{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T23:23:41Z","timestamp":1787009021425,"version":"build-2736575974"},"reference-count":119,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,3,23]],"date-time":"2026-03-23T00:00:00Z","timestamp":1774224000000},"content-version":"vor","delay-in-days":81,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000780","name":"European Union","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000780","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004837","name":"Spanish Ministerio de Ciencia e Innovaci\u00f3n","doi-asserted-by":"crossref","award":["MCIN\/AEI\/10.13039\/501100011033"],"award-info":[{"award-number":["MCIN\/AEI\/10.13039\/501100011033"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Ram\u00f3n y Cajal","award":["RyC2021\u2013031775-I"],"award-info":[{"award-number":["RyC2021\u2013031775-I"]}]},{"name":"H2020 MSCA-IF-TURBOMUD","award":["101018321"],"award-info":[{"award-number":["101018321"]}]},{"DOI":"10.13039\/100004807","name":"DFG","doi-asserted-by":"publisher","award":["ME 5011\/1\u20131"],"award-info":[{"award-number":["ME 5011\/1\u20131"]}],"id":[{"id":"10.13039\/100004807","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100004807","name":"DFG","doi-asserted-by":"publisher","award":["M149"],"award-info":[{"award-number":["M149"]}],"id":[{"id":"10.13039\/100004807","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"crossref","award":["EXC 2077\u2013390741603"],"award-info":[{"award-number":["EXC 2077\u2013390741603"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,14]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Deep marine sediments generate large amounts of methane, but most of this gas is consumed by the anaerobic oxidation of methane (AOM) mediated by microscopic consortia of anaerobic methane-oxidizing archaea (ANME) and sulfate-reducing bacteria (SRB). In this study, we investigated the AOM within a sulfate\u2013methane transition zone (SMTZ) at a depth of ~9.6\u00a0m at the rim of the Ginsburg mud volcano in the Gulf of C\u00e1diz. The SMTZ is supplied with sulfate from both overlying seawater and an underlying evaporitic deposit, and it coincides with a fracture zone that hosts a visible biofilm. Here, carbon dioxide shows the strongest 13C-depletion, indicating intense methane consumption. Metagenomic and lipid biomarker analysis of the biofilm revealed an exceptionally simple microbial community dominated by ANME-1b archaea (63%), which predominantly produce strongly 13C-depleted glycerol dialkyl glycerol tetraethers and, to a lesser extent, the less common macrocyclic archaeols. The putative partner bacterium Seep-SRB1c (Desulfobacterota) is less abundant (9%). Additionally, the biofilm contained five low-abundance heterotrophs that likely rely on biomass or metabolites released from the ANME-SRB consortium. Our study highlights the presence of active methanotrophic biofilms in subsurface sediments and suggests that these communities may play an overlooked role in mitigating seafloor methane emissions.<\/jats:p>","DOI":"10.1093\/ismeco\/ycag072","type":"journal-article","created":{"date-parts":[[2026,3,21]],"date-time":"2026-03-21T12:39:31Z","timestamp":1774096771000},"source":"Crossref","is-referenced-by-count":1,"title":["Anaerobic oxidation of methane supports a minimal microbial community in a subsurface biofilm at Ginsburg mud volcano"],"prefix":"10.1093","volume":"6","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-2266-0112","authenticated-orcid":false,"given":"Cleopatra","family":"Collado","sequence":"first","affiliation":[{"name":"Museo Nacional de Ciencias Naturales (MNCN), Spanish National Research Council (CSIC) , 28006 Madrid ,","place":["Spain"]},{"name":"Secci\u00f3n departamental de Nutrici\u00f3n y Ciencia de los Alimentos (NUTRYCIAL), Facultad de Veterinaria, Universidad Complutense de Madrid (UCM) , Madrid ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-9766-6145","authenticated-orcid":false,"given":"Pedro","family":"Romero-Tena","sequence":"additional","affiliation":[{"name":"Museo Nacional de Ciencias Naturales (MNCN), Spanish National Research Council (CSIC) , 28006 Madrid ,","place":["Spain"]},{"name":"Unit\u00e9 d' Ecologie Soci\u00e9t\u00e9 Evolution , Universit\u00e9 Paris-Saclay AgroParisTech, CNRS, Gif-sur-Yvette,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6819-373X","authenticated-orcid":false,"given":"Gunter","family":"Wegener","sequence":"additional","affiliation":[{"name":"Max Planck Institute for Marine Microbiology , 28359, Bremen ,","place":["Germany"]},{"name":"MARUM, Center for Marine Environmental Science, University of Bremen , 28359, Bremen ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3863-0975","authenticated-orcid":false,"given":"Marcus","family":"Elvert","sequence":"additional","affiliation":[{"name":"MARUM, Center for Marine Environmental Science, University of Bremen , 28359, Bremen ,","place":["Germany"]},{"name":"Faculty of Geosciences, University of Bremen , 28359, Bremen ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8407-7528","authenticated-orcid":false,"given":"Walter","family":"Menapace","sequence":"additional","affiliation":[{"name":"MARUM, Center for Marine Environmental Science, University of Bremen , 28359, Bremen ,","place":["Germany"]},{"name":"Institute of Marine Sciences (ICM), Spanish National Research Council (CSIC) , 