{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T18:57:28Z","timestamp":1781636248290,"version":"3.54.5"},"reference-count":75,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,2,24]],"date-time":"2026-02-24T00:00:00Z","timestamp":1771891200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,3,23]],"date-time":"2026-03-23T00:00:00Z","timestamp":1774224000000},"content-version":"vor","delay-in-days":27,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Deep-sea methane seeps fuel biodiverse habitats sustained by the release of hydrocarbon-rich fluids and associated microbial activity. Here, we describe the ecology of a seep-associated cold-water coral and provide evidence of its associations with chemosynthetic bacteria. High-resolution seafloor surveys revealed that the distribution of this coral was predominantly confined to actively seeping zones, and habitat suitability models confirmed that proximity to active seepage was an important factor influencing the coral\u2019s distribution. Stable carbon-isotope values were consistent with a nutritional strategy incorporating chemosynthetically derived carbon, likely as a supplement to suspension feeding on photosynthetically derived material. Microbial metabarcoding confirmed the presence of both thiotrophic and methanotrophic bacteria, including SUP05 and MMG-2 groups. Incubations with\n                    <jats:sup>13<\/jats:sup>\n                    C-labelled methane further revealed this species may also be capable of assimilating methane-derived carbon into its biomass. These findings provide new evidence of a previously underrecognized facultative symbiosis between cold-water corals and chemosymbiotic bacteria and suggest that these corals are not restricted to the periphery of seep habitats. Instead, they may exploit microbial associations, including contributions from both thiotrophic and potentially methanotrophic taxa, to persist in actively seeping areas.\n                  <\/jats:p>","DOI":"10.1038\/s41598-025-32153-0","type":"journal-article","created":{"date-parts":[[2026,2,24]],"date-time":"2026-02-24T04:35:13Z","timestamp":1771907713000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Microbially mediated carbon utilization by a cold-water coral inhabiting methane seeps"],"prefix":"10.1038","volume":"16","author":[{"given":"April","family":"Stabbins","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shana","family":"Goffredi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryan","family":"Gasbarro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katherine","family":"Dawson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John","family":"Magyar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amanda","family":"Glazier","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kelly","family":"Meinert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Victoria","family":"Orphan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Erik","family":"Cordes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,24]]},"reference":[{"issue":"10","key":"32153_CR1","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1038\/nrmicro1992","volume":"6","author":"N Dubilier","year":"2008","unstructured":"Dubilier, N., Bergin, C. & Lott, C. Symbiotic diversity in marine animals: The art of harnessing chemosynthesis. Nat. Rev. Microbiol. 6(10), 725\u2013740 (2008).","journal-title":"Nat. Rev. Microbiol."},{"issue":"14","key":"32153_CR2","doi-asserted-by":"crossref","first-page":"eaay8562","DOI":"10.1126\/sciadv.aay8562","volume":"6","author":"SK Goffredi","year":"2020","unstructured":"Goffredi, S. K. et al. Methanotrophic bacterial symbionts fuel dense populations of deep-sea feather duster worms (Sabellida, Annelida) and extend the spatial influence of methane seepage. Sci. Adv. 6(14), eaay8562 (2020).","journal-title":"Sci. Adv."},{"issue":"5","key":"32153_CR3","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1111\/j.1758-2229.2009.00081.x","volume":"1","author":"JM Petersen","year":"2009","unstructured":"Petersen, J. M. & Dubilier, N. Methanotrophic symbioses in marine invertebrates. Environ. Microbiol. Rep. 1(5), 319\u2013335 (2009).","journal-title":"Environ. Microbiol. Rep."