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Data may be preliminary. aiiygains(atln ta. 04 ebl ta. 04 Mu˜noz-Mart´ın2005). 2014; al., al., et et up Herbold reach 2014; fumaroles al., Island et Deception ascending (Bartolini volcanoes, hot gradients continental with salinity and Antarctic contact glaciers mass Unlike of water 100 combination 2019). and to This cryosphere al., the 2004). et between al., (Geyer interaction et emissions magmas the (Somoza continuous by sulfide basin, with produced hydrogen marginal fumaroles is and Strait active fumaroles dioxide Bransfield and the carbon glaciers of mostly Deception center permanent gases, pathways. spreading of of the metabolic ecosystems con- at the and contrasting Islands volcano have mechanisms Shetland harbors Island together adaptive South which that the Deception microbial in sources The of located energy variety unique is 2019). geochemical wide Island a al., and persists a providing et temperatures selecting life as extreme (Merino for how well of potential itself as Understanding combination life function, unique of biologists. a systems history tains fascinated living evolutionary how long the into has into insight environments window provides extreme extremes in environmental life at of study The Introduction and Nanoarchaeota Archaea. to capabilities and survival related and Bacteria metabolic those of the lineages especially of extremophilic understanding groups, were different diverse enhance taxonomic we results of less genomes, underrepresented Our showed of of lineages. communities reconstruction archaeal lifestyles glacier ammonia-oxidizing the novel thermophilic Through contrast, putative pathways. In clarify carbon strategies to conditions. and survival able heterotrophic and temperature mainly metabolic and diverse comprising redox more potentials, fluctuating possess metabolic sediments. to fumaroles as microbial glacier well oC responding how as and <80 strategies, of from fumarole explore molecular capable those hyperthermophilic in to while specific gradients pathways, harbor genomes fumarole autotrophic temperature enzymes oC metagenome-assembled and and 98 metabolic reductive and a little salinity key from been metagenomics However, communities geochemical, of microbial have extreme shotgun inhibition that diversity. observed active by to We distances used and shaped due and we short structure example is study, oC) very community diversity for this (<0 over microbial functional functioning, In ecosystem glaciers geochemistry effect affect to permanent and pathways. gradients shown salinity as or these Island been how such temperature, Deception have regarding ecosystems in known and geochemical pathways. contrasting gradients is Island, and metabolic in Steep Deception temperature and resulting oC). for remarkable mechanisms sea, 100 reported have adaptive the to landscape, microbial by (up of icy variety the fumaroles flooded wide of stratovolcano a rest a for the is select to could contrast that in gradients Antarctica, in volcanoes Active Abstract 2020 11, August 3 2 1 Bohannan Brendan BENDIA AMANDA Antarctica Deception volcano, on Island gradients temperature extreme microbial across of communities strategies survival and potential Metabolic nvriyo Oregon of University USP Paulo Sao of University o n aedrc aieiflec,cetn eakbecmiaino hra,gohmcland geochemical thermal, of combination remarkable a creating influence, marine direct have and C 3 n iinPellizari Vivian and , 1 enr acmnoLemos Nascimento Leandro , 1 1 2 ua Mendes Lucas , 1 aiaSignori Camila , 1 , Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. n ftmeaue n drse eta usin eadn h ucinlaatblt fextremophiles of adaptability functional the regarding gradi- steep questions a regions. central inhabiting polar addresses communities adds in and microbial study temperature, of This processes of temperature. vent ecological ent increasing hydrothermal the with deep-sea regarding for decrease these information interactions reported (ii) important was gradients; microbe-microbe strategies that temperature (iii) potential metabolic extreme and metabolic and and communities; survival of salinity diversity, redundancy geochemical, community the of for follow combination reported communities the been hypo- by has and we influenced metageno- what lifestyles Here, also shotgun putative profiles. to are functional similar microbes‘ performed community (i) the we with that reveal information this, thesize to genomic two For genomes the from reconstructed combining Antarctica. sediments and capabilities, glacier volcano, survival diversity and Island functional fumarole unveil Deception microbial in to profile a the mics co-occurrence functional in on microbial of place the sites 2018). analysis take assessed al., geothermal The we that ha- et study, rearrangements 2018). Mandakovic current others ecological the 2010; al., 2014), In the al., et al., et of Mandakovic et (Freilich interactions nature environments 2016; Sharp the microbe-microbe contrasting 2019; al., examining facing of al., et community for et frequency (Lin useful Merino the trend is 2013; in patterns opposite al., interac- the decrease et microbe-microbe observed a (Cole on ve patterns reported environments co-occurrence have extreme from of studies inferred impact some the Although in about thriving surroundings. tions. controversy of cold hydrothermal is capable the deep-sea there specialists tolerate 2019; Furthermore, (hyper)thermophilic and also favors other Dick, can volcanoes that (and that (e.g. pressure polar pressure but selective vents in of temperatures strong differences hydrothermal typical high a regarding deep-sea as range that, act in temperature suggests can which found vents wide cold 2001), the those surrounding al., effects), to et the Takai environmental 2018) with 2006; hydrothermal heat al., al., deep-sea of et et the Nakagawa contact of Signori, the exception temperature by (Bendia, the extent 0-4 the created members with where are what around volcanoes, ecosystems, temperatures to polar (temperature thermal contrasting in seawater unclear nonpolar the as on still which metabolic broad focused in and is too and have mechanisms vents, not it adaptive studies Price is their previous 2017), range 2017; as these al., such temperature al., community, of et microbial majority et a Ward taxonomic The Antranikian of pathways. microbial 2014; processes (e.g. of functional al., ecosystems driver the et affects primary hydrothermal Sharp a and 2017; is geothermal Giovannelli, temperature different that and in demonstrated Island. adaptive diversity have These Deception their 2018). studies on and al., found several structure gradients et Although microbial environmental Signori, between particular (Bendia, linkages niche the the temperature over strongly address by strategies such is not shaped geochemicals, metabolic Island did volcanic mainly however, Deception and is studies, salinity on diversity previous pH, archaeal structure (temperature, community while parameters sulfate), bacterial environmental as the of variety that a Archaea, combination reported by hyperthermophilic unique driven we 2019). (including a Detector Also vents al., select Life hydrothermal 2018). to et shallow the able al., (Lezcano and were applied Caliente deep Deception study as Cerro in on such second gradients found from steep groups a communities the taxonomic and of that through of 2018), showed functions diversity study al., taxonomic general previous determining et describe Our on Signori, to focused (Bendia, (LDChip) group, micro- sequencing Chip on our gradients gene by temperature rRNA analyzed. limited performed Deception were of 16S were first, sediments effect studies The cold the communities. previous understanding or on bial fumaroles These focused only 2019). have since studies al., extent previous et Two and (Centurion depth Bay sampling resistome Whalers to the using characterize respect from fumaroles to with sediments Deception metagenomics was cold shotgun on study diversity and in recent microbial 2011) profiles (Bendia, more al., described Mu˜noz gradients et a studies 2013; temperature 1967), al., molecular steep (Amen´abar or al., et the Previous DGGE hot among et 2018). from isolates Stanley al., isolates thermophilic et 1997; and bacterial Araujo, al., psychrophilic obtaining et both on Llarch recover primarily to 2011; able focused al., Carri´on Deception et (e.g. on ecosystems out cold carr yoitu spp. Pyrodictium ,gohra ytm n hs yia rmplreoytm Bni,Sgoi et Signori, (Bendia, ecosystems polar from typical those and systems geothermal ), o ) ned h eeto omnte rmfmrlshv similar have fumaroles from communities Deception the Indeed, C). 2 Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. ae(ak ta. 04 n otae( eeomn oeTa) sn h packages the using Team), Core Development 2012), (R soft- (STAMP) software al., Profiles R et Metagenomic and Wilke of 2014) 2016; Analysis al., al., Statistical et et using (Parks Keegan analyzed ware 2000; statistically Goto, were & MG-RAST MG- 1x10 by Kanehisa subsystem KEGG, to parameters: 2008; and through following uploaded al., database) generated the then protein et were with (non-redundant and (Aziz M5NR reads >30 subsystem, MG-RAST of SEED phred in profiles the available with taxonomic using classifications and 2011) Functional hit Fass, best 2016). & and al., (Joshi et Sickle (Keegan using RAST trimmed quality total reads Al, were Mg, metagenomic Reads Ca, of K, inference Na, Si, functional P, and pH, Taxonomic sediments carbon, humidity, organic sulfate. for conductivity, matter, and electrical parameters organic ammonia, granulometry, Zn), physical-chemical nitrate, included and nitrogen, which the Mn, Fe, 2018), used Cu, al., we (B, et micronutrients Signori, data, (Bendia, environmental by and measured biological correlate To paired-end bp 2x100 using the platform Universidade parameters following (LaCTAD), 2000 Physical-chemical Hiseq Coulter) Vida” Illumina da (Beckman the Ciencias on kit em (UNICAMP), system. beads Desempenho State at XP Campinas Alto conducted de de AMPure were Estadual Tecnologias sequencing using the de metagenomic shotgun purified following Central DNA and “Laboratorio sufficient USA), then constructions obtain same NJ, Library was to Piscataway, the instructions. DNA pooled manufacturer’s Healthcare, were using used sample (GE (MDA) several we sequencing. per after kit amplification reactions 2018), even for displacement amplification amplification mass al., Three multiple DNA DNA et low instructions. to V2 manufacturer’s to Signori, subjected GenomiPhi Due (Bendia, was illustra metagenomics. the by DNA the for extracted performed template efforts, a study as concentration the extractions water 2014. DNA with the April and in in results dataset Brazil depth our Paulo, cm Sao compare 50 intertidal of at To the University sequencing (submerged the in metagenomic zone at were arrival and subtidal fumaroles until the Extraction All degC in DNA apart. -20 was at km which 98 stored 10 FB, measuring were cm approximately from fumarole, Samples were few B hottest transects column). glacier, point the FB for the was and except from FBA WB zone, point the collected The and were were m, samples B 1d). 