (CARD-FISH) studies for the detection and quantification of bathyarchaeotal cells suggest that they are abundant in the center and marine invertebrate-inhabited layers in the Haakon Mosby Mud Volcano, and in the marine subsurface sediments in the Equatorial ODP site 1125 and Peru Basin ODP site 1231 (Kuboetal.2012). The 13C-depleted nature of butanetriol dibiphytanyl glycerol tetraethers found in the study implied that members of Bathyarchaeota might be autotrophs or fueled by 13C-depleted organic substrates (Meadoretal.2015). The members of Bathyarchaeota were positively and strongly correlated especially with the acetoclastic Methanosaeta; however, the second most abundant archaeal group, MG-I (subordinate to Thaumarchaeota) is negatively correlated with other groups, probably indicating segregation corresponding to two distinct lifestyles in this case (Liuetal.2014). 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. Bathyarchaeota, a recently proposed archaeal phylum, is globally distributed and highly abundant in anoxic sediments. The first comprehensive phylogenetic tree of Bathyarchaeota was constructed in 2012 (Kuboetal.2012); it was based on 4720 bathyarchaeotal sequences from the SILVA database (SSU Ref NR106 and SSU Parc106). Callac N, Rommevaux-Jestin C, Rouxel O et al. Inagaki F, Nunoura T, Nakagawa S et al. bathys, meaning deep as it locates deep branching with Thaumarchaeota and Aigarchaeaota, and frequently detected in the deep subsurface sediments; N.L. No methane metabolism genes were recovered from bathyarchaeotal genomic bins or any contigs from the WOR estuarine sediments, in contrast to an earlier study (Evansetal.2015). WebArchaea are tiny, simple organisms. S. butanivorans protein extracts; they are probably responsible for the initial step of butane activation to generate butyl-CoM. Microbial communities of deep marine subsurface sediments: molecular and cultivation surveys, Methanogenic archaea: ecologically relevant differences in energy conservation, Methylotrophic methanogenesis discovered in the archaeal phylum, Methanotrophic archaea possessing diverging methane-oxidizing and electron-transporting pathways, Prokaryotic community composition and biogeochemical processes in deep subseafloor sediments from the Peru Margin, Prokaryotic functional diversity in different biogeochemical depth zones in tidal sediments of ?the Severn Estuary, UK, revealed by stable-isotope probing, Enrichment and cultivation of prokaryotes associated with the sulphate-methane transition zone of diffusion-controlled sediments of Aarhus Bay, Denmark, under heterotrophic conditions, The physiology and habitat of the last universal common ancestor, Distribution of Bathyarchaeota communities across different terrestrial settings and their potential ecological functions, Uniting the classification of cultured and uncultured bacteria and archaea using 16S rRNA gene sequences, A large-scale evaluation of algorithms to calculate average nucleotide identity, High occurrence of Bathyarchaeota (MCG) in the deep-sea sediments of South China Sea quantified using newly designed PCR primers, Growth of sedimentary Bathyarchaeota on lignin as an energy source, Genomic and transcriptomic evidence for carbohydrate consumption among microorganisms in a cold seep brine pool, This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (, Illuminating the Oral Microbiome and its Host Interactions: Animal models of disease, Engineering lanthipeptides by introducing a large variety of RiPP modifications to obtain new-to-nature bioactive peptides, Meat fermentation at a crossroads: where the age-old interplay of human, animal, and microbial diversity and contemporary markets meet, Incorporation, fate, and turnover of free fatty acids in cyanobacteria, Ruminococcus gnavus: friend or foe for human health, About the Federation of European Microbiological Societies, GLOBAL DISTRIBUTION AND HIGH DIVERSITY OF BATHYARCHAEOTA, DISTRIBUTION PATTERN AND MOLECULAR DETECTION, PHYSIOLOGICAL AND GENOMIC CHARACTERIZATION, ECOLOGICAL FUNCTIONS AND EVOLUTION OF BATHYARCHAEOTA, https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model, Receive exclusive offers and updates from Oxford Academic, Copyright 2023 Federation of European Microbiological Societies. (B) The dendrogram and genome similarity heatmap based on pairwise OrthoANIu values of 24 bathyarchaeotal genomes (Yoonetal.2017). 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 . Logares R, Brate J, Bertilsson S et al. Due to their prevalence in the microbial community, we also performed phylogenetic analysis to understand the closeness of our Bathyarchaeota OTUs with The major bathyarchaeotal community comprises Subgroups-1, -8, -12 and -15, and is relatively stable during the hypoxic/oxic change, thus being independent of the sedimentary chemistry change, such as manganese and iron redox cycling during different seasons (Devereuxetal.2015). Materials and methods 2.1. It is one of the predominant groups in the marine subsurface archaeal community (Fryetal.2008; Teske and Srensen 2008; Lloydetal.2013). Bathy-15 (36.4% of all archaea), Recently, another meta-analysis using newly acquired global sediment bathyarchaeotal sequences resulted in the addition of two more subgroups, Subgroups-18 and -19, with high bootstrap supporting values (96% and 86%, respectively) (Filloletal.2016). The members of the Bathyarchaeota are the most abundant archaeal components of the transitional zone between the freshwater and saltwater benthic sediments along the Pearl River, with a central position within the co-occurrence network among other lineages (Liuetal.2014). 2017KZDXM071), and the Science and Technology Innovation Committee of Shenzhen (Grant No. the potential AOM metabolism of Bathyarchaeota in the flange of the hydrothermal vent would be consistent with the aforementioned genomic inferences (Evansetal.2015). The analysis of the stable isotopic-probed microcosms from Cheesequake salt marsh sediment revealed that all Crenarchaeota groups, which still include Bathyarchaeota and Thaumarchaeota (formerly Crenarchaeota MG 1.a) and other Crenarchaeota groups, are heterotrophic and do not incorporate 13C-bicarbonate (Seyler, McGuinness and Kerkhof 2014). 