},{"issue":"5","key":"32153_CR4","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1038\/s41396-019-0346-7","volume":"13","author":"M Rubin-Blum","year":"2019","unstructured":"Rubin-Blum, M. et al. Fueled by methane: deep-sea sponges from asphalt seeps gain their nutrition from methane-oxidizing symbionts. ISME J. 13(5), 1209\u20131225 (2019).","journal-title":"ISME J."},{"issue":"1","key":"32153_CR5","doi-asserted-by":"crossref","first-page":"11870","DOI":"10.1038\/ncomms11870","volume":"7","author":"T Thomas","year":"2016","unstructured":"Thomas, T. et al. Diversity, structure and convergent evolution of the global sponge microbiome. Nat. Commun. 7(1), 11870 (2016).","journal-title":"Nat. Commun."},{"issue":"9","key":"32153_CR6","doi-asserted-by":"crossref","first-page":"1465","DOI":"10.1038\/s41564-019-0475-9","volume":"4","author":"HR Gruber-Vodicka","year":"2019","unstructured":"Gruber-Vodicka, H. R. et al. Two intracellular and cell type-specific bacterial symbionts in the placozoan Trichoplax H2. Nat. Microbiol. 4(9), 1465\u20131474 (2019).","journal-title":"Nat. Microbiol."},{"issue":"1","key":"32153_CR7","first-page":"1","volume":"19","author":"SK Goffredi","year":"2021","unstructured":"Goffredi, S. K. et al. Mixotrophic chemosynthesis in a deep-sea anemone from hydrothermal vents in the Pescadero Basin, Gulf of California. BMC Biol. 19(1), 1\u201318 (2021).","journal-title":"BMC Biol."},{"issue":"1","key":"32153_CR8","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1111\/gcb.16447","volume":"29","author":"EO Osman","year":"2023","unstructured":"Osman, E. O. et al. Capacity of deep-sea corals to obtain nutrition from cold seeps aligned with microbiome reorganization. Glob. Change Biol. 29(1), 189\u2013205 (2023).","journal-title":"Glob. Change Biol."},{"issue":"11","key":"32153_CR9","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.1601122","volume":"2","author":"K Frankowiak","year":"2016","unstructured":"Frankowiak, K. et al. Photosymbiosis and the expansion of shallow-water corals. Sci. Adv. 2(11), e1601122 (2016).","journal-title":"Sci. Adv."},{"issue":"3","key":"32153_CR10","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1016\/S0012-8252(02)00104-6","volume":"60","author":"GD Stanley","year":"2003","unstructured":"Stanley, G. D. The evolution of modern corals and their early history. Earth-Sci. Rev. 60(3), 195\u2013225 (2003).","journal-title":"Earth-Sci. Rev."},{"issue":"3","key":"32153_CR11","first-page":"311","volume":"81","author":"SD Cairns","year":"2007","unstructured":"Cairns, S. D. Deep-water corals: An overview with special reference to diversity and distribution of deep-water scleractinian corals. Bull. Mar. Sci. 81(3), 311\u2013322 (2007).","journal-title":"Bull. Mar. Sci."},{"key":"32153_CR12","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1007\/3-540-27673-4_44","volume-title":"Cold-Water Corals and Ecosystems","author":"L Buhl-Mortensen","year":"2005","unstructured":"Buhl-Mortensen, L. & Mortensen, P. B. Distribution and diversity of species associated with deep-sea gorgonian corals off Atlantic Canada. In Cold-Water Corals and Ecosystems 849\u2013879 (Berlin, Heidelberg, Springer, Berlin Heidelberg, 2005)."},{"issue":"6","key":"32153_CR13","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1016\/j.dsr.2008.03.005","volume":"55","author":"EE Cordes","year":"2008","unstructured":"Cordes, E. E. et al. Coral communities of the deep Gulf of Mexico. Deep-Sea Res. Part Oceanogr. Res. Pap. 55(6), 777\u2013787 (2008).","journal-title":"Deep-Sea Res. Part Oceanogr. Res. Pap."},{"key":"32153_CR14","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1007\/3-540-27673-4_41","volume-title":"Cold-Water Corals and Ecosystems","author":"MJ Costello","year":"2005","unstructured":"Costello, M. J. et al. Role of cold-water Lophelia pertusa coral reefs as fish habitat in the NE Atlantic. In Cold-Water Corals and Ecosystems 771\u2013805 (Berlin, Heidelberg, Springer, Berlin Heidelberg, 2005)."