15 and and C ( A 1c surface point Points 0 sediment (Figure while between samples. the edge sediment fumaroles, temperatures glacier’s 18 in with the totaling collected below 1b), point, samples collected and each as 1a for defined (Figure sediment performed W) were surface triplicates 60deg33’27.3” collected and We S, (62deg58’45.1” 98 Maximiano. WB and Almirante and NPo. W) vessel 36.4” polar 2014), the January ( to & are from samples 2013 Zhang (December gases support 2004; Expedition sulfide Antarctic logistical al., Brazilian hydrogen et with XXXII (Somoza and the during sulfate dioxide performed and was Carbon sulfite Sampling as submerged 2004). such and in in al., products WB mostly km et to in 1993). distributed oxidized Somoza 9 40-60degC are Millero, are 1995; Bay, and from they Foster al., fumaroles varying and et by Port and temperatures emitted (WB), (Rey 2019), called with Bay FB al., zones), caldera Whalers in (intertidal et (FB), flooded 80-100degC Bay regions Geyer Fumarole a submerged 2007; at to partially al., mainly and rise ago found et years are (Fermani giving Fumaroles 10,000 Cove island approximately 1975). Pendulum the al., occurring et eruption of (Baker past part diameter A central Peninsula. the Antarctic collapsed the near Strait, Bransfield (62 Island Deception strategy sampling and site Study methods and Materials o .A ahst,w bandsmlsfo he ieetpit ihntetmeauegradient, temperature the within points different three from samples obtained we site, each At C. ca. m nfmrlsadgair tgohral ciestsi B(2e5’27 ,60deg42’ S, (62deg58’02.7” FB in sites active geothermally at glaciers and fumaroles in cm) 5 ° 8 ,6dg9 )i ope taoocn oae nteSuhSeln Islands, Shetland South the in located stratovolcano complex a is W) 60deg39’ S, 58’ ca. 0c) itne ewe uaoe n lcesa ahst eeapproximately were site each at glaciers and fumaroles between Distances cm). 20 -5 -au,mnmm5 painet n 0 dniy aagenerated Data identity. 60% and alignment, bp 50 minimum e-value, 3 vegan (Oksanen, o at C Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. oeteohlso soitdt ufradntoe eaoim.Antto falpeitdOF in ORFs predicted all of Annotation metabolisms. related nitrogen groups and selecting We sulfur preferably taxonomy, 2017). to and al., high-quality associated et or or (Bowers medium by extremophiles their metagenome suggested on to contamination) standards based MAGs <10% quality 11 genome complete, selected to (<50% according draft (>50% (MAG), low-quality draft medium-quality assembled-genome or contamination), <5% contamination) complete, <10% (>90% were draft complete, high-quality Bins a (MAGs) as defined genomes sets. were metagenome-assembled Bins gene of marker 2020). annotation lineage-specific al., functional et of and (Chaumeil representation Taxonomic GTDB-Tk the 1.0.7 using on phylogeny v. genome based CheckM on through is checked based which quality classified were 2015), taxonomically refined approaches manually al., co-assembly then et and used were assembly (Parks ‘anvi-summarize’. We the Bins using using by estimated parameters. generated bins. performed were Bins genome default was contamination identify with and binning and completeness program Genome and data ‘anvi-merge’ ‘anvi-refine’, kbp. merged the using 4 the through using visualize of 2013) KEGG profiled to length al., were against ‘anvi-interactive’ contig files et annotated minimum BAM (Altschul (Alneberg taxonomically a Individual v2.10.0+ CONCOCT and with the blastp 2016). of ‘anvi-profile’ functionally al., using release program were et 2014 2015) the sequences (Kanehisa December al., protein (genus_prokaryotes) the et GhostKOALA Predicted against and (Galperin with them bacterial 1990). database ‘anvi-run-ncbi-cogs’ for Single-copy program al., (COGs) searching The by (ORFs). Groups et ‘anvi-gen-contigs-database’. 2011). functions frames Orthologous al., the with reading of et genes using (Finn Clusters open annotate 3.1b2 generated predict al. to v. to et was used HMMER Eren was used using database by identified was parameters contig were described 2010) default genes workflow with A al., archaeal the bowtie2 et using following (Hyatt co-assembly 2012). 5 the Prodigal to Salzberg, v. mapped anvi’o & smaller were using contigs metagenome (Langmead Microbiomes discarding binned each 2015), were for & al., contigs Reads et Genomes Then (2015). (Li Microbial 1.0.2. bp. Integrated MaxBin 1000 v. the MEGAHIT through than using using performed co-assembled were was annotated reads (- binning Furthermore, were 2009). 2012) genomic Contigs al., al., then et et and (Markowitz 2016). (Peng system 1000) IDBA-ud al., (IMG/M) -min_contig using et 4, assembled (Wu -tep were 2.0 92, reads metagenomic -maxk First, eighteen the 50, volcano. of mink binning Island genomic Deception and assembly from metagenomic datasets for strategies different two used We reconstruction the genome with and modularity, calculated assembly edges, were Metagenomic and 2009). measurements nodes al., property of et and degree number (Bastian cumulative visualization the Gephi and Network software as diameter, length, genus such 2003). analysis path measures, the average (Newman, The ( analysis. connectivity, of at distribution node significant for nodes. set average taxa between communities, highly a used represent of correlation on and was number network negative based the level -0.9) or reconstructed was genus < positive complexity the significantly network the or in represent of to 0.9 nodes edges affiliated > the The whereas hits (SparCC level, taxa. of strong ‘SparCC’ microbial frequency only module between of considered Python correlations we the table network, using A each performed For 2012). were analyses Alm, network co-occurrence co-occurrence & used non-random (Friedman we site, this, sampling For each at interactions analysis. community correlations of complexity environmental Spearman the the investigate To addition, and profiles In functional relative and to samples. taxonomic normalized between across were parameters. relationships Values composition determine to software. taxonomic STAMP performed of 80 were in to comparison visualized up then for and fumaroles abundance subsystems glaciers, were SEED comparisons temperatures: of environmental Statistical functions to Newcombe-Wilson. according of samples method the o grouping the by using performed calculated were intervals confidence and 2007) n uaoea 98 at fumarole and C ggplot Wchm 01.The 2011). (Wickham, o .Picplcmoetaayi PA riainwspromdb sn ee 3 level using by performed was ordination (PCA) analysis component Principal C. p auswr acltduigFse’ xc w-ie etadthe and test two-sided exact Fisher’s using calculated were values 4 p 0.01) < Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. ftecmuiyicesdwt h eprtr.W lontdta omnte fFmrl a were complexity Bay the Fumarole (98 of general, FBA communities In The that MG-RAST. noted Bay. in also We annotated Whalers reads than temperature. microbial the metagenome between complex with correlations on more increased SparCC based microbial community calculated the level the we within genus of this, interactions For the of 2c). complexity at the (Figure taxa treatment explore each to analysis in network communities similar co-occurrence 1). showed used system Figure reads then (Supplementary IMG/M We using annotation the level through reads class contigs to of the compared annotation at when taxonomic classified patterns and The not Verrucumicrobi- MG-RAST. within Delta- were and in Acidobacteria, assignments Beta-, affiliated annotation within Thaumarchaeota by classes class followed Verrucomicrobia. abundant Al- main within the Proteobacteria, and was aea The within Solibacteres Gamma- only classes 2b). whereas were represented (Figure 2a). Sphingobacteria, Acidobacteria Epsilonbacteria most (Figure and and the Flavobacteria respectively) Verrucomicrobia were Cytophagia, phaproteobacteria 1-1.6%, were <80 Bay). phylum and in Fumarole Bacteroidetes (1.2-3.1% in found the glaciers predominantly 0.2-0.3% <80 was and in fumaroles Thaumarchaeota Bay detected distributed in Whalers example, uniquely 0.7-2% were For for phyla of (0.8-1% dominant abundance temperature. less relative Firmicutes), to and a according Proteobacteria, with Bacteroidetes, samples, (e.g. other glaciers the in dominant o less were 2a). Archaea (Figure were Bacteria Thermotogales) within (order classes Thermotogae and dominant Aquificales) the most (order 98 Aquificae the the Flavobacteria, Fusobacteria, represented in ria, (>0.1%) Methanomicrobia classes and archaeal also Methanopyri, abundant were Methanobacteria, (0.3-1.0%) 98 Archaeoglobi, Thermotogae the Methanococci, in and 98 Firmi- proportions the (0.3-4.6%), minor (0.1-0.4%). Aquificae from in Korarchaeota detected samples (0.6-15.3%), and in Bacteroidetes (2.5-7.5%) dominant Euryarchaeota (3.1-22.4%), by were 98 cutes followed Archaea the (23.8-79.3%), in Crenarchaeota glaciers. composition as and taxonomic fumaroles the other that with observed o comparison we in volcano reads, distinct Island of Deception annotation the the Through on database communities COG microbial the annotation 2). on of assembly Table based profile for (Supplementary genes 1,557 Taxonomic 842,798 KEGG databases and genes, on different rRNA Archaea, based used 16S genes within putative We 304,173 487 12,034 and in respectively. Bacteria, resulted viruses, that within and JGI/IMG assigned Eukarya through 353,731 within genes, 1,534 the putative and of under 1,396,820 strategies abundances in survival Relative resulted to related genes 2). (62.3%). novo samples. and Table De 296,818,692 glaciers potential (Supplementary were and metabolic viruses fumaroles reads the among as annotated compare in extremes taxonomically 79,111 to wereenvironmental predicted were used and 475,895,996 reads were proteins Eukarya genes which 162,755,88 as of detected of within 2,094,916 number total reads, Archaea, total A paired-end as The analyses. 