{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T13:42:58Z","timestamp":1792676578868,"version":"4.5.0"},"reference-count":99,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T00:00:00Z","timestamp":1791331200000},"content-version":"vor","delay-in-days":279,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100011033","name":"Spanish State Research Agency","doi-asserted-by":"publisher","award":["RyC2022-038245-I"],"award-info":[{"award-number":["RyC2022-038245-I"]}],"id":[{"id":"10.13039\/501100011033","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011033","name":"Spanish State Research Agency","doi-asserted-by":"publisher","award":["PID2023-152168NB-I00"],"award-info":[{"award-number":["PID2023-152168NB-I00"]}],"id":[{"id":"10.13039\/501100011033","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011033","name":"Spanish State Research Agency","doi-asserted-by":"publisher","award":["CNS2025-165952"],"award-info":[{"award-number":["CNS2025-165952"]}],"id":[{"id":"10.13039\/501100011033","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,7]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Host\u2013associated microbiomes play vital roles in species\u2019 ecology and evolution, yet little is known about their shift during the transition from aquatic to terrestrial habitats. We characterize the skin microbiomes of three combtooth blenny species (Blenniella paula, Praealticus labrovittatus, Alticus arnoldorum) that occupy distinct positions along the intertidal gradient\u2014from fully subtidal to intertidal and supratidal environments. Using 16S rRNA sequencing, we compared skin-associated bacterial communities with those in seawater and substrate biofilms. The divergence between skin and substrate microbiomes parallels fishes\u2019 distribution along progressively higher intertidal zones. The supratidal A. arnoldorum had the most divergent microbiome, characterized by higher Gammaproteobacteria abundance and enrichment of epiphytic and mucus-associated taxa. Fish skins were enriched in 27 microbial orders that included genera commonly associated with fish (e.g. Vibrio, Alteromonas, Cetobacterium) but others rarely reported (e.g. Rubritalea, Granulosicoccus). Together, our findings suggest that variation in fish skin microbiomes parallels the ecological transition from subtidal (aquatic) to supratidal (terrestrial) habitats, although host and habitat effects could not be disentangled. We hypothesize that microbial symbionts may contribute to adaptations enabling amphibious lifestyles. This is the first study on fish microbiome variation across the intertidal gradient and offers a framework for predicting microbiome responses to environmental change.<\/jats:p>","DOI":"10.1093\/femsle\/fnag123","type":"journal-article","created":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T11:52:30Z","timestamp":1791373950000},"source":"Crossref","is-referenced-by-count":0,"title":["Skin microbiome mirrors habitat divergence in amphibious combtooth blenny fish (Teleostei, Blenniidae)"],"prefix":"10.1093","volume":"373","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-3760-8725","authenticated-orcid":false,"given":"Ewelina","family":"Rubin","sequence":"first","affiliation":[{"name":"Marine Laboratory, University of Guam , Mangilao, GU 96913 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2494-1345","authenticated-orcid":false,"given":"Michele","family":"Felletti","sequence":"additional","affiliation":[{"name":"Max Planck Institute for Multidisciplinary Sciences Department of Physical Biochemistry, , 37077 G\u00f6ttingen ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-2890-2403","authenticated-orcid":false,"given":"Therese C","family":"Miller","sequence":"additional","affiliation":[{"name":"Institute of Marine Sciences, University of Auckland , 1010 Auckland ,","place":["New Zealand"]},{"name":"Cawthron Institute , 7010 Nelson ,","place":["New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9554-3125","authenticated-orcid":false,"given":"Bastian","family":"Bentlage","sequence":"additional","affiliation":[{"name":"Marine