},{"issue":"1","key":"32153_CR15","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1007\/s12526-021-01255-z","volume":"52","author":"PJ Etnoyer","year":"2022","unstructured":"Etnoyer, P. J. et al. Diversity and time-series analyses of Caribbean deep-sea coral and sponge assemblages on the tropical island slope of Isla de Roat\u00e1n, Honduras. Mar. Biodivers. 52(1), 8 (2022).","journal-title":"Mar. Biodivers."},{"key":"32153_CR16","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.dsr2.2013.05.023","volume":"99","author":"CL Doughty","year":"2014","unstructured":"Doughty, C. L., Quattrini, A. M. & Cordes, E. E. Insights into the population dynamics of the deep-sea coral genus Paramuricea in the Gulf of Mexico. Deep Sea Res. Part II Top. Stud. Oceanogr. 99, 71\u201382 (2014).","journal-title":"Deep Sea Res. Part II Top. Stud. Oceanogr."},{"issue":"23","key":"32153_CR17","doi-asserted-by":"crossref","first-page":"7108","DOI":"10.1111\/gcb.16415","volume":"28","author":"R Gasbarro","year":"2022","unstructured":"Gasbarro, R., Sowers, D., Margolin, A. & Cordes, E. E. Distribution and predicted climatic refugia for a reef-building cold-water coral on the southeast US margin. Glob Change Biol. 28(23), 7108\u20137125 (2022).","journal-title":"Glob Change Biol."},{"key":"32153_CR18","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1525\/elementa.349","volume":"7","author":"F Girard","year":"2019","unstructured":"Girard, F., Cruz, R., Glickman, O., Harpster, T. & Fisher, C. R. In situ growth of deep-sea octocorals after the Deepwater Horizon oil spill. Elem Sci. Anthr. 7, 12 (2019).","journal-title":"Elem Sci. Anthr."},{"key":"32153_CR19","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.biocon.2018.06.028","volume":"225","author":"F Girard","year":"2018","unstructured":"Girard, F. & Fisher, C. R. Long-term impact of the Deepwater Horizon oil spill on deep-sea corals detected after seven years of monitoring. Biol. Conserv. 225, 117\u2013127 (2018).","journal-title":"Biol. Conserv."},{"issue":"6","key":"32153_CR20","doi-asserted-by":"crossref","first-page":"e65394","DOI":"10.1371\/journal.pone.0065394","volume":"8","author":"M Lacharit\u00e9","year":"2013","unstructured":"Lacharit\u00e9, M. & Metaxas, A. Early life history of deep-water gorgonian corals may limit their abundance. PLoS ONE 8(6), e65394 (2013).","journal-title":"PLoS ONE"},{"issue":"4","key":"32153_CR21","doi-asserted-by":"crossref","first-page":"2181","DOI":"10.1111\/gcb.14996","volume":"26","author":"T Morato","year":"2020","unstructured":"Morato, T. et al. Climate-induced changes in the suitable habitat of cold-water corals and commercially important deep-sea fishes in the North Atlantic. Glob Change Biol. 26(4), 2181\u20132202 (2020).","journal-title":"Glob Change Biol."},{"issue":"129","key":"32153_CR22","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1016\/j.dsr2.2014.10.021","volume":"1","author":"NG Prouty","year":"2016","unstructured":"Prouty, N. G., Fisher, C. R., Demopoulos, A. W. J. & Druffel, E. R. M. Growth rates and ages of deep-sea corals impacted by the Deepwater Horizon oil spill. Deep Sea. Res. Part II Top. Stud. Oceanogr. 1(129), 196\u2013212 (2016).","journal-title":"Deep Sea. Res. Part II Top. Stud. Oceanogr."},{"issue":"8","key":"32153_CR23","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0022588","volume":"6","author":"E Ramirez-Llodra","year":"2011","unstructured":"Ramirez-Llodra, E. et al. Man and the last great wilderness: Human impact on the deep sea. PLoS ONE 6(8), e22588 (2011).","journal-title":"PLoS ONE"},{"issue":"7","key":"32153_CR24","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1038\/nclimate2611","volume":"5","author":"RE Thresher","year":"2015","unstructured":"Thresher, R. E., Guinotte, J. M., Matear, R. J. & Hobday, A. J. Options for managing impacts of climate change on a deep-sea community. Nat. Clim. Change 5(7), 635\u2013639 (2015).","journal-title":"Nat. Clim. Change"},{"issue":"10","key":"32153_CR25","doi-asserted-by":"crossref","first-page":"e76869","DOI":"10.1371\/journal.pone.0076869","volume":"8","author":"DA Bowden","year":"2013","unstructured":"Bowden, D. A. et