3,680,020 567,410,264 further Bacteria, of for as total (Q>30) 98 quality a of by of produced filtered temperatures metagenomes HiSeq2000 0 with the Illumina around fumaroles analyzed of temperatures we comprising with volcano, samples, glaciers Island eighteen and Deception extreme of the across strategies total of survival a gradients with associated geochemical genes and and potential temperature metabolic between links investigate framework To PICA the using predicted were ( Results PhenDB Phenotypes and 2015). SEED sequences 2015) the al., al., to in et et compared and (Feldbauer 2016) (Brettin were al., proteins RASTtk et Further, (Kanehisa through (genus_prokaryotes) GhostKOALA Subsystem 2014). through (Seemann, Database KEGG v.14.5 the prokka in using performed was MAGs n .-.%i lces lhuhsm oiatpyawr omnbten<80 between common were phyla dominant some Although glaciers. in 0.4-0.6% and C classified phyla archaeal abundant most the with 87.3%), and 31.5 between abundance (relative fumarole C sebiso h ult-lee ed eeae oa f5395cnis h rdcino ORFs of prediction The contigs. 543,945 of total a generated reads quality-filtered the of assemblies https://phendb.csb.univie.ac.at/ o uaoe okn ttecaslvl hrorti Thermococci, Thermoprotei, level, class the at Looking fumarole. 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Data may be preliminary. eprtr Fgr ) h xdtv tesrsos a akdyhge ntefmrl t98 at genes fumarole the in in abundant higher markedly equally was were response environmental accordingly stress samples observed oxidative were The responses all biosynthesis, 4). specific from of protein (Figure patterns repair, temperature we communities distinct very DNA gradients, Although response, response, stress geochemical stress chemotaxis. to related with and and involved are temperature transport that extreme metagenomes and our under in strategies genes survival selected community understand To extremes environmental under 4). phosphate strategies (Figure survival pentose fixation Community was carbon the regulators and storage as fermentation, carbon photosynthesis, such of as pathways, function such <80 The carbon fermentation. in and central higher respiration significantly and were as heterotrophy glycolysis, with and pathway associated mainly 98 the were <80 in the to higher potential in belonging genetic much detected genes The was reduction. of fixation nitrite abundance dissimilatory carbon similar and for ammonification, a nitrite showed and fumaroles nitrate oxidation, All pathways. sulfur nitrogen metabolic prevalent the ( denitrification ( 0.02), temperature assimilation < highest sulfur the <80 inorganic in in to prevalent dominant related was genes reduction of sulfate ( number were While high sulfur to temperature. a related pathways fumarole, the metabolic to different temperatures, according all observed ( among ( abundant reduction was fixation metabolism sulfate dioxide sulfur carbon with (98 and associated temperature 0.001) highest temperatures functions environmental the for to with according higher fumarole diversity The metabolic 4). of (Figure patterns partitioning different temperatures observed extreme We among partitioning metabolic of Patterns with 98 the associated with genes comparison more 3c). in showed categories glaciers defense” and for and observations 0.049 disease contrast, “virulence, In and glaciers. “carbohydrates” to “photosynthesis” compared when 1.07e-4), <80 ( 0.023) = in chemotaxis” (p values and significant metabolism” “motility highest “nitrogen the categories exhibited The of glaciers. abundance and ( highest fumaroles ( the ( other plasmids” showed metabolism” and with method, elements “DNA transposable Newcombe-Wilson categories calculated prophages, the SEED, the “phages, of and groups to 1 test level prosthetic belonging from two-sided genes vitamins, functions exact “cofactors, and groups Fisher’s and sample using between metabolism” differences “DNA Significant pigments”. metabolism”, and functions derivatives”, and of “RNA a acids core Further, prevalent “amino metabolism”, the 3b). “carbohydrates”, The “protein (Figure subsystems”, observed. both “clustering-based was ordination through were PCA temperature samples observed highest through Deception the level were among from functional groups samples SEED sample between and and pattern distinct 3a) distinct 98 test, (Figure three profile the hypothesis these 1 between and between Level observed SEED differences method were multivariate Clear levels glaciers. a functional and using all compared in volcano were variations correlations. Island significant metagenomes positive Deception of of the on number profiles communities the temperatures, Functional in microbial higher increase of at an profile was temperature; functional there the Comparative temperatures lower to in according while even, changed is also proportion correlations positive/negative (0 of WBC the whereas structure, modular p p au a o significant, not was value .7-) slu eaoim ( metabolism” “sulfur 5.07e-3), = p .3,rsetvl)ad<80 and respectively) 0.036, = o uaoe hncmae oohrsmls ihntaeadntieamnfiain( ammonification nitrite and nitrate with samples, other to compared when fumaroles C o uaoe n lces ngair,tegnsietfidwti abnmetabolism carbon within identified genes the glaciers, In glaciers. and fumaroles C p .1,ntoe xto ( fixation nitrogen 0.01), < p o p uaoe,i diint h bevto fohrcro-eae processes, carbon-related other of observation the to addition in fumaroles, C .)wr eetdi <80 in detected were 0.1) < .1-) RAmtbls”( metabolism” “RNA 2.71e-8), = p p .0)we oprdt te uaoe n lces Although glaciers. and fumaroles other to compared when 0.001) < .1) n mtbls faoai opud”( compounds” aromatic of “metabolism and 0.015), = º )st a h es ope ewr.Itrsigy h proportion the Interestingly, network. complex least the had site C) o uaoe ( fumaroles C o uaoe( fumarole C o uaoe hncmae ote98 the to compared when fumaroles C 6 p p .5 n moi siiain( assimilation ammonia and 0.05) < p o .4) poenmtbls”( metabolism” “protein 0.046), = p p uaoe.I eea,ntoe eaoimwas metabolism nitrogen general, In fumaroles. C .0) ismltr irt euto ( reduction nitrite dissimilatory 0.001), < .7-)i h 98 the in 3.97e-3) = .8- and 1.68e-6 = o p )ehbtdmtblcptnilsignificantly potential metabolic exhibited C) .1,weespooytei a mainly was photosynthesis whereas 0.01), < p .1,ad“rti eaoim ( metabolism” “protein and 0.01), = p p o uaoe <80 fumarole, C .1,rsetvl)(Figure respectively) 0.012, = .0)adslu oxidation sulfur and 0.001) < o uaoei comparison in fumarole C o o uaoe and fumarole, C uaoe( fumarole C p .4) and 0.049), = p o fumaroles C .0)as 0.001) < p o ,mainly C, 0.036) = p p p < = = p Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. rnpsbeeeet n lsis.I otat uft a oiieycreae ih“carbohydrates”, with correlated positively also was Nitrate sulfate prophages, contrast, pigments”. “phages, In and and groups sporulation” plasmids”. prosthetic and and vitamins, “dormancy elements “carbohydrates”, “cofactors, transposable with and functions correlations comprised derivatives” negative correlations com- and positive presented whereas “motil- acids aromatic metabolism”, “carbohydrates”, “amino of as “RNA such and as functions “metabolism “sulfur “respiration” with negative chemotaxis”, correlated and acquisition”, negatively and exhibited response”, was ity “iron Ammonia which “stress 5b). isoprenoids”, metabolism”, those (Figure “secondary and metabolism” and “photosynthesis”, lipids metabolism”, metabolism”, “phosphorus acids, “RNA “DNA pounds”, were “fatty and temperature were with nucleotides” correlations correlations Al and positive presented and “nucleosides which Mg 1) Ca, metabolism”, level (SEED K, categories Na, functional organic correlations), The matter, negative organic (uniquely 3). Other Table Fe as Supplementary such correlations), 5a, phyla, negative others. (Figure several (uniquely among with Cu Verrucomicrobia, correlations B, negative and carbon, and Actinobacteria, with positive correlations Cyanobacteria, exhibited as negative Acidobacteria, parameters such Proteobac- and as phyla with Crenarchaeota, including correlations such and positive positive, phyla Deferribacteres significant were Euryarchaeota, showed correlations Thaumarchaeota, Sulfate nitrate teria, etc. significant Thermotogae, All Nitrospirae, Cyanobacteria, Firmicutes, others. Eu- Acidobacteria, Thaumarchaeota, among Firmicutes, Proteobacteria, were Spirochaetes, were correlated and ammonia positively with those Eu- others) Acti- correlated Deferribacteres; negatively were and among Spirochaetes, that ryarchaeota, temperature Bacteroidetes, phyla Gemmatimonadetes, The Acidobacteria, with and 5a). correlated Deinococcus-Thermus, (Figure (e.g. several positively Nitrospirae, temperature whereas that Verrucumicrobia, with Aquificae, phyla nobacteria, correlated and the negatively le- Thermotogae, general, were (SEED Nanoarchaeota, In phyla function Korarchaeota, ( and considered. Crenarchaeota, significant level) were only ryarchaeota, phylum then 0.6) (at calculated; or were taxonomy -0.6 correlations community Spearman of 1), drivers vel environmental key diversity identify functional To and taxonomic with on (all influence tungstate) Physical-chemical and oligopeptide acid, amino branched-chain <80 (e.g. prevalent transporters also least ABC were at and genes Chemotaxis Zn) samples. and (98 other Co, fumarole the temperature to highest compared the when in higher significantly were 0.001) (98 (98 with fumarole (all temperature glaciers highest the of communities using in also recombination found were repair, notably chemotaxis excision repair and Base 2). transport Figure biosynthesis, (Supplementary through protein temperatures repair, environmental DNA across of observed patterns abundance in Differences ( ( family function