Laboratory, University of Guam , Mangilao, GU 96913 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8161-2620","authenticated-orcid":false,"given":"Diego F B","family":"Vaz","sequence":"additional","affiliation":[{"name":"Natural History Museum , South Kensington, SW7 5BD London ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2608-2150","authenticated-orcid":false,"given":"Terry\u00a0J","family":"Ord","sequence":"additional","affiliation":[{"name":"Evolution and Ecology Research Center, and the School of Biological, Earth and Environmental Sciences, University of New South Wales , 2033 Kensington, New South Wales ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3628-1137","authenticated-orcid":false,"given":"Iker","family":"Irisarri","sequence":"additional","affiliation":[{"name":"Museo Nacional de Ciencias Naturales (MNCN-CSIC) Department of Biodiversity and Evolutionary Biology, , 28006 Madrid ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,10,7]]},"reference":[{"key":"2026102209164141700_bib1","doi-asserted-by":"publisher","first-page":"3543","DOI":"10.1002\/ar.24908","article-title":"Comparison of vertebrate skin structure at class level: a review","volume":"305","author":"Akat","year":"2022","journal-title":"Anat Rec"},{"key":"2026102209164141700_bib2","doi-asserted-by":"publisher","first-page":"21269","DOI":"10.1038\/s41598-023-48533-3","article-title":"Growth performance, gut microbiota composition, health and welfare of European sea bass (Dicentrarchus labrax) fed an environmentally and economically sustainable low marine protein diet in sea cages","volume":"13","author":"Alfonso","year":"2023","journal-title":"Sci Rep"},{"key":"2026102209164141700_bib3","doi-asserted-by":"publisher","first-page":"129","DOI":"10.3354\/ame01753","article-title":"Minor revision to V4 region SSU rRNA 806R gene primer greatly increases detection of SAR11 bacterioplankton","volume":"75","author":"Apprill","year":"2015","journal-title":"Aquat Microb Ecol"},{"key":"2026102209164141700_bib4","doi-asserted-by":"publisher","first-page":"1","DOI":"10.25177\/JAFFS.2.2.RA.459","article-title":"Fish are not alone: characterization of the gut and skin microbiomes of Largemouth Bass (Micropterus salmoides), Bluegill (Lepomis macrochirus), and Spotted Gar (Lepisosteus oculatus)","volume":"2","author":"Arias","year":"2019","journal-title":"SDRP-JAFFS"},{"key":"2026102209164141700_bib5","doi-asserted-by":"publisher","first-page":"21033","DOI":"10.1038\/s41598-023-48308-w","article-title":"Species-specific gill\u2019s microbiome of eight crab species with different breathing adaptations","volume":"13","author":"Bacci","year":"2023","journal-title":"Sci Rep"},{"key":"2026102209164141700_bib6","doi-asserted-by":"publisher","first-page":"4103","DOI":"10.1099\/ijs.0.070045-0","article-title":"Granulosicoccus marinus sp. nov., isolated from Antarctic seawater, and emended description of the genus Granulosicoccus","volume":"64","author":"Baek","year":"2014","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026102209164141700_bib7","doi-asserted-by":"publisher","first-page":"3201","DOI":"10.21105\/joss.03201","article-title":"MicroViz: an R package for microbiome data visualization and statistics","volume":"6","author":"Barnett","year":"2021","journal-title":"JOSS"},{"key":"2026102209164141700_bib8","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1016\/j.cell.2016.12.019","article-title":"A periplasmic polymer curves Vibrio cholerae and promotes pathogenesis","volume":"168","author":"Bartlett","year":"2017","journal-title":"Cell"},{"key":"2026102209164141700_bib9","doi-asserted-by":"publisher","first-page":"fiae021","DOI":"10.1093\/femsec\/fiae021","article-title":"Influence of host phylogeny and water physicochemistry on microbial assemblages of the fish skin microbiome","volume":"100","author":"Bell","year":"2024","journal-title":"FEMS Microbiol Ecol"},{"key":"2026102209164141700_bib10","doi-asserted-by":"publisher","first-page":"3137","DOI":"10.1093\/bioinformatics\/btx373","article-title":"FQC Dashboard: integrates FastQC results into a web-based, interactive, and