08003, Barcelona ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6912-7865","authenticated-orcid":false,"given":"Rafael","family":"Laso-P\u00e9rez","sequence":"additional","affiliation":[{"name":"Museo Nacional de Ciencias Naturales (MNCN), Spanish National Research Council (CSIC) , 28006 Madrid ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,3,23]]},"reference":[{"key":"2026042501113680300_ref1","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1021\/cr050362v","article-title":"Oceanic methane biogeochemistry","volume":"107","author":"Reeburgh","year":"2007","journal-title":"Chem Rev"},{"key":"2026042501113680300_ref2","doi-asserted-by":"publisher","first-page":"697","DOI":"10.5194\/essd-8-697-2016","article-title":"The global methane budget 2000\u20132012","volume":"8","author":"Saunois","year":"2016","journal-title":"Earth System Science Data"},{"key":"2026042501113680300_ref3","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1146\/annurev-micro-111021-045911","article-title":"Anaerobic degradation of alkanes by marine archaea","volume":"76","author":"Wegener","year":"2022","journal-title":"Ann Rev Microbiol"},{"key":"2026042501113680300_ref4","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1038\/s41561-018-0122-8","article-title":"Global diffusive fluxes of methane in marine sediments","volume":"11","author":"Egger","year":"2018","journal-title":"Nat Geosci"},{"key":"2026042501113680300_ref5","doi-asserted-by":"publisher","first-page":"e3001508","DOI":"10.1371\/journal.pbio.3001508","article-title":"Comparative genomics reveals electron transfer and syntrophic mechanisms differentiating methanotrophic and methanogenic archaea","volume":"20","author":"Chadwick","year":"2022","journal-title":"PLoS Biol"},{"key":"2026042501113680300_ref6","doi-asserted-by":"publisher","first-page":"e3002292","DOI":"10.1371\/journal.pbio.3002292","article-title":"Physiological potential and evolutionary trajectories of syntrophic sulfate-reducing bacterial partners of anaerobic methanotrophic archaea","volume":"21","author":"Murali","year":"2023","journal-title":"PLoS Biol"},{"key":"2026042501113680300_ref7","doi-asserted-by":"publisher","first-page":"1651","DOI":"10.1111\/1462-2920.14077","article-title":"Gene expression and ultrastructure of meso- and thermophilic methanotrophic consortia","volume":"20","author":"Krukenberg","year":"2018","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref8","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1038\/nature15512","article-title":"Single cell activity reveals direct electron transfer in methanotrophic consortia","volume":"526","author":"McGlynn","year":"2015","journal-title":"Nature."},{"key":"2026042501113680300_ref9","doi-asserted-by":"publisher","first-page":"eadw4289","DOI":"10.1126\/sciadv.adw4289","article-title":"Redox conduction facilitates direct interspecies electron transport in anaerobic methanotrophic consortia. Science","volume":"11","author":"Yu","year":"2025","journal-title":"Advances."},{"key":"2026042501113680300_ref10","doi-asserted-by":"publisher","first-page":"988871","DOI":"10.3389\/fmicb.2022.988871","article-title":"Deep-branching ANME-1c archaea grow at the upper temperature limit of anaerobic oxidation of methane","volume":"13","author":"Benito Merino","year":"2022","journal-title":"Front Microbiol"},{"key":"2026042501113680300_ref11","doi-asserted-by":"publisher","first-page":"422","DOI":"10.1111\/j.1462-2920.2009.02083.x","article-title":"Metagenome and mRNA expression analyses of anaerobic methanotrophic archaea of the ANME-1 group","volume":"12","author":"Meyerdierks","year":"2010","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref12","doi-asserted-by":"publisher","DOI":"10.1128\/mmbr.00074-18","article-title":"Physiology and distribution of archaeal Methanotrophs that couple anaerobic oxidation of methane with Sulfate reduction","volume":"83","author":"Bhattarai","year":"2019","journal-title":"Microbiol Mol Biol Rev"},{"key":"2026042501113680300_ref13","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1038\/s41396-020-00757-1","article-title":"Experimentally-validated correlation analysis reveals new anaerobic methane oxidation partnerships with consortium-level heterogeneity in diazotrophy","volume":"15","author":"Metcalfe","year":"2021","journal-title":"The ISME Journal"},{"key":"2026042501113680300_ref14","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1038\/s41396-021-01047-0","article-title":"Sulfate differentially stimulates but is not respired by diverse anaerobic methanotrophic archaea","volume":"16","author":"Yu","year":"2022","journal-title":"The ISME Journal."},{"key":"2026042501113680300_ref15","doi-asserted-by":"publisher","first-page":"3073","DOI":"10.1111\/1462-2920.13283","article-title":"Candidatus Desulfofervidus auxilii, a hydrogenotrophic sulfate-reducing bacterium involved in the thermophilic anaerobic oxidation of methane","volume":"18","author":"Krukenberg","year":"2016","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref16","doi-asserted-by":"publisher","first-page":"2327","DOI":"10.1111\/j.1462-2920.2010.02275.x","article-title":"Identification of the dominant sulfate-reducing bacterial partner of anaerobic methanotrophs of the ANME-2 clade","volume":"12","author":"Schreiber","year":"2010","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref17","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1146\/annurev.micro.61.080706.093130","article-title":"Anaerobic oxidation of methane: Progress with an unknown process","volume":"63","author":"Knittel","year":"2009","journal-title":"Ann Rev Microbiol"},{"key":"2026042501113680300_ref18","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1007\/s10347-005-0059-4","article-title":"Methane-derived carbonate build-ups and associated microbial communities at cold seeps on the lower Crimean shelf (Black Sea)","volume":"51","author":"Reitner","year":"2005","journal-title":"Facies."