protein sporulation and dormancy ( of glaciers by followed fumaroles, ( heat-shocks chaperonin and fumaroles thermosome among as distributed distinctly was (including response) proteins heat-shock (thermal o samples, of responses function cold glaciers General and glaciers. in heat to prevalent ( related were osmoregulation genes genes as ( such stress with Z aquaporin functions (all osmotic by functions Contrastingly, mainly rubrerythrin samples). represented and other cycle) with (redox comparison glutathione glutaredoxins, by represented uaoe ( fumaroles C o ) ucin uha nvra Tae ( GTPases universal as such functions C); o uvrABC uaoe alwt tleast at with (all fumaroles C p recU .5.M rnpr n h B rnpreso rnadppie eesgicnl ihrin higher significantly were peptides and iron of transporters ABC the and transport Mn 0.05). < groEL p n ees yae(l with (all gyrase reverse and ope,rcmiainthrough recombination complex, p .5 n ytei fomrgltdprpamtcguas( glucans periplasmatic osmoregulated of synthesis and 0.05) = p .0) hra pcfi rheltemlrsossdmntdte98 the dominated responses thermal archaeal specific whereas 0.001), < p .1 Fgr 4). (Figure 0.01) < / .0) rti isnhssgnswr oiati h ihs eprtr fumarole temperature highest the in dominant were genes biosynthesis Protein 0.001). < groES ( p p .0) h eaieaudneo odshock cold of abundance relative The 0.001). < .0,07 fttlrltv bnac) swr atra n archaeal and bacterial were as abundance), relative total of 0.7% 0.001, < p .0) lcesad<80 and Glaciers 0.001). < p .5 SplmnayFgr 2). Figure (Supplementary 0.05) < p o p )( C) .1 n l uaoe a rvlneo h nvra stress universal the of prevalence a had fumaroles all and 0.01) < .1 eetemi taeiso N oiiesproln and supercoiling positive DNA of strategies main the were 0.01) < p p recA .0) swr eea rnpr ytm tasoto Ni, of (transport systems transport several were as 0.001), < .1 n rnlto lnainfcosi rhe ( Archaea in factors elongation translation and 0.01) < n htlaewr oiati <80 in dominant were photolyase and 7 hsp70 p .0) sortcat( osmoprotectant 0.001), < o uaoe xiie h ihs abundance highest the exhibited fumaroles C / dnaK p .5 n togcreain ( correlations strong and 0.05) < eepeaeti lcesad<80 and glaciers in prevalent were ) o p ) taeiso N repair DNA of Strategies C). .0) h bnac of abundance The 0.001). < cspA a ihri <80 in higher was yehX o o uaoe such fumarole, C uaoe and fumaroles C ( ) p .0 in 0.001 < p 0.01), = p r o > < C Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. es ee o irt euto ( reduction nitrate for (Hydrogenothermaceae/ genes DI_MAG_00004 ness) high-quality the in identified We 1). (Table (Calditrichia) 12 (Py- (Nitrososphaerales/ DI_MAG_00020 DI_MAG_00049 assigned ex- GW2011_AR20), were (Chitinophagaceae), to MAGs DI_MAG_00019 related selected rodictiaceae/ groups (Thermonemataceae), 11 on DI_MAG_00011 These based ), and metabolisms. quality (Promineofilaceae/ nitrogen their MAG_00006 ( and by sulfur DI_MAG_00003 annotation to drafts. as functional quality associated for medium and as selected tremophiles 82 total were and A quality MAGs 4). high Table The as Supplementary 4, considered were Figure (92), (Supplementary MAGs Bacteroidota (35) 13 (2), Proteobacteria of through Nitrospirota Aquificota and (1), classified Firmicutes (1), (4) (1), were Patescibacteria Acidobacteriota Cyanobacteriota (2), MAGs (3), phyla: Chloroflexota bacterial (1), following Campylobacterota The the (5), Calditrichota as (3). GhostKoala (Nanoarchaeota) and Woesearchaeia GTDB-Tk and eighteen (2) ( our Pyrodictiaceae Nitrososphaerales to for belonging GhostKoala, spumilus results and MAGs, GTDB-Tk not 159 binning was Nitrosocaldus the through through From which Archaea tus produced (Calditrichia) best 4). as MAG taxon Table Supplementary assigned a 1 the 3, were to only Figure 12 belonged (Supplementary analyses showed MAG pipeline our this anvi’o in the since co-assembly through included binning achieved We MaxBin using and MAGs. assembler 158 idba-ud pipeline of the total anvi’o a generating the metagenomes, general, 3). MAGs Table in In Supplementary strategies 5b, B, survival (Figure carbon, and Al organic matter, and potential organic Mg Metabolic ex- as Ca, such parameters K, functions, Other Na, several Si, metabolism”. with Fe, “RNA correlations Cu, negative and and “respiration” positive chemotaxis”, hibited and “motility metabolism”, “DNA ieetcl-hc ee eedtce mn As IMG006wsteoewihpeetdmore presented which one the was DI_MAG_00006 MAGs; among csp detected were genes and DI_MAG_00004 cold-shock Different for except fixation, carbon 6a). for in pathways (Figure DI_- incomplete detected DI_MAG_00021 and had DI_MAG_00022 also MAGs MAGs DI_MAG_00021, were for DI_MAG_00020, All except MAGs genes MAG_FBB2_12. MAGs, for reduction selected except all MAGs, Sulfate almost selected (e.g. different in DI_MAG_00022. identified response and were thermal cycle DI_MAG_00021, and nitrogen DI_MAG_00020, the rubredoxin) with and involved genes rubrerythrin oxidative general, especially dismutase, response, and stress with reductase groES associated superoxide genes several had (e.g. MAG stress This 6a). (Figure fixation carbon ( reduction fate h rmr olo u td a ouvi o omnte ucinlyrsodt h combination the to respond functionally communities how unveil to was study our of goal primary The 6b). (Figure Discussion DI_MAG_00004) in ( (only genes repair photolyase recombination wereand several ( identified genes were repair as mechanisms rubredoxin excision such repair nucleotide and DNA MAGs, potential Different rubrerythrin selected the DI_MAG_00003. DI_MAG_00049), in and showed (except DI_MAG_00004 MAGs in response observed all stress only Although oxidative heat- of different DI_MAG_00049). to presence and related DI_MAG_00022, were that (DI_MAG_00020, MAGs selected ( all Thermosome in including genes responses, observed shock we However, DI_MAG_00049). ee.W i o n any find not did We genes. , hsp20 1,Dsluoocls( Desulfurococcales (1), Pyrodictium thsA and sat codn ihGotol aooy (2), taxonomy) GhostKoala with according , hspR n ees yaegnswr dnie nalteslce Asasge sArchaea as assigned MAGs selected the all in identified were genes gyrase reverse and ) cysH Sulfurimonas ,a ieetDArpi ehnss nldn htlaerpi Fgr b.In 6b). (Figure repair photolyase including mechanisms, repair DNA different as ), groEL/groES , ,D_A_02 Djaatra,D_A_02 (Woesearchaeia/archaeon DI_MAG_00022 (Dojkabacteria), DI_MAG_00021 ), sir uvr ,slu n houft xdto ( oxidation thiosulfate and sulfur ), ee) obesrn ra ear( repair break double-strand genes), addtsPromineofilum Candidatus Aeropyrum narGHI csp ,D_A_00 (Hydrogenothermaceae/ DI_MAG_00004 ), ee nacaa As(IMG000 IMG002 and DI_MAG_00022, (DI_MAG_00020, MAGs archaeal in genes ee nD_A_00,D_A_02,adDI_MAG_00021. and DI_MAG_00020, DI_MAG_00004, in genes and codn ihGotol aooy 1,Aiioaee(1), Acidilobaceae (1), taxonomy) GhostKoala with according nirA ,dnticto ( denitrification ), 8 addtsNitrosocaldus Candidatus rec ,D_A_01 (Caldilineaceae/ DI_MAG_00010 ), Nitrosoarchaeum ee) N imthrpi ( repair mismatch DNA genes), soxAXBYZ her narGHI ,ol nacaa-eetdMAGs) archaeal-selected in only A, Persephonella ,adicmlt ahasfor pathways incomplete and ), ,nticto ( nitrification ), (1), n DI_MAG_FBB2_- and ) Nitrosotenius Persephonella 7 complete- 97% ~ , narGH mut (1), Caldilinea Candida- ,DI_- ), genes), Nitro- ,sul- ), Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. ufie hc a togrdciepwrcpbeo asn xdtv tes n yetempii tem- hyperthermophilic and stress, hydrogen oxidative of causing production of fumarolic capable the power reductive including strong factors, a stress re- has primaries heat-shock which its archaeal sulfide, reflects specific This stress, ( oxidative transporters. recombination with ABC repair, (98 associated excision fumarole those autotrophic base hottest as and sponses, reductive such the with strategies, from this microorganisms adaptive communities selecting of different addition, preferably conditions and In hyperthermophilic gradient and redox pathways. steep emissions a sulfide creating hydrogen ers, and the reduction, (98 that nitrite and fumarole (98 dissimilatory suggest fumarole location, and hottest We ammonification, the geographic reduction, fixation. in by sulfate carbon as potential 98 than such metabolic by pathways, rather reductive predominant grouped with are The temperature, Island by Deception glaciers. samples on and communities the microbial selective clustered that showed strong pattern the microbial the functional and that environment, reflects The marine indicates that the activity, results content volcanic our the genetic between Gordon, in diverse interaction & cryosphere. observed remarkable remarkably Turnbaugh a heterogeneity a 2018; by metabolic caused al., harbor pressures The et Deception Tully gut 2017). on 2019; and freshwater al., al., communities ecosystems, et et Trembath-Reichert subseafloor Varbiro cold envi- 2016; several back-arc, 2009; al., in Mariana et observed gradient. the et been (Louca from temperature Reveillaud has which microbiomes fluids variation sharp 2019; venting vents taxonomic the al., as the hydrothermal such across et despite ronments at heterogeneity redundancy (Galambos environ- functional out metabolic level steep of showed community observation carried the The communities the studies among Deception at of previous diversity 2016), redundancy diversity Unlike metabolic al., functional results functional of metabolic and These Island. partition to taxonomic Deception pointed a ammonia. both on observed with shapes gradients we correlated parameters mental Indeed, positively environmental fumaroles, were of communities. in glaciers mosaic microbial abundant in mainly the example, prevalent those that For were functions indicate sodium groups. and and functional groups sulfate, and temperature, while taxonomic by both influenced among positively varied groups drivers Island. environmental environmental Deception with these on al., although Correlation hyperthermophilic found et 2016), (Mandakovic community those al., that et Desert from of suggest (Lin Atacama different digestion patterns the results are anaerobic Similar in in conditions Our temperature conditions increasing stressors. stressful with environmental 2014). in and observed 2018) strong al., previously facing et been when have Sharp modulate interaction resilience to increase 2019; and members