extensible FASTQ quality control tool","volume":"33","author":"Brown","year":"2017","journal-title":"Bioinformatics"},{"key":"2026102209164141700_bib11","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1016\/j.tree.2012.03.011","article-title":"Speciation by symbiosis","volume":"27","author":"Brucker","year":"2012","journal-title":"Trends Ecol Evol"},{"key":"2026102209164141700_bib12","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1038\/nmeth.3869","article-title":"DADA2: high-resolution sample inference from Illumina amplicon data","volume":"13","author":"Callahan","year":"2016","journal-title":"Nat Methods"},{"key":"2026102209164141700_bib13","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1007\/s40257-020-00551-x","article-title":"Skin Microbiome and its Interplay with the Environment","volume":"21","author":"Callewaert","year":"2020","journal-title":"Am J Clin Dermatol"},{"key":"2026102209164141700_bib14","doi-asserted-by":"publisher","first-page":"964","DOI":"10.3390\/microorganisms9050964","article-title":"Alteration of the immune response and the microbiota of the skin during a natural infection by Vibrio harveyi in European seabass (Dicentrarchus labrax)","volume":"9","author":"C\u00e1mara-Ruiz","year":"2021","journal-title":"Microorganisms"},{"key":"2026102209164141700_bib15","doi-asserted-by":"publisher","first-page":"575372","DOI":"10.3389\/fmicb.2020.575372","article-title":"To the land and beyond: crab microbiomes as a paradigm for the evolution of terrestrialization","volume":"11","author":"Cannicci","year":"2020","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib16","doi-asserted-by":"publisher","first-page":"1758990","DOI":"10.3389\/fmicb.2026.1758990","article-title":"Advances in microbial sulfoquinovose catabolism","volume":"17","author":"Chen","year":"2026","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib17","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1186\/s40168-018-0530-4","article-title":"Skin microbiome of coral reef fish is highly variable and driven by host phylogeny and diet","volume":"6","author":"Chiarello","year":"2018","journal-title":"Microbiome"},{"key":"2026102209164141700_bib18","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1007\/s12088-020-00900-8","article-title":"A preliminary assessment of skin microbiome diversity of zebrafish (Danio rerio): south African pet shop fish","volume":"61","author":"Coetzer","year":"2021","journal-title":"Indian J Microbiol"},{"key":"2026102209164141700_bib19","doi-asserted-by":"publisher","first-page":"2945","DOI":"10.1007\/s10499-023-01117-4","article-title":"Tilapia aquaculture, emerging diseases, and the roles of the skin microbiomes in health and disease","volume":"31","author":"Debnath","year":"2023","journal-title":"Aquacult Int"},{"key":"2026102209164141700_bib20","doi-asserted-by":"publisher","first-page":"1090","DOI":"10.1016\/j.marpolbul.2006.05.002","article-title":"Polycyclic aromatic hydrocarbons (PAHs) in small island coastal environments: a case study from harbours in Guam, Micronesia","volume":"52","author":"Denton","year":"2006","journal-title":"Mar Pollut Bull"},{"key":"2026102209164141700_bib21","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1038\/s41587-020-0548-6","article-title":"PICRUSt2 for prediction of metagenome functions","volume":"38","author":"Douglas","year":"2020","journal-title":"Nat Biotechnol"},{"key":"2026102209164141700_bib22","doi-asserted-by":"publisher","first-page":"1083","DOI":"10.1099\/mic.0.26111-0","article-title":"Genetic relatedness and phenotypic characteristics of Treponema associated with human periodontal tissues and ruminant foot disease","volume":"149","author":"Edwards","year":"2003","journal-title":"Microbiology"},{"key":"2026102209164141700_bib23","doi-asserted-by":"publisher","first-page":"obab004","DOI":"10.1093\/iob\/obab004","article-title":"Patterns of body shape diversity and evolution in intertidal and subtidal lineages of combtooth blennies (Blenniidae)","volume":"3","author":"Egan","year":"2021","journal-title":"Integr Org