},{"key":"2026042501113680300_ref19","doi-asserted-by":"publisher","first-page":"1013","DOI":"10.1126\/science.1072502","article-title":"Microbial reefs in the Black Sea Fueled by anaerobic oxidation of methane","volume":"297","author":"Michaelis","year":"2002","journal-title":"Science."},{"key":"2026042501113680300_ref20","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1128\/AEM.71.1.467-479.2005","article-title":"Diversity and distribution of Methanotrophic archaea at cold seeps","volume":"71","author":"Knittel","year":"2005","journal-title":"Appl Environ Microbiol"},{"key":"2026042501113680300_ref21","doi-asserted-by":"publisher","first-page":"9725","DOI":"10.1038\/s41598-019-46209-5","article-title":"Methane-fuelled biofilms predominantly composed of methanotrophic ANME-1 in Arctic gas hydrate-related sediments","volume":"9","author":"Gr\u00fcndger","year":"2019","journal-title":"Sci Rep"},{"key":"2026042501113680300_ref22","doi-asserted-by":"publisher","first-page":"6780","DOI":"10.1128\/AEM.00288-11","article-title":"Macroscopic biofilms in fracture-dominated sediment that anaerobically oxidize methane","volume":"77","author":"Briggs","year":"2011","journal-title":"Appl Environ Microbiol"},{"key":"2026042501113680300_ref23","doi-asserted-by":"publisher","first-page":"6296","DOI":"10.1038\/s41467-021-26549-5","article-title":"Distinct methane-dependent biogeochemical states in Arctic seafloor gas hydrate mounds","volume":"12","author":"Klasek","year":"2021","journal-title":"Nat Commun"},{"key":"2026042501113680300_ref24","first-page":"1","article-title":"Seafloor chemosynthetic habitats and AOM-influenced sediment microbiome at a cold-water coral site off the Vester&aring;len coast, northern Norway","author":"Argentino","year":"2025","journal-title":"EGUsphere."},{"key":"2026042501113680300_ref25","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/j.epsl.2008.12.005","article-title":"The quest for the Africa\u2013Eurasia plate boundary west of the strait of Gibraltar","volume":"280","author":"Zitellini","year":"2009","journal-title":"Earth Planet Sci Lett"},{"key":"2026042501113680300_ref26","doi-asserted-by":"publisher","DOI":"10.1029\/2011GC003576","article-title":"Insights into active deformation in the Gulf of Cadiz from new 3-D seismic and high-resolution bathymetry data","volume":"12","author":"Crutchley","year":"2011","journal-title":"Geochem Geophys Geosyst"},{"key":"2026042501113680300_ref27","doi-asserted-by":"publisher","first-page":"850","DOI":"10.1111\/j.1365-246X.2011.05328.x","article-title":"Neotectonics of the SW Iberia margin, gulf of Cadiz and Alboran Sea: a reassessment including recent structural, seismic and geodetic data","volume":"188","author":"Cunha","year":"2012","journal-title":"Geophys J Int"},{"key":"2026042501113680300_ref28","doi-asserted-by":"publisher","first-page":"2-1-2","DOI":"10.1029\/2000RG000093","article-title":"Significance of mud volcanism","volume":"40","author":"Kopf","year":"2002","journal-title":"Rev Geophys"},{"key":"2026042501113680300_ref29","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1007\/1-4020-3204-8_13","volume-title":"Mud Volcanoes, Geodynamics and Seismicity","author":"Judd","year":"2005"},{"key":"2026042501113680300_ref30","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1007\/1-4020-3204-8_3","volume-title":"Mud Volcanoes, Geodynamics and Seismicity","author":"Milkov","year":"2005"},{"key":"2026042501113680300_ref31","doi-asserted-by":"publisher","first-page":"105188","DOI":"10.1016\/j.marpetgeo.2021.105188","article-title":"Mud volcanoes in the Gulf of Cadiz as a manifestation of tectonic processes and deep-seated fluid mobilization","volume":"132","author":"Xu","year":"2021","journal-title":"Mar Pet Geol"},{"key":"2026042501113680300_ref32","doi-asserted-by":"publisher","first-page":"101653","DOI":"10.1016\/j.jog.2019.101653","article-title":"Recent deformational state from morphological analysis of mud volcanoes in the Gulf of Cadiz (southwestern part of the Iberian Atlantic margin)","volume":"132","author":"Maestro","year":"2019","journal-title":"J Geodyn"},{"key":"2026042501113680300_ref33","doi-asserted-by":"publisher","first-page":"1232","DOI":"10.1016\/j.gca.2006.11.022","article-title":"Sources of mud volcano fluids in the Gulf of Cadiz\u2014indications for hydrothermal imprint","volume":"71","author":"Hensen","year":"2007","journal-title":"Geochim Cosmochim Acta"},{"key":"2026042501113680300_ref34","doi-asserted-by":"publisher","first-page":"B05101","DOI":"10.1029\/2010JB008061","article-title":"Linked halokinesis and mud volcanism at the Mercator mud volcano, gulf of Cadiz","volume":"116","author":"Perez-Garcia","year":"2011","journal-title":"Journal of Geophysical Research: Solid Earth"},{"key":"2026042501113680300_ref35","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/j.marpetgeo.2005.11.001","article-title":"Molecular