community temperature between resistance al., interactions with their ecological et in decreases trigger probably temperatures, Merino complexity Deception lowest community on 2013; the temperatures that (98 to al., showed fumarole compared et that hottest when (Cole studies interactions the positive previous in fewer to interaction presented community contrast and 99 the complex to that 2014). more 7.5 showed was al., from analysis et a ranging network (Sharp with gradient our Zealand consistent Surprisingly, temperature New also a and were along Canada patterns in members taxonomic phyla, areas similar our Thermotogae Furthermore, geothermal observed and Aquificae that method. the that report for analysis diversity by except previous for 2018), detected set al., not sample et were same Signori, the which (Bendia, using sequencing) Island gene Deception rRNA on hyperthermophilic we (16S out carried the addition, work previous 98 In temperature: with the consistent environmental in and phyla salinity. the <80 Thermotoga Firmicutes, and in and to Thaumarchaeota Aquificae Proteobacteria, geochemistry according Crenarchaeota/Thermoprotei, groups phyla to varied temperature, bacterial belonging the that members in some notably groups detected gradients the (most specific We for strong by sediments observed observed driven the fumarole 2018). previously and despite strongly al., we glacier et is what Bacteroidetes), proximity both Signori, to function of in similar (Bendia, community regardless diversity present occurred the that and as show Island, composition factors, results Deception community environmental Our contrasting on of volcanoes. glaciers combination marine and polar fumaroles of between typical factors environmental of o )(ooae l,20)mydces h isle xgnee ntesprca eietlay- sediment superficial the in even oxygen dissolved the decrease may 2004) al., et (Somoza C) o uaoe,adAioatraadVruoirbai lces hs atrsare patterns These glaciers. in Verrucomicrobia and Acidobacteria and fumaroles, C recU ,rvregrs,poenboytei,ceoai,and chemotaxis, biosynthesis, protein gyrase, reverse ), 9 o )ehbtdsvrlgnsrltdto related genes several exhibited C) o uaoe <80 fumarole, C o )wsmsl associated mostly was C) o o fumaroles C fumarole, C o in C o C) Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. oeta ettlrnefrti ieg u ote()lc fcl-hc ee,(hs ee r mostly and are thermosome genes gyrase, (these reverse archaeal of genes, psychrophilic/mesophilic presence cold-shock in the of (ii) present (e.g. and lack genes usually 2007), (i) heat-shock while al., other the et archaea, least Giaquinto to thermophilic at 2006; due of (Cavicchioli, or members lineage genomes MAG lifestyle and Woesearchaeia this the thermophilic size our for in genome of putative absent tolerance small analysis novel heat the genome a to The potential due suggests a 2015). lifestyle GW2011_AR20, DI_MAG_00022) al., pathogenic et archaeon or GW2011_AR20, (Castelle symbiotic to pathways (archaeon a broad biosynthesis related have some a closely to of previously to appears was lack described microorganisms and MAGs member samples these Nanoarchaeota/Woesearchaeia our aquifer underrepresented of in of and adaptation uncultivated One and an survival is parameters. which enabling periplas- extreme likely and of oxidative damage, cells ecosystems. starvation, protect combination Deception DNA volcanic carbon that and heat-shock, on marine proteins and gene-encoding stress, in presence cold with mic role including their equipped conditions, their are 2008), stressful MAGs Ar- understand several selected al., ammonia-oxidizing against better our thermophilic et of to of Torre majority investigation distribution the 2018; Furthermore, further ecological the encourages al., for and et outcome chaea as Daebeler novel path- a such 2018; represents nitrogen al., data, fumaroles ther- and et sulfur and genome (Abby predicted hyperthermophilic in environments different included ar- underrepresented of Since annotation of homologs groups range for chaeota/Woesearchaeia. containing archaeal phylogenetic selected lineages broad and MAGs as a 11 well ways, comprised as The volcano of lineages, Island phyla. effect mophilic Deception direct bacterial microbial the the different and from active elucidate ecosystems. chaeal recovered on truly volcanic to MAGs limits Antarctic studies the 159 in temperature more The revealing processes encourage potential biogeochemical in pathways and specific the metagenomics or level on of community extremes enzymes of the understanding temperature metabolic limitation at about increase key the known metabolism results of is Despite microbial our little inhibition 2014), pathways, to 2015). al., metabolic due et al., Sharp functioning et 2013; ecosystem al., (Hedlund affects et in (Cole increases temperature temperature ecosystems how as hydrothermal diversity Antarctic microbial and the in geothermal decrease of quantitative both conditions a shown freezing have studies predominant several the Although to 2016). stress, due osmotic al., availability with to et water associated (Wei compared liquid genes sulfide ecosystems low more when hydrogen exhibited and the variable communities temperatures reflects more glacier Further, hyperthermophilic which are the periods. that maintains long which conditions for fumarole, emissions redox hottest and of temperatures stability the fluctuating through to <80 mechanisms exposure in repair strategies nity DNA survival Diverse (Sup- and glaciers photolyase. functions, in towards and sporulation occurs sulfate) as and sources, dormancy (e.g. energy for genes, geochemicals different diversity pathways exploiting volcanic <80 metabolic for potential and The unavailable lowest of marine substrates fumaroles. This these of majority making heterotrophy. decrease the 1), as and Table the glaciers, plementary genes well by metabolism in Although as explained carbon pathways, detected be to level. oxidative were can community related pathways and were local nitrogen reductive communities the and both glacier at sulfur for processes dis- to trend biogeochemical and related a complex fixation, and suggests versatility nitrogen This metabolic denitrification, reduction. ammonification, nitrite oxidation, similatory sulfur <80 pathways: chemolithotrophic 2018; contrast, al., often et have Lemmens In mechanisms 2000; al., thermosome detected et and (Forterre 2014). also Archaea gyrase Klostermeier, were hyper(thermophilic) reverse & of transporters Lulchev 2017); groups ABC several Sun, of in were & mechanisms described types (Wang repair been Different DNA fields different hydrothermal metagenomes metagenomes. but Ilheya in 2011), Deception observed al., in to also et was compared (Xie chemotaxis al., Ridge when Fuca with et involved de found (Hedlund Juan genes at denaturation in vents Enrichment cell-component hydrothermal and from 2019). al., processes et metabolic Merino to 2015; disturbance induces that perature o o uaoe n lce omnte abrdmcaim o ohha n cold-shock and heat both for mechanisms harbored communities glacier and fumaroles C uaoe eedmntdb ee novdwt ieeteegtcand energetic different with involved genes by dominated were fumaroles C groES a Nitrosocaldus Ca. htaeesnilyrltdt hprtempie n etresponse. heat and (hyper)thermophiles to related essentially are that ) o uaoe n lcesmgtb xlie ycommu- by explained be might glaciers and fumaroles C a rvosyrpre nyi ersra geothermal terrestrial in only reported previously was 10 a Nitrosocaldus Ca. uvrABC complex, n Nanoar- and recA , Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. oa r ai uea n r noi alsRcaCmo o hi upr nsmln.W are We sampling. in support. support scientific Henrique their their Luiz for for Gamba Campos Dr. C. Rocha Maximiano, Rosa Almirante Carlos and vessel team Antonio research polar LECOM’s Dr. research to and the thankful Duleba, of very Wania crew the Dr. and Rosa, captain the thank relation- We financial interest. or of commercial conflict any potential of a absence Acknowledgments as the construed in be conducted could was research that the ships – that Foundation declare (CNPq Research authors The Paulo INCT-Criosfera Sao Develop- Statement and The Scientific Interest of 407816/2013-5) and (ProAntar). Technological Conflict (2012/23241-0). (CNPq Program of fellowship Antarctic Doctoral Microsfera Council ABs Brazilian National supported projects the FAPESP Brazilian and the the (CNPq) by of ment supported part and was 028306/2009) and study wrote data, This materials, paper. the reagents, the discussed contributed of BB experiments, drafts the reviewed CS paper. Funding designed and paper. the wrote and the of conceived and of drafts VP tools, drafts reviewed analysis reviewed paper. and and and the paper, paper, of reviewed the contributed the drafts and wrote LL reviewed wrote paper, analysis, network the data, tables. performed wrote the and LM data, discussed figures the paper. prepared discussed the analysis, and of the genome drafts paper, analyzed metagenome-assembled the experiments, of the wrote the discussion performed to analysis, regarding experiments, bioinformatic the insights designed performed new and data, conceived provides samples, the it collected and AB volcanoes. ecosystems Antarctic these the functional from inhabiting Contributions microbial extremophiles MAGs Author different of recover of response to capabilities the survival first a metagenomics and 98 the for metabolic shotgun to of lifestyle through (0 thermophilic one gradient reveal novel temperature was to extreme putative study the a first to a clarify diversity the for to know, lifestyle able we marine were as a we and MAGs, member of nega- Woesearchaeia Furthermore, and reconstruction members. (98 complex community the among fumarole more associations through hottest found competitive and We the trigger probably temperatures from conditions. stressors communities environmental hyperthermophilic temperature the and the among redox to interactions fluctuating tive responding to responding in microorganisms metabolic of specialized of capable with <80 strategies very Survival communities while are stress, heterotrophy. harbor fumarole oxidative and glaciers hottest metabolism and the carbon pathways, from to oxidative related and especially <80 reductive processes fumaroles both from communities with (98 while environmen- pathways, versatility temperatures steep autotrophic by hyperthermophilic and shaped that reductive are Island that with observed Deception processes metagenomic We the metabolic inhabiting from and gradients. microorganisms reconstruction strategies that tal adaptive genome evidence of variety with genomic a and together possess genetic contigs volcano first and the reads provide we of data, annotation the combining By hyperthermophile. or thermophile 2019), Forterre, likely & a Conclusion (Catchpole as mesophiles member in Woesearchaeia absent Halobacte- this but within to organisms, lineages pointing hyperthermophilic archaeal in mesophilic ubiquitously some found in detected and also Thaumarchaeota, were ria, genes heat-shock these Although Methanosarcina o uaoe n lce omnte osse ait fsrtge htare that strategies of variety a possesses communities glacier and fumaroles C p.