Biol"},{"key":"2026102209164141700_bib24","doi-asserted-by":"publisher","first-page":"1","DOI":"10.5402\/2012\/853470","article-title":"An overview of the immunological defenses in fish skin","volume":"2012","author":"Esteban","year":"2012","journal-title":"ISRN Immunol"},{"key":"2026102209164141700_bib25","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1080\/07388551.2021.1922351","article-title":"Marine algal flavonoids and phlorotannins; an intriguing frontier of biofunctional secondary metabolites","volume":"42","author":"Fernando","year":"2022","journal-title":"Crit Rev Biotechnol"},{"key":"2026102209164141700_bib26","first-page":"1369","article-title":"Current state of knowledge in microbial degradation of polycyclic aromatic hydrocarbons (PAHs): a review","volume":"7","author":"Ghosal","year":"2016","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib27","doi-asserted-by":"publisher","first-page":"877","DOI":"10.1007\/s00360-013-0761-4","article-title":"The skin of fish as a transport epithelium: a review","volume":"183","author":"Glover","year":"2013","journal-title":"J Comp Physiol B"},{"key":"2026102209164141700_bib28","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1007\/s00248-020-01648-w","article-title":"A slimy business: the future of fish skin microbiome studies","volume":"82","author":"Gomez","year":"2021","journal-title":"Microb Ecol"},{"key":"2026102209164141700_bib29","doi-asserted-by":"publisher","first-page":"2847","DOI":"10.1093\/bioinformatics\/btw313","article-title":"Complex heatmaps reveal patterns and correlations in multidimensional genomic data","volume":"32","author":"Gu","year":"2016","journal-title":"Bioinformatics"},{"key":"2026102209164141700_bib30","doi-asserted-by":"publisher","first-page":"214","DOI":"10.3390\/fishes8040214","article-title":"Environmental impacts on skin microbiomes of sympatric high Arctic salmonids","volume":"8","author":"Hamilton","year":"2023","journal-title":"Fishes"},{"key":"2026102209164141700_bib31","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1111\/j.1574-6968.1999.tb08775.x","article-title":"Conservation of the pyrrolnitrin biosynthetic gene cluster among six pyrrolnitrin-producing strains","volume":"180","author":"Hammer","year":"1999","journal-title":"FEMS Microbiol Lett"},{"key":"2026102209164141700_bib32","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1007\/s00248-024-02467-z","article-title":"Phylosymbiosis in seven wild fish species collected off the southern coast of Korea: skin microbiome most strongly reflects evolutionary pressures","volume":"87","author":"Han","year":"2024","journal-title":"Microb Ecol"},{"key":"2026102209164141700_bib33","doi-asserted-by":"publisher","first-page":"eabj0829","DOI":"10.1126\/sciadv.abj0829","article-title":"The lungfish cocoon is a living tissue with antimicrobial functions","volume":"7","author":"Heimroth","year":"2021","journal-title":"Sci Adv"},{"key":"2026102209164141700_bib34","doi-asserted-by":"publisher","first-page":"e11197","DOI":"10.1371\/journal.pone.0011197","article-title":"A locomotor innovation enables water-land transition in a marine fish","volume":"5","author":"Hsieh","year":"2010","journal-title":"PLoS One"},{"key":"2026102209164141700_bib35","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1111\/1758-2229.12686","article-title":"Coral microbiome database: integration of sequences reveals high diversity and relatedness of coral-associated microbes","volume":"11","author":"Huggett","year":"2019","journal-title":"Environ Microbiol Rep"},{"key":"2026102209164141700_bib36","first-page":"249","article-title":"The diet and gut microbial communities of two closely related combtooth blennies, Chasmodes saburrae and Scartella cristata","volume":"105","author":"Hundt","year":"2017","journal-title":"Ichthyol Herpetol"},{"key":"2026102209164141700_bib37","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.ympev.2013.09.001","article-title":"A multilocus molecular phylogeny of combtooth blennies (Percomorpha: blennioidei: blenniidae): multiple invasions of intertidal habitats","volume":"70","author":"Hundt","year":"2014","journal-title":"Mol