and carbon isotopic variability of hydrocarbon gases from mud volcanoes in the Gulf of Cadiz, NE Atlantic","volume":"23","author":"Stadnitskaia","year":"2006","journal-title":"Mar Pet Geol"},{"key":"2026042501113680300_ref36","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1016\/j.margeo.2019.01.005","article-title":"Formation and migration of hydrocarbons in deeply buried sediments of the Gulf of Cadiz convergent plate boundary\u2014insights from the hydrocarbon and helium isotope geochemistry of mud volcano fluids","volume":"410","author":"Nuzzo","year":"2019","journal-title":"Mar Geol"},{"key":"2026042501113680300_ref37","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/S0025-3227(02)00688-6","article-title":"Gas hydrates in the Ginsburg and Yuma mud volcano sediments (Moroccan margin): results of chemical and isotopic studies of pore water","volume":"195","author":"Mazurenko","year":"2003","journal-title":"Mar Geol"},{"key":"2026042501113680300_ref38","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1080\/01490450802258196","article-title":"Shallow microbial recycling of deep-sourced carbon in gulf of Cadiz mud volcanoes","volume":"25","author":"M","year":"2008","journal-title":"Geomicrobiol J"},{"key":"2026042501113680300_ref39","doi-asserted-by":"publisher","first-page":"350","DOI":"10.1016\/j.chemgeo.2009.06.023","article-title":"Origin of light volatile hydrocarbon gases in mud volcano fluids, gulf of Cadiz\u2014evidence for multiple sources and transport mechanisms in active sedimentary wedges","volume":"266","author":"Nuzzo","year":"2009","journal-title":"Chem Geol"},{"key":"2026042501113680300_ref40","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1016\/j.margeo.2008.10.007","article-title":"Tectonics and mud volcano development in the Gulf of C\u00e1diz","volume":"261","author":"Medialdea","year":"2009","journal-title":"Mar Geol"},{"key":"2026042501113680300_ref41","first-page":"53","volume-title":"Fluid Seepages\/Mud Volcanism in the Mediterranean and Adjacent Domains: CIESM Workshop Monographs","author":"Pinheiro","year":"2006"},{"key":"2026042501113680300_ref42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.margeo.2005.04.007","article-title":"The El Arraiche mud volcano field at the Moroccan Atlantic slope, gulf of Cadiz","volume":"219","author":"Van Rensbergen","year":"2005","journal-title":"Mar Geol"},{"key":"2026042501113680300_ref43","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1007\/s00367-016-0440-z","article-title":"Evidence of episodic long-lived eruptions in the Yuma, Ginsburg, Jes\u00fas Baraza and Tasyo mud volcanoes, gulf of C\u00e1diz","volume":"36","author":"Toyos","year":"2016","journal-title":"Geo-Mar Lett"},{"key":"2026042501113680300_ref44","doi-asserted-by":"publisher","first-page":"3889","DOI":"10.1130\/B37954.1","article-title":"Complex architecture of mud volcano systems: new insights on flow pathways unravel intricate fluid circulation","volume":"137","author":"Xu","year":"2025","journal-title":"GSA Bull"},{"key":"2026042501113680300_ref45","doi-asserted-by":"publisher","first-page":"22","DOI":"10.5194\/sd-4-22-2007","article-title":"The IODP expedition 302 scientists. Rhizon sampling of pore waters on scientific drilling expeditions: an example from the IODP expedition 302, Arctic coring expedition (ACEX)","volume":"4","author":"Dickens","year":"2007","journal-title":"Sci Drill"},{"key":"2026042501113680300_ref46","volume-title":"Preliminary Rresults of R\/V METEOR cruise M149: Shipboard and Post-Cruise Analysis, Recurrence of Ttsunamigenic Hhazards from MeBo Ddrilling Rrecords and Hhazard Mmitigation Uusing MeBo Oobservatories, Las Palmas (Canary Islands) \u2013 Cadiz (Spain), 24.07.2018\u201324.08.2018","author":"H\u00fcpers","year":"2020"},{"key":"2026042501113680300_ref47","doi-asserted-by":"publisher","first-page":"6033","DOI":"10.1016\/j.gca.2010.08.006","article-title":"The biogeochemistry of sorbed methane in marine sediments","volume":"74","author":"Ertefai","year":"2010","journal-title":"Geochim Cosmochim Acta"},{"key":"2026042501113680300_ref48","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.01814-19","article-title":"Anaerobic degradation of non-methane alkanes by \u201cCandidatus Methanoliparia\u201d in hydrocarbon seeps of the Gulf of Mexico","volume":"10","author":"Laso-P\u00e9rez","year":"2019","journal-title":"mBio"},{"key":"2026042501113680300_ref49","author":"Bushnell","year":"2014"},{"key":"2026042501113680300_ref50","doi-asserted-by":"publisher","first-page":"e102","DOI":"10.1002\/cpbi.102","article-title":"Using SPAdes De novo assembler","volume":"70","author":"Prjibelski","year":"2020","journal-title":"Curr Protoc Bioinformatics"},{"key":"2026042501113680300_ref51","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1186\/2049-2618-2-26","article-title":"MaxBin: an automated binning method to recover individual genomes from metagenomes using an expectation-maximization algorithm","volume":"2","author":"Wu","year":"2014","journal-title":"Microbiome."},{"key":"2026042501113680300_ref52","doi-asserted-by":"publisher","first-page":"e7359","DOI":"10.7717\/peerj.7359","article-title":"MetaBAT 2: an adaptive binning algorithm for robust and efficient genome reconstruction from metagenome assemblies","volume":"7","author":"Kang","year":"2019","journal-title":"PeerJ."