al. Cold seep epifaunal communities on the hikurangi margin, New Zealand: Composition, succession, and vulnerability to human activities. PLoS ONE 8(10), e76869 (2013).","journal-title":"PLoS ONE"},{"issue":"10","key":"32153_CR26","doi-asserted-by":"crossref","first-page":"e0139904","DOI":"10.1371\/journal.pone.0139904","volume":"10","author":"AM Quattrini","year":"2015","unstructured":"Quattrini, A. M. et al. Exploration of the canyon-incised continental margin of the Northeastern United States reveals dynamic habitats and diverse communities. PLoS ONE 10(10), e0139904 (2015).","journal-title":"PLoS ONE"},{"issue":"5","key":"32153_CR27","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1016\/j.dsr.2008.12.006","volume":"56","author":"EL Becker","year":"2009","unstructured":"Becker, E. L., Cordes, E. E., Macko, S. A. & Fisher, C. R. Importance of seep primary production to Lophelia pertusa and associated fauna in the Gulf of Mexico. Deep-Sea Res. Part Oceanogr. Res. Pap. 56(5), 786\u2013800 (2009).","journal-title":"Deep-Sea Res. Part Oceanogr. Res. Pap."},{"issue":"3","key":"32153_CR28","doi-asserted-by":"crossref","first-page":"407","DOI":"10.11646\/zootaxa.4671.3.6","volume":"4671","author":"O Breedy","year":"2019","unstructured":"Breedy, O., Rouse, G. W., Stabbins, A., Cort\u00e9s, J. & Cordes, E. E. New records of Swiftia (Cnidaria, Anthozoa, Octocorallia) from off the Pacific Costa Rican margin, including a new species from methane seeps. Zootaxa 4671(3), 407\u2013419 (2019).","journal-title":"Zootaxa"},{"issue":"19\u201320","key":"32153_CR29","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1016\/j.dsr2.2009.05.002","volume":"56","author":"KL Erickson","year":"2009","unstructured":"Erickson, K. L., Macko, S. A. & Van Dover, C. L. Evidence for a chemoautotrophically based food web at inactive hydrothermal vents (Manus Basin). Deep Sea Res. Part II Top. Stud. Oceanogr. 56(19\u201320), 1577\u20131585 (2009).","journal-title":"Deep Sea Res. Part II Top. Stud. Oceanogr."},{"issue":"15","key":"32153_CR30","doi-asserted-by":"crossref","first-page":"4123","DOI":"10.1111\/mec.12370","volume":"22","author":"AM Quattrini","year":"2013","unstructured":"Quattrini, A. M. et al. Niche divergence by deep-sea octocorals in the genus Callogorgia across the continental slope of the Gulf of Mexico. Mol. Ecol. 22(15), 4123\u20134140 (2013).","journal-title":"Mol. Ecol."},{"issue":"4","key":"32153_CR31","doi-asserted-by":"crossref","first-page":"361","DOI":"10.2307\/3514822","volume":"7","author":"CK Paull","year":"1992","unstructured":"Paull, C. K. et al. Indicators of methane-derived carbonates and chemosynthetic organic carbon deposits: Examples from the florida escarpment. Palaios 7(4), 361\u2013375 (1992).","journal-title":"Palaios"},{"key":"32153_CR32","doi-asserted-by":"publisher","DOI":"10.1101\/2020.02.27.968453","author":"SA Vohsen","year":"2020","unstructured":"Vohsen, S. A. et al. Deep-sea corals near cold seeps associate with chemoautotrophic bacteria that are related to the symbionts of cold seep and hydrothermal vent mussels. BioRxiv https:\/\/doi.org\/10.1101\/2020.02.27.968453 (2020).","journal-title":"BioRxiv"},{"issue":"1","key":"32153_CR33","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.epsl.2005.03.009","volume":"235","author":"P Linke","year":"2005","unstructured":"Linke, P., Wallmann, K., Suess, E., Hensen, C. & Rehder, G. In situ benthic fluxes from an intermittently active mud volcano at the Costa Rica convergent margin. Earth Planet Sci. Lett. 235(1), 79\u201395 (2005).","journal-title":"Earth Planet Sci. Lett."},{"key":"32153_CR34","first-page":"49","volume-title":"Martinelli Mud Volcanoes","author":"T Moerz","year":"2005","unstructured":"Moerz, T. et al. Styles and Productivity of Mud Diapirism along the Middle American Margin. In Martinelli Mud Volcanoes 49\u201376 (Springer, Geodynamics and Seismicity. Berlin\/Heidelberg, 2005)."