Phylogenet Evol"},{"key":"2026102209164141700_bib38","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1016\/S1049-9644(02)00044-0","article-title":"Pyrrolnitrin production by Burkholderia cepacia and biocontrol of Rhizoctonia stem rot of poinsettia","volume":"25","author":"Hwang","year":"2002","journal-title":"Biol Control"},{"key":"2026102209164141700_bib39","doi-asserted-by":"publisher","first-page":"e0322261","DOI":"10.1371\/journal.pone.0322261","article-title":"Effect of plastic and seaweed shelters on the skin microbiome of lumpfish Cyclopterus lumpus used as cleaner fish in aquaculture pens","volume":"20","author":"Jacobsen","year":"2025","journal-title":"PLoS One"},{"key":"2026102209164141700_bib40","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":"2026102209164141700_bib41","doi-asserted-by":"publisher","first-page":"5112","DOI":"10.1128\/AEM.01043-13","article-title":"Development of a dual-index sequencing strategy and curation pipeline for analyzing amplicon sequence data on the MiSeq Illumina sequencing platform","volume":"79","author":"Kozich","year":"2013","journal-title":"Appl Environ Microb"},{"key":"2026102209164141700_bib42","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1186\/s40168-020-0784-5","article-title":"Dissecting the factors shaping fish skin microbiomes in a heterogeneous inland water system","volume":"8","author":"Krotman","year":"2020","journal-title":"Microbiome"},{"key":"2026102209164141700_bib43","doi-asserted-by":"publisher","first-page":"972","DOI":"10.1099\/ijs.0.013516-0","article-title":"Granulosicoccus coccoides sp. nov., isolated from leaves of seagrass (Zostera marina)","volume":"60","author":"Kurilenko","year":"2010","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026102209164141700_bib44","first-page":"1","article-title":"Revisiting the genus Photobacterium: taxonomy, ecology and pathogenesis","volume":"20","author":"Labella","year":"2017","journal-title":"Int Microbiol"},{"key":"2026102209164141700_bib45","doi-asserted-by":"publisher","DOI":"10.18129\/B9.bioc.microbiome","volume-title":"Microbiome R package. Version 3.23","author":"Lahti","year":"2012\u20132019"},{"key":"2026102209164141700_bib46","doi-asserted-by":"publisher","first-page":"e0004722","DOI":"10.1128\/msystems.00047-22","article-title":"Integrated omics approaches revealed the osmotic stress-responsive genes and microbiota in gill of marine medaka","volume":"7","author":"Lai","year":"2022","journal-title":"mSystems"},{"key":"2026102209164141700_bib47","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1111\/1574-6941.12136","article-title":"Diversity of the skin microbiota of fishes: evidence for host species specificity","volume":"85","author":"Larsen","year":"2013","journal-title":"FEMS Microbiol Ecol"},{"key":"2026102209164141700_bib48","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1002\/evl3.219","article-title":"From scales to armor: scale losses and trunk bony plate gains in ray-finned fishes","volume":"5","author":"Lemopoulos","year":"2021","journal-title":"Evol Lett"},{"key":"2026102209164141700_bib49","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1186\/s42523-020-00071-3","article-title":"Differential response of digesta- and mucosa-associated intestinal microbiota to dietary insect meal during the seawater phase of Atlantic salmon","volume":"3","author":"Li","year":"2021","journal-title":"Anim Microbiome"},{"key":"2026102209164141700_bib50","first-page":"20192900","article-title":"An introduction to phylosymbiosis","volume":"287","author":"Lim","year":"2020","journal-title":"Proc Biol Sci"},{"key":"2026102209164141700_bib51","doi-asserted-by":"publisher","first-page":"207","DOI":"10.3389\/fmicb.2014.00207","article-title":"Teleost microbiomes: the state of the art in their characterization, manipulation and importance in aquaculture and fisheries","volume":"5","author":"Llewellyn","year":"2014","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib52","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1038\/nrmicro3400","article-title":"Ecology and exploration of the rare