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. The currently available bathyarchaeotal genomes shared 63.5% similarity on average, indicating a wide phylogenetic diversity at the genome scale (Fig. In this study, the abundance and The current genomic and physiological information of these subgroups also suggests their potential ecological strategies and functions in specific habitats, further highlighting their important roles in global biogeochemical cycling (Xiangetal.2017). 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. (i) The 13C signature of the archaeal biomass suggests that only a small fraction of local archaea in SMTZ utilize methane, which might be explained by the contribution of Bathyarchaeota in the biomass; until now, only one line of evidence points to the acquisition of methane metabolism by Bathyarchaeota (Lloydetal.2013; Evansetal.2015; Lazaretal.2015; Heetal.2016). CARD-FISH can be utilized for the detection, identification and enumeration of microorganisms in various environments, independently of culturing (Kubota 2013). The metagenomic binning of WOR estuarine sediment DNA led to the reconstruction of draft genomes of four widespread Bathyarchaeota, with the genome completeness in the range of 4898% (Lazaretal.2016). (2016) reconstructed six nearly complete bathyarchaeotal genomes (Subgroups-13, -15, -16, -18 and -19) from the Guaymas Basin subsurface sediment. Further membrane lipid characterization of enriched or pure bathyarchaeotal cultures will help to validate this discovery. Metagenomic evidence of sulfate reductase-encoding genes in the upper region of SMTZ of the OPD site 1229 provides more hints to the potential synergistic metabolism of AOM coupled with sulfate reduction (Biddleetal.2008). Bathyarchaeota, reflecting its phylogenetic position as deeply branching with Aigarchaeota and Thaumarchaeota, and its prevalence in subsurface sediments (Mengetal.2014). is bathyarchaeota multicellular. Three fosmid clones harboring bathyarchaeotal genomic fragments were screened from the South China Sea sediments (05 cm depth) (Lietal.2012). Fillol M, Snchez-Melsi A, Gich F et al. Reconsideration of the potential methane-oxidizing contribution of Bathyarchaeota would refine the congruency between the predicted and observed microbial communities, i.e. It is known that a methane microbiome can be established in methane seeps sites; however, they are still poorly characterised. Considering the bathyarchaeotal community structure, depth is the first variable responsible for the high degree of absolute subgroup separation, followed by sulfide concentration (reflecting the redox conditions), which is responsible for a low degree of subgroup separation (Lazaretal.2015). The gene for cytoplasmic flavin adenine dinucleotide-containing dehydrogenase (glcD) co-located with hdrD, indicating that BA1 uses lactate to reduce heterodisulfide in methanogenesis. A meta-analysis of the distribution of sediment archaeal communities towards environmental eco-factors (7098 archaeal operational taxonomic units from 207 sediment sites worldwide) was performed and a multivariate regression tree was constructed to depict the relationship between archaeal lineages and the environmental origin matrix (Filloletal.2016). The picked genomes are of high completeness (>70%) and good quality (excluding genomes with numerous long breaking parts with N). Although the accumulated information paves the way for further clarification of the adaptation of different lineages to various environments, systematic understanding of the distribution pattern of bathyarchaeotal subgroups and influential factors is still needed. (2016) demonstrated that half of the bathyarchaeotal genomes encode a set of phosphate acetyltransferase (Pta) and acetate kinase (Ack) for acetate production or assimilation, usually observed in bacteria. Members of the archaeal phylum Bathyarchaeota are widespread and abundant in the energy-deficient marine subsurface sediments. Several pre-/non-enriched sediment cultures afforded preliminary evidence for the trophic properties and metabolic capacities of Bathyarchaeota. 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. 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). Tree building intermediate files are publicly available (https://github.com/ChaoLab/Bathy16Stree). Subgroup-5b was further split into 5b and 5bb, as additional sequences were added. Recently, two more bathyarchaeotal fosmid clones were screened from estuarine mangrove sediments (Mengetal.2014). the classification proposed by Zhou et al. 2). n. Bathyarchaeota Gender: neuter Bathyarchaeota are believed to have roles in the carbon cycle in marine systems. A model based on the thermodynamic considerations of chemicals and temperatures may be used to offer a framework linking the distribution of microbial groups and energy landscapes (Amendetal.2011; LaRowe and Amend 2014; Dahleetal.2015). Recent genomic evidence suggests that Bathyarchaeota might potentially be involved in methane metabolism, a property that had only been confirmed to date in the Euryarchaeota domain (Evansetal.2015; Lloyd 2015). A group called Peat MCG (pMCG) (Xiangetal.2017) was also listed on the tree; however, because there was only one represented sequence after dereplication at 90% similarity of all bathyarchaeotal 16S rRNA gene sequences, we did not list pMCG as a separate subgroup in this tree (Fig. ( 2012) conducted a comprehensive analysis of the biogeographical distribution of Bathyarchaeota and found that it was the dominant archaeal population in anoxic, low-activity subsurface sediments. 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). Biddle JF, Fitz-Gibbon S, Schuster SC et al. A pair of primers (Bathy-442F/Bathy-644R) was recently designed to target Subgroups-15 and -17; the in silico primer testing indicates that Bathy-442F can also adequately cover Subgroups-2, -4, -9 and -14, with Bathy-644R covering nearly all subgroups, except for Subgroups-6 and -11 (Yuetal.2017).
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