},{"issue":"7","key":"32153_CR35","doi-asserted-by":"crossref","first-page":"1845","DOI":"10.1007\/s00531-014-1045-2","volume":"103","author":"V Liebetrau","year":"2014","unstructured":"Liebetrau, V. et al. Cold-seep-driven carbonate deposits at the Central American forearc: contrasting evolution and timing in escarpment and mound settings. Int. J. Earth Sci. 103(7), 1845\u20131872 (2014).","journal-title":"Int. J. Earth Sci."},{"key":"32153_CR36","first-page":"36","volume":"1","author":"S Kutterolf","year":"2008","unstructured":"Kutterolf, S. et al. Lifetime and cyclicity of fluid venting at forearc mound structures determined by tephrostratigraphy and radiometric dating of authigenic carbonates. Geology 1, 36 (2008).","journal-title":"Geology"},{"issue":"3","key":"32153_CR37","doi-asserted-by":"crossref","first-page":"94","DOI":"10.3390\/geosciences8030094","volume":"8","author":"S Walbridge","year":"2018","unstructured":"Walbridge, S., Slocum, N., Pobuda, M. & Wright, D. Unified geomorphological analysis workflows with benthic terrain modeler. Geosciences 8(3), 94 (2018).","journal-title":"Geosciences"},{"key":"32153_CR38","doi-asserted-by":"crossref","first-page":"145","DOI":"10.3354\/meps10816","volume":"506","author":"SE Georgian","year":"2014","unstructured":"Georgian, S. E., Shedd, W. & Cordes, E. E. High-resolution ecological niche modelling of the cold-water coral Lophelia pertusa in the Gulf of Mexico. Mar. Ecol. Prog. Ser. 506, 145\u2013161 (2014).","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"32153_CR39","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1146\/annurev.marine.010908.163912","volume":"1","author":"EE Cordes","year":"2009","unstructured":"Cordes, E. E., Bergquist, D. C. & Fisher, C. R. Macro-ecology of gulf of Mexico cold seeps. Annu. Rev. Mar. Sci. 1, 143\u2013168 (2009).","journal-title":"Annu. Rev. Mar. Sci."},{"issue":"5","key":"32153_CR40","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1111\/ddi.12892","volume":"25","author":"T Hao","year":"2019","unstructured":"Hao, T., Elith, J., Guillera-Arroita, G. & Lahoz-Monfort, J. J. A review of evidence about use and performance of species distribution modelling ensembles like BIOMOD. Divers. Distrib. 25(5), 839\u2013852 (2019).","journal-title":"Divers. Distrib."},{"issue":"1","key":"32153_CR41","doi-asserted-by":"publisher","first-page":"e014485","DOI":"10.1002\/ecm.1486","volume":"92","author":"R Valavi","year":"2021","unstructured":"Valavi, R., Guillera-Arroita, G., Lahoz-Monfort, J. J. & Elith, J. Predictive performance of presence-only species distribution models: a benchmark study with reproducible code. Ecol. Monogr. 92(1), e014485. https:\/\/doi.org\/10.1002\/ecm.1486 (2021).","journal-title":"Ecol. Monogr."},{"issue":"1","key":"32153_CR42","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1146\/annurev.ecolsys.110308.120159","volume":"40","author":"J Elith","year":"2009","unstructured":"Elith, J. & Leathwick, J. R. Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40(1), 677\u2013697 (2009).","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"32153_CR43","doi-asserted-by":"publisher","DOI":"10.1214\/ss\/1009213726.full","author":"L Breiman","year":"2001","unstructured":"Breiman, L. Statistical modeling: The two cultures (with comments and a rejoinder by the author). Stat. Sci. https:\/\/doi.org\/10.1214\/ss\/1009213726.full (2001).","journal-title":"Stat. Sci."},{"issue":"2","key":"32153_CR44","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1111\/2041-210X.13107","volume":"10","author":"R Valavi","year":"2019","unstructured":"Valavi, R., Elith, J., Lahoz-Monfort, J. J. & Guillera-Arroita, G. block CV\u202f: An r package for generating spatially or environmentally separated folds for k -fold cross-validation of species distribution models. Methods Ecol Evol. 10(2), 225\u2013232 (2019).","journal-title":"Methods Ecol Evol."},{"key":"32153_CR45","unstructured":"R Core Team. R: A Language and Environment for Statistical Computing [Internet]. Vienna, Austria: R Foundation for Statistical Computing. https:\/\/www.R-project.org\/. (2023)."},{"key":"32153_CR46","unstructured":"Thullier W, Georges D, Maya G, Engler R, Breiner F. biomod2: Ensemble platform for species distribution modeling. R package version 3.5.1. (2020)."