(emn ta. 08,rvregrs steol protein only the is gyrase reverse 2018), al., et (Lemmens spp. 11 Ca.Nitrosocaldus o )o nAtrtcvlao utemr,orstudy our Furthermore, volcano. Antarctic an of C) o ieg.Orwr ersns sfar as represents, work Our lineage. )peeal eetmicroorganisms select preferably C) o ) hc niaeta h strong the that indicate which C), o hwahg metabolic high a show C Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. eda .G,Sgoi .N,Fac,D . ure .T . oann .J . elzr,V H. V. Pellizari, & M., J. B. Bohannan, D., Deception T. on Structure R. Community Microbial Duarte, Shapes C., Features Marine D. Antarctica. and Volcano, Soft- Franco, Island Geothermal Island Deception of N., & Source Mosaic from C. D., A thermophiles Galante, Signori, (2018). Open and F., G., Psychrophiles Rodrigues, A. An D., cold: T. Bendia, and R. hot Duarte, Gephi: in B., Contro, Surviving Antarctica. A., volcano, (2018). A. Pulschen, H. 19). G., V. G. Pellizari, marco Araujo, G., A. (2009, Bendia, Networks. M. Media Manipulating Jacomy, Social https://www.aaai.org/ocs/index.php/ICWSM/09/paper/view/154 and & and Exploring S., blogs for Heymann, Deception ware on M., hazard Volcanic Bastian, (2014). G. Aguirre-Diaz, & (1975). https://doi.org/10.1016/j.jvolgeores.2014.08.009 Antarctica). D., 168. G. Islands, Pedrazzi, Shetland T. J., (South Davies, Marti, Island A., & Island Geyer, J., Deception S., M. Bartolini, of Roobol, evolution Volcanic R., V. M. http://nora.nerc.ac.uk/id/eprint/509208/ L. Harvey, S., Islands: McNeil, Annotations I., Gerdes, Rapid A., Shetland K., R. McReath, Server: South RAST Formsma, Overbeek, E., The L., A., P. (2008). A. R. O. Baker, Osterman, Zagnitko, Edwards, R., T., Olson, . . Disz, J., . Technology. of B., G. Subsystems M., Diversity Parrello, Olsen, using T., DeJongh, F., Paczian, (2017). A., Meyer, D., Paarmann, O. M., A. K., K. Kubal, Best, M., Stetter, D., E. Bartels, Glass, . K., . Island. . R. 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All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. niov ieieadaerpeetdwt h encvrg fcnis n h Asredundancy, MAGs the dendrogram and clustering The contigs, reported. of SNVs coverage of mean number the and with mapped through recovered represented reads were total are that content, (MAGs) and GC genomes three pipeline metagenome-assembled completeness, the 158 5 the following v. of temperature, view anvi’o circular environmental A to S3. Figure Newcombe- according from colored method on the and 98 by based groups: calculated clustered STAMP intervals different are confidence through the Samples topoisomerase, visualized with Wilson. test, and were The two-sided helicase subsystems. exact Profiles SEED repair, Fisher’s chemotaxis. using using DNA reads and regarding metagenomic profiles of transport annotation functional and for biosynthesis, plots Integrated protein error the repre- Extended using contigs, S2. annotated of and Figure system. annotation IDBA-ud (IMG/M) through on Microbiomes constructed based & were taxonomy Genomes Contigs level. community Microbial phylum microbial the of at sented abundances Relative S1. Figure Tables and CheckM), on Figures described. (based Supplementary completeness are GhostKoala. and status, and redundancy quality GTDB-Tk content, genome on GC based the N50, classification and length, taxonomic their genome and total MAGs of selected Characteristics 11 the of List and 1. Table GTDB-Tk on Clusters (EC). (KO), based Orthology numbers legends KEGG classification Commission of Table Enzyme taxonomic identifiers are or with their codes (COG) here presented whereas MAGs Groups are pathway. Orthologous figures, Genes or of bottom. complete of cluster the or side at gene genes are of upper incomplete GhostKoala presence represents the the circle represents on me- sulfate, circle yellow including represented Mg, black a (MAGs), A Ca, and genomes (B). cluster/pathway, K, metagenome-assembled strategies adaptive gene selected Na, and 11 Si, (A) the P, potential of tabolic carbon), annotation Functional (organic 6. OC clay. Figure and matter), corre- (electrical silt, a (organic EC sand, in OM pH, nitrate, 0.05 (temperature), Zn, ammonia, < Temp Fe, nitrogen, p are: 1 Cu, parameters exhibited (level environmental B, that profile The conductivity), parameters functional represented. Only and are parameters. level) analysis environmental (phylum lation (A) and profile (B) taxonomic subsystem) between SEED correlation according Spearman colored 5. Figure and 98 clustered groups: are different three Samples the Newcombe-Wilson. glaciers. following from temperature, method environmental to the subsystems. by SEED calculated using reads intervals metagenomic of responses. annotation shock on heat/cold The based and STAMP osmotic, nitrogen through and sulfur, visualized oxidative including were including pathways, Profiles response, metabolic stress regarding and environmental profiles metabolisms, to functional carbon for and according plots colored error Extended and representing 4. 98 clustered is Figure groups: are Heatmap different (B). Samples three subsystem (C). SEED the functions the following 1 of temperature, 3 level at level of significant at abundances considered functions relative on were based Differences represented performed Newcombe-Wilson. through was are from ordination generated subsystems, method communities SEED the microbial on by based calculated of STAMP profiles The through general (A). visualized in functional reads, for the metagenomic modularity, plots of degree edges, annotation error cumulative and Extended and nodes 3. diameter, of Figure number length, the path as average such connectivity, Com- measures, complexity. node of distribution. and average set temperatures communities, environmental a of of on number increases based the calculated highlighting was and plexity point sampling each of p ausaerpeetdadwr acltduigFse’ xc w-ie et ihteconfidence the with test, two-sided exact Fisher’s using calculated were and represented are values p auswr acltduigFse’ xc w-ie etadtecndneitraswere intervals confidence the and test two-sided exact Fisher’s using calculated were values o uaoe <80 fumarole, C o uaoe n glaciers. and fumaroles C 18 o uaoe <80 fumarole, C p ausaerpeetdadwr calculated were and represented are values o o uaoe,<80 fumaroles, C uaoe n glaciers. and fumaroles C o uaoe and fumaroles C p .5 PCA 0.05. < Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. A oeKG/hs ol aooyGD-ktaxonomy GTDB-Tk taxonomy Koala KEGG/Ghost DI_MAG_00003 Code MAG completeness, and status. redundancy quality content, genome GC 1 the TABLE N50, and length, CheckM, based genome and classification anvi’o total taxonomic characteristics: on the their based GhostKoala, with MAGs and reconstructed GTDB-Tk all of on list complete A S4. Table data. nitrogen, mental sulfate, annotation. Mg, contigs Ca, S3. and Table K, reads Na, about Si, information P, General carbon), S2. (organic Table clay. conduc- OC and (electrical matter), silt, EC (organic sand, pH, OM nitrate, temperature, Zn, ammonia, including Fe, sample Cu, per B, tivity), data parameters mapping Physical-chemical read S1. on Table following based temperature, MAGs 98 environmental to quality their groups: according medium different colored and 82 and three clustered composition, and the are quality Samples sequence Z-score. high with their 13 sample the on per representing based Heatmap contigs S4. Figure of clustering hierarchical samples. the across shows distribution center the in IMG001UkonBcei;Ptsiatra okbcei 3,7 7,7 15 72 .2Medium 1.72 Medium 77.27 0.00 31.53 176,471 79.44 Medium High 44.63 7.88 83,443 0.55 8 734,971 62.85 88.43 40 37.53 49.00 1,155,662 10,795 22,003 324 152 3,082,789 2,518,210 Dojkabacteria Patescibacteria; Bacteria; Chitinophagaceae Chitinophagales; Bacteroidia; Bacteroidota; Thermonemataceae Bacteria; Cytophagales; Bacteroidia; Bacteroidota; Bacteria; Woesearchaeia Nanoarchaeota; Archaea; GW2011_AR20 archaeon DI_MAG_FBB2_12 Unknown DI_MAG_00049 DI_MAG_00022 Unknown DI_MAG_00021 Unknown DI_MAG_00020 DI_MAG_00019 DI_MAG_00011 DI_MAG_00010 DI_MAG_00006 DI_MAG_00004 P vle fSera orltoscmaigtxnmcadfntoa rfie ihteenviron- the with profiles functional and taxonomic comparing correlations Spearman of -values Sulfurimonas Caldithrix Nitrosocaldus Candidatus Pyrodictium Caldilinea Promineofilum Candidatus Persephonella o uaoe <80 fumarole, C o uaoe n glaciers. and fumaroles C 19 atra aplbceoa aplbcei;Cmyoatrls Thiovulaceae; Campylobacterales; Campylobacteria; Campylobacterota; Bacteria; atra adtiht;Clirci ,3,4 7 1243.09.400 High 0.00 95.54 Medium 39.10 1.94 21,244 60.52 Medium High 176 32.09 High 2,931,548 0.47 0.91 6,877 High 2.00 75.27 96.36 0.22 145 47.76 92.73 57.82 68,816 90,197 982,006 98.37 52.27 102,288 30.32 44,923 28 92 1,071,537 4,473,267 56 4,660,983 67 Calditrichia Calditrichota; 1,682,998 Bacteria; Nitrososphaerales Nitrososphaeria; Crenarchaeota; Archaea; Pyrodictiaceae Desulfurococcales; Thermoprotei; Crenarchaeota; Archaea; Caldilineaceae Caldilineales; Anaerolineae; Chloroflexota; Promineofilaceae Bacteria; Promineofilales; Anaerolineae; Chloroflexota; Hydrogenothermaceae Bacteria; Hydrogenothermales; Aquificae; Aquificota; Bacteria; Sulfurimonas oa eghCnisnme 5 CcnetCmltns eudnyDatQuality Draft Redundancy Completeness content GC N50 number Contigs Length Total ,1,7 43,7 94 79 .1High 0.41 97.95 39.42 32,276 94 1,912,170 Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary.