},{"key":"2026042501113680300_ref53","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1038\/s41564-020-00834-3","article-title":"Community-led, integrated, reproducible multi-omics with anvi\u2019o","volume":"6","author":"Eren","year":"2021","journal-title":"Nat Microbiol"},{"key":"2026042501113680300_ref54","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1038\/s41564-022-01297-4","article-title":"Evolutionary diversification of methanotrophic ANME-1 archaea and their expansive virome","volume":"8","author":"Laso-P\u00e9rez","year":"2023","journal-title":"Nat Microbiol"},{"key":"2026042501113680300_ref55","doi-asserted-by":"crossref","DOI":"10.1101\/gr.186072.114","article-title":"CheckM: assessing the quality of microbial genomes recovered from isolates, single cells, and metagenomes","volume":"25","author":"Parks","year":"2015","journal-title":"Genome Res"},{"key":"2026042501113680300_ref56"},{"key":"2026042501113680300_ref57","article-title":"CoverM: rRead alignment statistics for metagenomics","volume-title":"Bioinformatics","author":"Aroney","year":"2025"},{"key":"2026042501113680300_ref58","doi-asserted-by":"publisher","first-page":"2068","DOI":"10.1093\/bioinformatics\/btu153","article-title":"Prokka: rapid prokaryotic genome annotation","volume":"30","author":"Seemann","year":"2014","journal-title":"Bioinformatics."},{"key":"2026042501113680300_ref59","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1038\/nmeth.3176","article-title":"Fast and sensitive protein alignment using DIAMOND","volume":"12","author":"Buchfink","year":"2015","journal-title":"Nat Methods"},{"key":"2026042501113680300_ref60","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1093\/nar\/28.1.33","article-title":"The COG database: a tool for genome-scale analysis of protein functions and evolution","volume":"28","author":"Tatusov","year":"2000","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref61","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1186\/1745-6150-2-33","article-title":"Clusters of orthologous genes for 41 archaeal genomes and implications for evolutionary genomics of archaea","volume":"2","author":"Makarova","year":"2007","journal-title":"Biol Direct"},{"key":"2026042501113680300_ref62","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1093\/nar\/28.1.27","article-title":"KEGG: Kyoto Encyclopedia of genes and genomes","volume":"28","author":"Kanehisa","year":"2000","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref63","doi-asserted-by":"publisher","first-page":"D227","DOI":"10.1093\/nar\/gkp971","article-title":"MEROPS: the peptidase database","volume":"38","author":"Rawlings","year":"2010","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref64","doi-asserted-by":"publisher","first-page":"W200","DOI":"10.1093\/nar\/gky448","article-title":"HMMER web server: 2018 update","volume":"46","author":"Potter","year":"2018","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref65","doi-asserted-by":"publisher","first-page":"138D","DOI":"10.1093\/nar\/gkh121","article-title":"The Pfam protein families database","volume":"32","author":"Bateman","year":"2004","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref66","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1093\/nar\/gkg128","article-title":"The TIGRFAMs database of protein families","volume":"31","author":"Haft","year":"2003","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref67","doi-asserted-by":"publisher","first-page":"34212","DOI":"10.1038\/srep34212","article-title":"HydDB: a web tool for hydrogenase classification and analysis","volume":"6","author":"S\u00f8ndergaard","year":"2016","journal-title":"Sci Rep"},{"key":"2026042501113680300_ref68","doi-asserted-by":"publisher","first-page":"1312","DOI":"10.1093\/bioinformatics\/btu033","article-title":"RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies","volume":"30","author":"Stamatakis","year":"2014","journal-title":"Bioinformatics."},{"key":"2026042501113680300_ref69","doi-asserted-by":"publisher","first-page":"1160","DOI":"10.1093\/bib\/bbx108","article-title":"MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization","volume":"20","author":"Katoh","year":"2019","journal-title":"Brief Bioinform"},{"key":"2026042501113680300_ref70","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1186\/s12859-023-05311-2","article-title":"CAGECAT: the CompArative GEne cluster analysis toolbox for rapid search and visualisation of homologous gene clusters","volume":"24","author":"van den Belt","year":"2023","journal-title":"BMC Bioinformatics"},{"key":"2026042501113680300_ref71","doi-asserted-by":"publisher","first-page":"3150","DOI":"10.1093\/bioinformatics\/bts565","article-title":"CD-HIT: accelerated for clustering the next-generation sequencing data","volume":"28","author":"Fu","year":"2012","journal-title":"Bioinformatics."