},{"issue":"1","key":"32153_CR47","doi-asserted-by":"crossref","first-page":"4516","DOI":"10.1073\/pnas.1000080107","volume":"108","author":"JG Caporaso","year":"2011","unstructured":"Caporaso, J. G. et al. Global patterns of 16S rRNA diversity at a depth of millions of sequences per sample. Proc. Natl. Acad. Sci. 108(1), 4516\u20134522 (2011).","journal-title":"Proc. Natl. Acad. Sci."},{"issue":"5","key":"32153_CR48","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1038\/nmeth.f.303","volume":"7","author":"JG Caporaso","year":"2010","unstructured":"Caporaso, J. G. et al. QIIME allows analysis of high-throughput community sequencing data. Nat. Methods 7(5), 335\u2013336 (2010).","journal-title":"Nat. Methods"},{"key":"32153_CR49","doi-asserted-by":"crossref","first-page":"147","DOI":"10.3354\/meps208147","volume":"208","author":"Y Fujiwara","year":"2000","unstructured":"Fujiwara, Y. et al. Phylogenetic characterization of endosymbionts in three hydrothermal vent mussels: Influence on host distributions. Mar. Ecol. Prog. Ser. 208, 147\u2013155 (2000).","journal-title":"Mar. Ecol. Prog. Ser."},{"issue":"12","key":"32153_CR50","doi-asserted-by":"crossref","first-page":"3608","DOI":"10.1111\/1462-2920.12379","volume":"16","author":"KM Fontanez","year":"2014","unstructured":"Fontanez, K. M. & Cavanaugh, C. M. Evidence for horizontal transmission from multilocus phylogeny of deep-sea mussel (Mytilidae) symbionts. Environ. Microbiol. 16(12), 3608\u20133621 (2014).","journal-title":"Environ. Microbiol."},{"issue":"1970","key":"32153_CR51","doi-asserted-by":"crossref","first-page":"20212137","DOI":"10.1098\/rspb.2021.2137","volume":"289","author":"M Perez","year":"2022","unstructured":"Perez, M. et al. Divergent paths in the evolutionary history of maternally transmitted clam symbionts. Proc. R Soc. B Biol. Sci. 289(1970), 20212137 (2022).","journal-title":"Proc. R Soc. B Biol. Sci."},{"key":"32153_CR52","doi-asserted-by":"crossref","first-page":"141","DOI":"10.3354\/meps254141","volume":"254","author":"M Ribes","year":"2003","unstructured":"Ribes, M., Coma, R. & Rossi, S. Natural feeding of the temperate asymbiotic octocoral-gorgonian Leptogorgia sarmentosa (Cnidaria: Octocorallia). Mar. Ecol. Prog. Ser. 254, 141\u2013150 (2003).","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"32153_CR53","doi-asserted-by":"publisher","DOI":"10.1029\/2008GC001978","author":"H Sahling","year":"2008","unstructured":"Sahling, H. et al. Fluid seepage at the continental margin offshore Costa Rica and southern Nicaragua. Geochem. Geophys. Geosyst. https:\/\/doi.org\/10.1029\/2008GC001978 (2008).","journal-title":"Geochem. Geophys. Geosyst."},{"issue":"19","key":"32153_CR54","doi-asserted-by":"crossref","first-page":"R1137","DOI":"10.1016\/j.cub.2020.07.050","volume":"30","author":"EM Sogin","year":"2020","unstructured":"Sogin, E. M., Leisch, N. & Dubilier, N. Chemosynthetic symbioses. Curr. Biol. 30(19), R1137\u2013R1142 (2020).","journal-title":"Curr. Biol."},{"key":"32153_CR55","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1111\/1758-2229.12909","volume":"13","author":"C Brzechffa","year":"2021","unstructured":"Brzechffa, C. & Goffredi, S. K. Contrasting influences on bacterial symbiont specificity by co-occurring deep-sea mussels and tubeworms. Environ. Microbiol. Rep. 13, 104\u2013111 (2021).","journal-title":"Environ. Microbiol. Rep."},{"issue":"1992","key":"32153_CR56","first-page":"337","volume":"30","author":"JJ Childress","year":"1992","unstructured":"Childress, J. J. & Fisher, C. R. The biology of hydrothermal vent animals: Physiology, biochemistry, and autotrophic symbioses. Oceanogr. Mar. Biol. Annu. Rev. 30(1992), 337\u2013441 (1992).","journal-title":"Oceanogr. Mar. Biol. Annu. Rev."},{"issue":"1\/2","key":"32153_CR57","doi-asserted-by":"crossref","first-page":"45","DOI":"10.3354\/meps103045","volume":"103","author":"CR Fisher","year":"1994","unstructured":"Fisher, C. R., Childress, J. J., Macko, S. A. & Brooks, J. M. Nutritional interactions in Galapagos Rift hydrothermal vent communities: Inferences from stable carbon and nitrogen isotope analyses. Mar. Ecol. Prog. Ser. 103(1\/2), 45\u201355 (1994).","journal-title":"Mar. Ecol. Prog. Ser."