as.numeric(Abundance) 0.00 0.25 0.50 0.75 1.00 B1FA B3FB B2FB B1FC B3WA B2WA B1WB WBB3WBC1WBC2WBC3 WBB2 WBB1 WBA3 WBA2 WBA1 FBC3 FBC2 FBC1 FBB3 FBB2 FBB1 FBA3 FBA2 FBA1 A

C Relative abundance (%) as.numeric(Abundance)

0.00 0.25 0.50 as.numeric(Abundance)0.75 1.00 0.00 0.25 0.50 0.75 1.00

Whalers Bay Fumarola Bay WBB3WBC1WBC2WBC3 WBB2 WBB1 WBA3 WBA2 WBA1 FBC3 FBC2 FBC1 FBB3 FBB2 FBB1 FBA3 FBA2 FBA1

Whalers Bay Fumarole Bay WBB3WBC1WBC2WBC3 WBB2 WBB1 WBA3 WBA2 WBA1 FBC3 FBC2 FBC1 FBB3 FBB2 FBB1 FBA3 FBA2 FBA1 98 o B C A complexity temperature Fumarole Bay Samples WBA FBA 80 o

C

0 o C Samples Samples 50 o C Whalers Bay WBB FBB 10

o

C Phylum Fibrobacteres Euryarchaeota Elusimicrobia Dictyoglomi Deinococcus-Thermus Deferribacteres Cyanobacteria Crenarchaeota Chrysiogenetes Chloroflexi Chlorobi Chlamydiae Candidatus Poribacteria Bacteroidetes Aquificae Actinobacteria Acidobacteria 0 o C Phylum Phylum Fibrobacteres Euryarchaeota Elusimicrobia Dictyoglomi Deinococcus-Thermus Deferribacteres Cyanobacteria Crenarchaeota Chrysiogenetes Chloroflexi Chlorobi Chlamydiae Candidatus Poribacteria Bacteroidetes Aquificae Actinobacteria Acidobacteria Fibrobacteres Euryarchaeota Elusimicrobia Dictyoglomi Deinococcus-Thermus Deferribacteres Cyanobacteria Crenarchaeota Chrysiogenetes Chloroflexi Chlorobi Chlamydiae Candidatus Poribacteria Bacteroidetes Aquificae Actinobacteria Acidobacteria Verrucomicrobia unclassified (derived fromBacteria) Thermotogae Thaumarchaeota Tenericutes Synergistetes Spirochaetes Proteobacteria Planctomycetes Nitrospirae Nanoarchaeota Lentisphaerae Korarchaeota Gemmatimonadetes Fusobacteria Firmicutes WBC FBC

D C 20 Verrucomicrobia unclassified (derived fromBacteria) Thermotogae Thaumarchaeota Tenericutes Synergistetes Spirochaetes Proteobacteria Planctomycetes Nitrospirae Nanoarchaeota Lentisphaerae Korarchaeota Gemmatimonadetes Fusobacteria Firmicutes Verrucomicrobia unclassified (derived fromBacteria) Thermotogae Thaumarchaeota Tenericutes Synergistetes Spirochaetes Proteobacteria Planctomycetes Nitrospirae Nanoarchaeota Lentisphaerae Korarchaeota Gemmatimonadetes Fusobacteria Firmicutes unclassified (derived fromCandidatusPoribacteria) B unclassified (derived fromThaumarchaeota) unclassified (derived fromVerrucomicrobia) unclassified (derived fromNanoarchaeota) unclassified (derived fromProteobacteria) unclassified (derived fromEuryarchaeota) unclassified (derived fromCyanobacteria) unclassified (derived fromLentisphaerae) unclassified (derived fromBacteroidetes) unclassified (derived fromAcidobacteria) unclassified (derived fromKorarchaeota) unclassified (derived fromElusimicrobia) unclassified (derived fromBacteria) Gemmatimonadetes (class) Thermomicrobia (class) Thermomicrobia Chrysiogenetes (class) Chrysiogenetes Deferribacteres (class) Gammaproteobacteria Epsilonproteobacteria Actinobacteria (class) Actinobacteria Fibrobacteres (class) Acidobacteria (class) Acidobacteria Thermotogae (class) Thermotogae Elusimicrobia (class) Spirochaetes (class) Fusobacteria (class) Fusobacteria Alphaproteobacteria Deltaproteobacteria Chlamydiae (class) Betaproteobacteria Dehalococcoidetes Zetaproteobacteria Chloroflexi (class) Planctomycetacia Verrucomicrobiae Methanomicrobia Thermoplasmata Nitrospira (class) Methanobacteria Aquificae (class) Ktedonobacteria Sphingobacteria Spartobacteria Erysipelotrichi Gloeobacteria Methanococci Negativicutes Flavobacteria Thermoprotei Archaeoglobi Thermococci Dictyoglomia Halobacteria Solibacteres Methanopyri Cytophagia Bacteroidia Synergistia Deinococci Mollicutes Chlorobia Clostridia Opitutae Bacilli

FBA1 FBA2 FBA3 FBB1 FBB2 FBB3 FBC1 FBC2 FBC3 WBA1 WBA2 WBA3 WBB1 WBB2 WBB3 WBC1 WBC2 WBC3 SampleType Abundance abundance (%) Fumaroles <80 Glaciers Fumarole 98 Relative Relative Hyperhot fumarole Glacier Fumarole 40 30 20 10 0 o C o C Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. gradients-on-deception-island-volcano-antarctica potential-and-survival-strategies-of-microbial-communities-across-extreme-temperature- Figure_4.pdf file Hosted vial at available A https://authorea.com/users/350067/articles/475020-metabolic- 21 Fumaroles <80 Fumarole 98 Glaciers o C o C C B Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. Phages ProphagesTransposable elementsPlasmids Cofactors VitaminsProstheticGroupsPigments A B Metabolism ofAromaticCompounds Fatty AcidsLipidsandIsoprenoids Iron acquisitionandmetabolism Virulence DiseaseandDefenseVirulence unclassified (derived fromBacteria) Clustering basedsubsystems Clustering Regulation andCellsignaling Nucleosides andNucleotides Amino AcidsandDerivatives Cell DivisionandCycle Dormancy andSporulation Dormancy Phosphorus Metabolism Phosphorus Motility andChemotaxis Secondary Metabolism Secondary Potassium metabolism Cell Wall andCapsule Candidatus Poribacteria Membrane Transport Nitrogen Metabolism Deinococcus Protein Metabolism Gemmatimonadetes Sulfur Metabolism Stress Response RNA Metabolism DNA Metabolism Photosynthesis Thaumarchaeota Carbohydrates Verrucomicrobia Miscellaneous Planctomycetes Chrysiogenetes Nanoarchaeota Deferribacteres Crenarchaeota Proteobacteria Euryarchaeota Cyanobacteria Actinobacteria Lentisphaerae Bacteroidetes Fibrobacteres Acidobacteria Synergistetes Thermotogae Korarchaeota Elusimicrobia Spirochaetes Fusobacteria Respiration Chlamydiae Dictyoglomi Tenericutes Nitrospirae Chloroflexi Firmicutes − Aquificae Thermus Chlorobi