},{"key":"2026042501113680300_ref72","doi-asserted-by":"publisher","first-page":"22505","DOI":"10.1073\/pnas.1909306116","article-title":"GDGT cyclization proteins identify the dominant archaeal sources of tetraether lipids in the ocean","volume":"116","author":"Zeng","year":"2019","journal-title":"Proc Natl Acad Sci"},{"key":"2026042501113680300_ref73","doi-asserted-by":"publisher","first-page":"1792","DOI":"10.1093\/nar\/gkh340","article-title":"MUSCLE: multiple sequence alignment with high accuracy and high throughput","volume":"32","author":"Edgar","year":"2004","journal-title":"Nucleic Acids Res"},{"key":"2026042501113680300_ref74","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1002\/rcm.1378","article-title":"Intact polar membrane lipids in prokaryotes and sediments deciphered by high-performance liquid chromatography\/electrospray ionization multistage mass spectrometry\u2014new biomarkers for biogeochemistry and microbial ecology","volume":"18","author":"Sturt","year":"2004","journal-title":"Rapid Commun Mass Spectrom"},{"key":"2026042501113680300_ref75","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1080\/01490450303894","article-title":"Characterization of specific membrane fatty acids as chemotaxonomic markers for sulfate-reducing bacteria involved in anaerobic oxidation of methane","volume":"20","author":"Elvert","year":"2003","journal-title":"Geomicrobiol J"},{"key":"2026042501113680300_ref76","doi-asserted-by":"publisher","first-page":"335","DOI":"10.5194\/bg-2-335-2005","article-title":"Methane emission and consumption at a North Sea gas seep (Tommeliten area)","volume":"2","author":"Niemann","year":"2005","journal-title":"Biogeosciences."},{"key":"2026042501113680300_ref77","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.orggeochem.2014.02.005","article-title":"Identification of isoprenoid glycosidic glycerol dibiphytanol diethers and indications for their biosynthetic origin","volume":"69","author":"Meador","year":"2014","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref78","doi-asserted-by":"publisher","first-page":"2817","DOI":"10.1002\/rcm.4707","article-title":"Intramolecular stable carbon isotopic analysis of archaeal glycosyl tetraether lipids","volume":"24","author":"Lin","year":"2010","journal-title":"Rapid Commun Mass Spectrom"},{"key":"2026042501113680300_ref79","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/femsec\/fiz131","article-title":"Substrate characteristic bacterial fatty acid production based on amino acid assimilation and transformation in marine sediments","volume":"95","author":"Aepfler","year":"2019","journal-title":"FEMS Microbiol Ecol"},{"key":"2026042501113680300_ref80","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.orggeochem.2013.03.004","article-title":"Application of two new LC\u2013ESI\u2013MS methods for improved detection of intact polar lipids (IPLs) in environmental samples","volume":"59","author":"W\u00f6rmer","year":"2013","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref81","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.orggeochem.2013.05.007","article-title":"An improved method for the analysis of archaeal and bacterial ether core lipids","volume":"61","author":"Becker","year":"2013","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref82","doi-asserted-by":"publisher","first-page":"519","DOI":"10.1038\/s41586-019-1916-6","article-title":"Isolation of an archaeon at the prokaryote\u2013eukaryote interface","volume":"577","author":"Imachi","year":"2020","journal-title":"Nature."},{"key":"2026042501113680300_ref83","doi-asserted-by":"publisher","first-page":"991","DOI":"10.1038\/nature07174","article-title":"Significant contribution of archaea to extant biomass in marine subsurface sediments","volume":"454","author":"Lipp","year":"2008","journal-title":"Nature."},{"key":"2026042501113680300_ref84","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/j.orggeochem.2016.02.004","article-title":"Tracing the production and fate of individual archaeal intact polar lipids using stable isotope probing","volume":"95","author":"Kellermann","year":"2016","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref85","doi-asserted-by":"publisher","first-page":"1545","DOI":"10.1038\/s41467-022-29264-x","article-title":"Identification of a protein responsible for the synthesis of archaeal membrane-spanning GDGT lipids","volume":"13","author":"Zeng","year":"2022","journal-title":"Nat Commun"},{"key":"2026042501113680300_ref86","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1038\/s41586-022-05120-2","article-title":"Discovery, structure and mechanism of a tetraether lipid synthase","volume":"609","author":"Lloyd","year":"2022","journal-title":"Nature."},{"key":"2026042501113680300_ref87","doi-asserted-by":"publisher","first-page":"11111","DOI":"10.1073\/pnas.0401188101","article-title":"Membrane lipid patterns typify distinct anaerobic methanotrophic consortia","volume":"101","author":"Blumenberg","year":"2004","journal-title":"Proc Natl Acad Sci"},{"key":"2026042501113680300_ref88","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.margeo.2005.02.023","article-title":"Biomarker and 16S rDNA evidence for anaerobic oxidation of methane and related carbonate precipitation in deep-sea mud volcanoes of the Sorokin trough, Black Sea","volume":"217","author":"Stadnitskaia","year":"2005","journal-title":"Mar Geol"},{"key":"2026042501113680300_ref89","doi-asserted-by":"publisher","first-page":"1668","DOI":"10.1016\/j.orggeochem.2007.11.003","article-title":"Diagnostic lipid biomarker and stable carbon isotope signatures of microbial communities mediating the anaerobic oxidation of methane with sulphate","volume":"39","author":"Niemann","year":"2008","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref90","doi-asserted-by":"publisher","first-page":"113349","DOI":"10.1016\/j.palaeo.2025.113349","article-title":"Sulfate-driven anaerobic oxidation of methane recorded by lower cretaceous (Barremian) authigenic carbonate