},{"issue":"5","key":"32153_CR58","doi-asserted-by":"crossref","first-page":"1336","DOI":"10.4319\/lo.2002.47.5.1336","volume":"47","author":"LA Levin","year":"2002","unstructured":"Levin, L. A. & Michener, R. H. Isotopic evidence for chemosynthesis-based nutrition of macrobenthos: The lightness of being at Pacific methane seeps. Limnol. Oceanogr. 47(5), 1336\u20131345 (2002).","journal-title":"Limnol. Oceanogr."},{"issue":"12","key":"32153_CR59","doi-asserted-by":"crossref","first-page":"1937","DOI":"10.1016\/j.dsr.2004.07.010","volume":"51","author":"K Kiriakoulakis","year":"2004","unstructured":"Kiriakoulakis, K., Bett, B. J., White, M. & Wolff, G. A. Organic biogeochemistry of the Darwin Mounds, a deep-water coral ecosystem, of the NE Atlantic. Deep Sea Res. Part Oceanogr. Res. Pap. 51(12), 1937\u20131954 (2004).","journal-title":"Deep Sea Res. Part Oceanogr. Res. Pap."},{"issue":"21","key":"32153_CR60","doi-asserted-by":"crossref","first-page":"1957","DOI":"10.1016\/j.dsr2.2010.05.002","volume":"57","author":"EL Becker","year":"2010","unstructured":"Becker, E. L., Lee, R. W., Macko, S. A., Faure, B. M. & Fisher, C. R. Stable carbon and nitrogen isotope compositions of hydrocarbon-seep bivalves on the Gulf of Mexico lower continental slope. Deep Sea Res. Part II Top. Stud. Oceanogr. 57(21), 1957\u20131964 (2010).","journal-title":"Deep Sea Res. Part II Top. Stud. Oceanogr."},{"issue":"1","key":"32153_CR61","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1002\/ece3.4712","volume":"9","author":"C Ferrier-Pag\u00e8s","year":"2018","unstructured":"Ferrier-Pag\u00e8s, C. & Leal, M. C. Stable isotopes as tracers of trophic interactions in marine mutualistic symbioses. Ecol. Evol. 9(1), 723\u2013740 (2018).","journal-title":"Ecol. Evol."},{"issue":"3","key":"32153_CR62","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/0009-2541(92)90009-T","volume":"101","author":"MC Kennicutt","year":"1992","unstructured":"Kennicutt, M. C. et al. Stable isotope partitioning in seep and vent organisms: Chemical and ecological significance. Chem. Geol. Isot. Geosci. Sect. 101(3), 293\u2013310 (1992).","journal-title":"Chem. Geol. Isot. Geosci. Sect."},{"issue":"4","key":"32153_CR63","doi-asserted-by":"crossref","first-page":"1694","DOI":"10.1128\/AEM.71.4.1694-1700.2005","volume":"71","author":"S Duperron","year":"2005","unstructured":"Duperron, S. et al. Dual symbiosis in a Bathymodiolus sp. mussel from a methane seep on the Gabon Continental Margin (Southeast Atlantic): 16S rRNA phylogeny and distribution of the symbionts in gills. Appl. Environ. Microbiol. 71(4), 1694\u20131700 (2005).","journal-title":"Appl. Environ. Microbiol."},{"issue":"2","key":"32153_CR64","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1128\/AEM.70.2.1190-1198.2004","volume":"70","author":"M Sunamura","year":"2004","unstructured":"Sunamura, M., Higashi, Y., Miyako, C., Ishibashi, J. I. & Maruyama, A. Two bacteria phylotypes are predominant in the suiyo seamount hydrothermal plume. Appl. Environ. Microbiol. 70(2), 1190\u20131198 (2004).","journal-title":"Appl. Environ. Microbiol."},{"issue":"1","key":"32153_CR65","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41467-018-04041-x","volume":"9","author":"CM Callbeck","year":"2018","unstructured":"Callbeck, C. M. et al. Oxygen minimum zone cryptic sulfur cycling sustained by offshore transport of key sulfur oxidizing bacteria. Nat. Commun. 9(1), 1\u201311 (2018).","journal-title":"Nat. Commun."},{"issue":"26","key":"32153_CR66","first-page":"1","volume":"2022","author":"B Dede","year":"2022","unstructured":"Dede, B. et al. Niche differentiation of sulfur-oxidizing bacteria (SUP05) in submarine hydrothermal plumes. ISME J. 2022(26), 1\u201312 (2022).","journal-title":"ISME J."},{"issue":"5","key":"32153_CR67","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1038\/s41579-019-0160-2","volume":"17","author":"GJ Dick","year":"2019","unstructured":"Dick, G. J. The microbiomes of deep-sea hydrothermal vents: Distributed globally, shaped locally. Nat. Rev. Microbiol. 