Temp Temp pH pH OM OM OC OC

22 B B Cu Cu Fe Fe Mn Mn Zn Zn P P Si Si N N NH4 NH4 NO3 NO3 SO4 SO4 Na Na K K Ca Ca

Spearman correlation Mg Spearman correlation Mg Al Al EC EC Sand Sand Silt Silt Clay Clay − − − − − 0.25 0.75 − 0.25 0.75 0.5 0.75 0.25 0.5 0.75 0.25 − − 0.5 0.5 0 1 0 1 1 1 Posted on Authorea 11 Aug 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159714983.38747883 | This a preprint and has not been peer reviewed. Data may be preliminary. gradients-on-deception-island-volcano-antarctica potential-and-survival-strategies-of-microbial-communities-across-extreme-temperature- Table_1.xlsx file Hosted soxAXBYZ (K17222,K17223,K17224,K17226,K17227) Dissimilatory Sulfate Reduction AndOxidation Reduction Dissimilatory Sulfate narGHJI (K00370,K00370,K00371,K00373,K00374) Phosphate Acetyltransferase-AcetateKinasePathway vial at available A cysNC, cysN,cysD(K00955,K00956,K00957) Hydroxypropionate-Hydroxybutylate Cycle Dicarboxylate-Hydroxybutyrate Cycle Dissimilatory NitrateReduction Assimilatory NitrateReduction amoABC (K10944,K10945,K10946) Assimilatory Sulfate Reduction Assimilatory Sulfate Reductive PentosePhosphateCycle narGHI (K00370,K00371,K00374) narGHI(K00370, K00371,K00374) Methylamine/Dimethylamine/Trimethylamine =>Methane 3-Hydroxypropionate Bi-Cycle soxAXBYZ (K17222,K17223,K17224,K17226,K17227) Dissimilatory Sulfate Reduction AndOxidation Reduction Dissimilatory Sulfate Reductive CitricAcidCycle Phosphate Acetyltransferase-AcetateKinasePathway A Thiosulfate Oxidation Thiosulfate napAB (K02567,K02568) napAB (K02567,K02568) nasAB (K00372,K00360) Wood-Ljundahl Pathway narGH(K00370, K00371) nirBD (K00362,K00363) cysJI (K00380,K00381) Methane Oxidation-=>Formaldehyde cysNC, cysN,cysD(K00955,K00956,K00957) Carbon Fixation Denitrification Hydroxypropionate-Hydroxybutylate Cycle norC (K02305) nosZ (K00376) cysH (K00390) norB (K04561) cysC (K00860) Nitrification nirA (K00366) Hydrogenothermaceae/ nirK (K00368) sat (K00958) sat (K00958) sir (K00392) Dicarboxylate-Hydroxybutyrate Cycle Dissimilatory NitrateReduction Assimilatory NitrateReduction amoABC (K10944,K10945,K10946) Assimilatory Sulfate Reduction Assimilatory Sulfate Reductive PentosePhosphateCycle narGHI (K00370,K00371,K00374) narGHI(K00370, K00371,K00374) gene cluster/pathway Presence ofgenesorcomplete

Persephonella 3-Hydroxypropionate Bi-Cycle Reductive CitricAcidCycle Thiosulfate Oxidation Thiosulfate Methane Metabolism Methane napAB (K02567,K02568) napAB (K02567,K02568) nasAB (K00372,K00360) Wood-Ljundahl Pathway narGH(K00370, K00371) nirBD (K00362,K00363) cysJI (K00380,K00381) Methanol =>Methane Calditrichia Acetate =>Methane DI_MAG_00004 Carbon Fixation CO

Sulfurimonas Denitrification norC (K02305) nosZ (K00376) cysH (K00390) norB (K04561) cysC (K00860) Nitrification narB (K00367) Ca. 2 DI_MAG_FBB2_12 nirA (K00366) nirK (K00368)

Caldilineaceae/ =>Methane sat (K00958) sat (K00958)

Promineofilum sir (K00392) https://authorea.com/users/350067/articles/475020-metabolic- DI_MAG_00003 Thermonemataceae Caldilinea DI_MAG_00006

Pyrodictiaceae/Chitinophagaceae DI_MAG_00004DI_MAG_00010

DI_MAG_FBB2_12DI_MAG_00011 archaeon GW2011_AR20 Pyrodictium Woesearchaeia/ DI_MAG_00003DI_MAG_00019 Nitrososphaerales/ Ca. DI_MAG_00006DI_MAG_00020 Nitrosocaldus DI_MAG_00010DI_MAG_00022 pathway Incomplete genecluster/ Dojkabacteria DI_MAG_00011DI_MAG_00049

DI_MAG_00019DI_MAG_00021

DI_MAG_00020

DI_MAG_00022 recJFQGR (COG0608,COG1195,COG0514,COG1200,COG0353)

23 DI_MAG_00049

DI_MAG_00021 Carbon starvationprotein A,Carbonstorageregulator Peroxide stress regulator PerR,FURfamily(COG0735) DNA protection duringstarvationprotein (COG2406) Superoxide dismutase(EC1.15.1.1)(COG0605) Superoxide reductase (EC1.15.1.2)(COG2033) B recANRO (K03553,K03631,K06187,K03584) uvrABCD (K03701,K03702,K03703,K03657) recJFQGR (COG0608,COG1195,COG0514,COG1200,COG0353) Transcriptional regulator, Crp/Fnr family Thermosome thsA(arCOG01257) Thermosome Protease/chaperone protein htrA DNA RepairandSupercoiling Periplasmic Stress Response mutXT (COG0494,COG0406) Carbon starvationprotein A,Carbonstorageregulator Peroxide stress regulator PerR,FURfamily(COG0735) DNA protection duringstarvationprotein (COG2406) Reverse gyrase(COG1110) Universal Stress Proteins B Rubrerythrin (COG1592) mutSL (K07456,K03572) Photolyase (EC4.1.99.3) Rubredoxin (COG1773) Heat-shock Response Cold-shock Response Superoxide dismutase(EC1.15.1.1)(COG0605) Superoxide reductase (EC1.15.1.2)(COG2033) recANRO (K03553,K03631,K06187,K03584) uvrABCD (K03701,K03702,K03703,K03657) hsp16.5 (COG0071) Carbon Starvation mutYM (COG0266) cspG (COG1278) Oxidative StressOxidative hspA (COG0071) yceD (COG2310) cspC (COG1278) cspA (COG1278) dps2 (COG0783) yocK (COG1734) teaD (COG0589) cspV (COG1278) cspE (COG1278) hrcA (COG1420) cspJ (COG1278) surA (COG0760) herA (COG0433) nreC (COG2197) nreA (COG1602) groEL (K04077) hsp20 (K13993) groES (K04078) ctc (COG1825) grpE (K03687) xpf (COG1948) dnaK (K04043) hspR (K13640) dnaJ (K03686) Transcriptional regulator, Crp/Fnrfamily narJ (K00373) Hydrogenothermaceae/ Thermosome thsA(arCOG01257) Thermosome Periplasmic Stress Response Protease/chaperone protein htrA mutXT (COG0494,COG0406)

Persephonella Universal Stress Proteins mutSL (K07456,K03572) Photolyase (EC4.1.99.3) Rubrerythrin (COG1592) Heat-shock proteins Cold-shock proteins Rubredoxin (COG1773) Carbon Starvation hsp16.5 (COG0071) mutYM (COG0266) Calditrichia StressOxidative DI_MAG_00004 herA (COG0433) nreC (COG2197) nreA (COG1602) cspG (COG1278) hspA (COG0071) yceD (COG2310) cspC (COG1278) cspA (COG1278) dps2 (COG0783) yocK (COG1734) teaD (COG0589) cspV (COG1278) cspE (COG1278) hrcA (COG1420) cspJ (COG1278) surA (COG0760) groEL (K04077) xpf (COG1948) hsp20 (K13993) groES (K04078) radA (K04485) ctc (COG1825) grpE (K03687) DNA Repair Sulfurimonas dnaK (K04043) hspR (K13640) dnaJ (K03686) Ca. narJ (K00373) DI_MAG_FBB2_12 Caldilineaceae/ Promineofilum DI_MAG_00003

Thermonemataceae Caldilinea DI_MAG_00006

DI_MAG_00004 Pyrodictiaceae/Chitinophagaceae DI_MAG_00010

DI_MAG_FBB2_12 DI_MAG_00011 archaeon GW2011_AR20 Pyrodictium DI_MAG_00003 Woesearchaeia/ DI_MAG_00019 DI_MAG_00006 Nitrososphaerales/ DI_MAG_00020 Ca. Nitrosocaldus DI_MAG_00010 DI_MAG_00022 Dojkabacteria DI_MAG_00011 DI_MAG_00049 DI_MAG_00019 DI_MAG_00021

DI_MAG_00020

DI_MAG_00022

DI_MAG_00049

DI_MAG_00021