deposits from the Kuhnpasset beds of Wollaston Forland, Northeast Greenland","volume":"680","author":"Wang","year":"2025","journal-title":"Palaeogeogr Palaeoclimatol Palaeoecol"},{"key":"2026042501113680300_ref91","doi-asserted-by":"crossref","first-page":"006435","DOI":"10.1099\/ijsem.0.006435","article-title":"Promethearchaeum syntrophicum gen. Nov., sp. nov., an anaerobic, obligately syntrophic archaeon, the first isolate of the lineage \u2018Asgard\u2019 archaea, and proposal of the new archaeal phylum Promethearchaeota phyl. Nov. and kingdom Promethearchaeati regn. Nov","volume":"74","author":"Imachi","year":"2024","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026042501113680300_ref92","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1111\/j.1472-4669.2005.00051.x","article-title":"Spatial variations of methanotrophic consortia at cold methane seeps: implications from a high-resolution molecular and isotopic approach","volume":"3","author":"Elvert","year":"2005","journal-title":"Geobiology."},{"key":"2026042501113680300_ref93","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1016\/S0146-6380(01)00015-8","article-title":"Three series of non-isoprenoidal dialkyl glycerol diethers in cold-seep carbonate crusts","volume":"32","author":"Pancost","year":"2001","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref94","doi-asserted-by":"publisher","first-page":"3157","DOI":"10.1128\/AEM.03794-14","article-title":"Mono- and Dialkyl glycerol ether lipids in anaerobic bacteria: biosynthetic insights from the mesophilic Sulfate reducer Desulfatibacillum alkenivorans PF2803T","volume":"81","author":"Grossi","year":"2015","journal-title":"Appl Environ Microbiol"},{"key":"2026042501113680300_ref95","doi-asserted-by":"publisher","first-page":"1685","DOI":"10.1016\/S0146-6380(00)00106-6","article-title":"Molecular and isotopic analysis of anaerobic methane-oxidizing communities in marine sediments","volume":"31","author":"Hinrichs","year":"2000","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref96","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/j.orggeochem.2016.05.015","article-title":"The alkyl glycerol ether lipid composition of heterotrophic sulfate reducing bacteria strongly depends on growth substrate","volume":"98","author":"Vin\u00e7on-Laugier","year":"2016","journal-title":"Org Geochem"},{"key":"2026042501113680300_ref97","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1111\/j.1462-2920.2006.01127.x","article-title":"In vitro cell growth of marine archaeal-bacterial consortia during anaerobic oxidation of methane with sulfate","volume":"9","author":"Nauhaus","year":"2007","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref98","doi-asserted-by":"publisher","first-page":"e70213","DOI":"10.1111\/1462-2920.70213","article-title":"Marine cold seep ANME-2\/SRB consortia produce their lipid biomass from inorganic carbon","volume":"27","author":"Stock","year":"2025","journal-title":"Environ Microbiol"},{"key":"2026042501113680300_ref99","doi-asserted-by":"publisher","first-page":"912299","DOI":"10.3389\/fmicb.2022.912299","article-title":"Activity of ancillary heterotrophic community members in anaerobic methane-oxidizing cultures","volume":"13","author":"Zhu","year":"2022","journal-title":"Front Microbiol"},{"key":"2026042501113680300_ref100","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/S0304-4203(00)00099-2","article-title":"Molecular signals for anaerobic methane oxidation in Black Sea seep carbonates and a microbial mat","volume":"73","author":"Thiel","year":"2001","journal-title":"Mar Chem"},{"key":"2026042501113680300_ref101","doi-asserted-by":"publisher","first-page":"596","DOI":"10.1038\/s41597-023-02521-4","article-title":"A comprehensive genomic catalog from global cold seeps","volume":"10","author":"Han","year":"2023","journal-title":"Sci Data"},{"key":"2026042501113680300_ref102","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/j.tim.2012.10.003","article-title":"Integrative mobile elements exploiting Xer recombination","volume":"21","author":"Das","year":"2013","journal-title":"Trends Microbiol"},{"key":"2026042501113680300_ref103","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/978-1-0716-3515-5_1","volume-title":"Viral Metagenomics: Methods and Protocols","author":"Zhou","year":"2024"},{"key":"2026042501113680300_ref104","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1111\/nyas.12675","article-title":"Single-stranded DNA viruses employ a variety of mechanisms for integration into host genomes","volume":"1341","author":"Krupovic","year":"2015","journal-title":"Ann N Y Acad Sci"},{"key":"2026042501113680300_ref105","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1146\/annurev-micro-041020-024935","article-title":"Diversity and evolution of methane-related pathways in archaea","volume":"76","author":"Garcia","year":"2022","journal-title":"Ann Rev Microbiol"},{"key":"2026042501113680300_ref106","doi-asserted-by":"publisher","first-page":"eabj1453","DOI":"10.1126\/sciadv.abj1453","article-title":"A methylotrophic origin of methanogenesis and early divergence of anaerobic multicarbon alkane metabolism","volume":"7","author":"Wang","year":"2021","journal-title":"Sci Adv"},{"key":"2026042501113680300_ref107","doi-asserted-by":"publisher","first-page":"e03490","DOI":"10.1128\/spectrum.03490-23","article-title":"Potential coupling of microbial methane, nitrogen, and