17(5), 271\u2013283 (2019).","journal-title":"Nat. Rev. Microbiol."},{"issue":"1","key":"32153_CR68","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1086\/BBLv223n1p123","volume":"223","author":"JM Petersen","year":"2012","unstructured":"Petersen, J. M., Wentrup, C., Verna, C., Knittel, K. & Dubilier, N. Origins and evolutionary flexibility of chemosynthetic symbionts from deep-sea animals. Biol. Bull. 223(1), 123\u2013137 (2012).","journal-title":"Biol. Bull."},{"issue":"12","key":"32153_CR69","doi-asserted-by":"crossref","first-page":"2487","DOI":"10.1038\/s41564-019-0572-9","volume":"4","author":"R Ansorge","year":"2019","unstructured":"Ansorge, R. et al. Functional diversity enables multiple symbiont strains to coexist in deep-sea mussels. Nat. Microbiol. 4(12), 2487\u20132497 (2019).","journal-title":"Nat. Microbiol."},{"issue":"22","key":"32153_CR70","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1128\/AEM.00992-19","volume":"85","author":"K Zhou","year":"2019","unstructured":"Zhou, K. et al. Potential interactions between clade SUP05 sulfur-oxidizing bacteria and phages in hydrothermal vent sponges. Appl. Environ. Microbiol. 85(22), 1\u201320 (2019).","journal-title":"Appl. Environ. Microbiol."},{"key":"32153_CR71","doi-asserted-by":"publisher","first-page":"1781","DOI":"10.3389\/fmicb.2016.01781\/full","volume":"7","author":"JT Hestetun","year":"2016","unstructured":"Hestetun, J. T., Dahle, H., J\u00f8rgensen, S. L., Olsen, B. R. & Rapp, H. T. The microbiome and occurrence of methanotrophy in carnivorous sponges. Front. Microbiol. 7, 1781. https:\/\/doi.org\/10.3389\/fmicb.2016.01781\/full (2016).","journal-title":"Front. Microbiol."},{"issue":"3","key":"32153_CR72","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1007\/s10126-009-9253-7","volume":"12","author":"M Nishijima","year":"2010","unstructured":"Nishijima, M. et al. Association of thioautotrophic bacteria with deep-sea sponges. Mar. Biotechnol. N Y N. 12(3), 253\u2013260 (2010).","journal-title":"Mar. Biotechnol. N Y N."},{"issue":"4","key":"32153_CR73","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1264\/jsme2.ME10135","volume":"25","author":"TO Watsuji","year":"2010","unstructured":"Watsuji, T. O. et al. Diversity and function of epibiotic microbial communities on the galatheid crab, Shinkaia crosnieri. Microbes Environ. 25(4), 288\u2013294 (2010).","journal-title":"Microbes Environ."},{"key":"32153_CR74","doi-asserted-by":"crossref","first-page":"276","DOI":"10.3389\/fmars.2016.00276","volume":"3","author":"AL Pasulka","year":"2017","unstructured":"Pasulka, A. L. et al. Colonial tube-dwelling ciliates influence methane cycling and microbial diversity within methane seep ecosystems. Front. Mar. Sci. 3, 276 (2017).","journal-title":"Front. Mar. Sci."},{"issue":"1","key":"32153_CR75","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1007\/s10482-011-9567-0","volume":"100","author":"CF Rodrigues","year":"2011","unstructured":"Rodrigues, C. F., Hil\u00e1rio, A., Cunha, M. R., Weightman, A. J. & Webster, G. Microbial diversity in Frenulata (Siboglinidae, Polychaeta) species from mud volcanoes in the Gulf of Cadiz (NE Atlantic). Antonie Van Leeuwenhoek 100(1), 83\u201398 (2011).","journal-title":"Antonie Van Leeuwenhoek"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-32153-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-32153-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-32153-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T18:44:42Z","timestamp":1781635482000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-32153-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,24]]},"references-count":75,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["32153"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-32153-0","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,24]]},"assertion":[{"value":"11 September 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 December 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Authors declare that they have no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"9603"}}