Sulphur cycling in the Okinawa trough cold seep sediments","volume":"12","author":"Chen","year":"2024","journal-title":"Microbiol Spectrum"},{"key":"2026042501113680300_ref108","doi-asserted-by":"publisher","first-page":"46","DOI":"10.3389\/fmicb.2016.00046","article-title":"Metabolic capabilities of microorganisms involved in and associated with the anaerobic oxidation of methane","volume":"7","author":"Wegener","year":"2016","journal-title":"Front Microbiol"},{"key":"2026042501113680300_ref109","doi-asserted-by":"publisher","first-page":"5485","DOI":"10.1099\/ijsem.0.001545","article-title":"Description of \u2018Candidatus Marispirochaeta associata\u2019 and reclassification of Spirochaeta bajacaliforniensis, Spirochaeta smaragdinae and Spirochaeta sinaica to a new genus Sediminispirochaeta gen. Nov. as Sediminispirochaeta bajacaliforniensis comb. nov., Sediminispirochaeta smaragdinae comb. nov. and Sediminispirochaeta sinaica comb. nov","volume":"66","author":"Shivani","year":"2016","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026042501113680300_ref110","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1007\/BF00491897","article-title":"Spirochaeta bajacaliforniensis sp. n. from a microbial mat community at Laguna Figueroa, Baja California Norte, Mexico","volume":"142","author":"Fracek","year":"1985","journal-title":"Arch Microbiol"},{"key":"2026042501113680300_ref111","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1111\/j.1574-6968.1997.tb13876.x","article-title":"Spirochaeta smaragdinae sp. nov., a new mesophilic strictly anaerobic spirochete from an oil field","volume":"155","author":"Magot","year":"1997","journal-title":"FEMS Microbiol Lett"},{"key":"2026042501113680300_ref112","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1016\/j.biortech.2018.03.018","article-title":"Enrichment of sulfate reducing anaerobic methane oxidizing community dominated by ANME-1 from Ginsburg mud volcano (gulf of Cadiz) sediment in a biotrickling filter","volume":"259","author":"Bhattarai","year":"2018","journal-title":"Bioresour Technol"},{"key":"2026042501113680300_ref113","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1038\/nrmicro3260","article-title":"A re-evaluation of the archaeal membrane lipid biosynthetic pathway","volume":"12","author":"Villanueva","year":"2014","journal-title":"Nat Rev Microbiol"},{"key":"2026042501113680300_ref114","doi-asserted-by":"publisher","first-page":"1696","DOI":"10.1038\/ismej.2015.233","article-title":"Genomic reconstruction of a novel, deeply branched sediment archaeal phylum with pathways for acetogenesis and sulfur reduction","volume":"10","author":"Seitz","year":"2016","journal-title":"The ISME Journal."},{"key":"2026042501113680300_ref115","doi-asserted-by":"publisher","first-page":"2815","DOI":"10.1073\/pnas.0511033103","article-title":"Biogeographical distribution and diversity of microbes in methane hydrate-bearing deep marine sediments on the Pacific Ocean margin","volume":"103","author":"Inagaki","year":"2006","journal-title":"Proc Natl Acad Sci"},{"key":"2026042501113680300_ref116","doi-asserted-by":"publisher","first-page":"16022","DOI":"10.1038\/npjbiofilms.2016.22","article-title":"Plasmid transfer in biofilms: a perspective on limitations and opportunities","volume":"2","author":"Stalder","year":"2016","journal-title":"NPJ Biofilms Microbiomes"},{"key":"2026042501113680300_ref117","doi-asserted-by":"publisher","DOI":"10.1093\/femsec\/fiaa031","article-title":"Biofilms: hot spots of horizontal gene transfer (HGT) in aquatic environments, with a focus on a new HGT mechanism","volume":"96","author":"Abe","year":"2020","journal-title":"FEMS Microbiol Ecol"},{"key":"2026042501113680300_ref118","doi-asserted-by":"publisher","first-page":"2025.07.24.666526","DOI":"10.1101\/2025.07.24.666526v1","article-title":"Phylogenomics of Asgard archaea reveals a unique blend of prokaryotic-like horizontal transfer and eukaryotic-like gene duplication","author":"Manzano-Morales","year":"2025","journal-title":"bioRxiv"},{"key":"2026042501113680300_ref119","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1038\/s41586-021-03494-3","article-title":"Expanded diversity of Asgard archaea and their relationships with eukaryotes","volume":"593","author":"Liu","year":"2021","journal-title":"Nature."}],"container-title":["ISME Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/ismecommun\/advance-article-pdf\/doi\/10.1093\/ismeco\/ycag072\/67483126\/ycag072.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismecommun\/article-pdf\/6\/1\/ycag072\/67483126\/ycag072.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismecommun\/article-pdf\/6\/1\/ycag072\/67483126\/ycag072.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,25]],"date-time":"2026-04-25T05:11:48Z","timestamp":1777093908000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/ismecommun\/article\/doi\/10.1093\/ismeco\/ycag072\/8537621"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1]]},"references-count":119,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,1,14]]}},"URL":"https:\/\/doi.org\/10.1093\/ismeco\/ycag072","relation":{"has-preprint":[{"id-type":"doi","id":"10.64898\/2026.01.09.698606","asserted-by":"object"}]},"ISSN":["2730-6151"],"issn-type":[{"value":"2730-6151","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1]]},"published":{"date-parts":[[2026,1]]},"article-number":"ycag072"}}