biosphere","volume":"13","author":"Lynch","year":"2015","journal-title":"Nat Rev Micro"},{"key":"2026102209164141700_bib53","doi-asserted-by":"publisher","first-page":"101459","DOI":"10.1016\/j.aqrep.2022.101459","article-title":"Pathology and pathogenesis of Vibrio infection in fish: a review","volume":"28","author":"Manchanayake","year":"2023","journal-title":"Aquac Rep"},{"key":"2026102209164141700_bib54","doi-asserted-by":"publisher","first-page":"723","DOI":"10.2307\/1445501","article-title":"Aerial CO2 and O2 exchange during terrestrial activity in an amphibious fish, Alticus kirki (Blenniidae)","volume":"1989","author":"Martin","year":"1989","journal-title":"Copeia"},{"key":"2026102209164141700_bib55","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1214\/19-AOAS1283","article-title":"Modeling microbial abundances and dysbiosis with beta-binomial regression","volume":"14","author":"Martin","year":"2020","journal-title":"Ann Appl Stat"},{"key":"2026102209164141700_bib56","doi-asserted-by":"publisher","first-page":"e61217","DOI":"10.1371\/journal.pone.0061217","article-title":"Phyloseq: an R package for reproducible interactive analysis and graphics of microbiome census data","volume":"8","author":"McMurdie","year":"2013","journal-title":"PLoS One"},{"key":"2026102209164141700_bib57","doi-asserted-by":"publisher","first-page":"fuaf027","DOI":"10.1093\/femsre\/fuaf027","article-title":"The ecology and plasticity of fish skin and gill microbiomes: seeking what matters in health and disease","volume":"49","author":"McMurtrie","year":"2025","journal-title":"FEMS Microbiol Rev"},{"key":"2026102209164141700_bib58","doi-asserted-by":"publisher","first-page":"3537","DOI":"10.1007\/978-1-4757-2191-1","article-title":"The genus Treponema","volume-title":"The Prokaryotes","author":"Miller","year":"1992"},{"key":"2026102209164141700_bib59","doi-asserted-by":"publisher","first-page":"6978","DOI":"10.1038\/s41467-022-34557-2","article-title":"Host biology, ecology and the environment influence microbial biomass and diversity in 101 marine fish species","volume":"13","author":"Minich","year":"2022","journal-title":"Nat Commun"},{"key":"2026102209164141700_bib60","doi-asserted-by":"publisher","first-page":"4371","DOI":"10.1007\/s00253-017-8300-y","article-title":"The biology and the importance of Photobacterium species","volume":"101","author":"Moi","year":"2017","journal-title":"Appl Microbiol Biotechnol"},{"key":"2026102209164141700_bib61","doi-asserted-by":"crossref","unstructured":"Mu\u00f1oz-S\u00e1nchez \u00c1, Kulikov N, Irisarri I. \u00a0Stitch or cluster? A comparison of alternative phylogenomic dataset assembly strategies for blenny fish. Bull Soc Syst Biol. 2026;5. 10.18061\/bssb.6848.","DOI":"10.18061\/bssb.6848"},{"key":"2026102209164141700_bib62","volume-title":"vegan: Community Ecology Package.","author":"Oksanen","year":"2026"},{"key":"2026102209164141700_bib63","doi-asserted-by":"publisher","first-page":"1747","DOI":"10.1111\/evo.12971","article-title":"Repeated evolution of amphibious behavior in fish and its implications for the colonization of novel environments","volume":"70","author":"Ord","year":"2016","journal-title":"Evolution"},{"key":"2026102209164141700_bib64","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1111\/j.1439-0310.2011.01949.x","article-title":"A highly social, land-dwelling fish defends territories in a constantly fluctuating environment","volume":"117","author":"Ord","year":"2011","journal-title":"Ethology"},{"key":"2026102209164141700_bib65","doi-asserted-by":"publisher","first-page":"570","DOI":"10.1086\/691155","article-title":"Ecological release from aquatic predation is associated with the emergence of marine blenny fishes onto land","volume":"189","author":"Ord","year":"2017","journal-title":"Am Nat"},{"key":"2026102209164141700_bib66","doi-asserted-by":"publisher","first-page":"2648","DOI":"10.1111\/1365-2435.14672","article-title":"Abiotic factors that prompt major ecological transitions: are fish on land to escape an intolerable aquatic environment?","volume":"38","author":"Ord","year":"2024","journal-title":"Functional Ecol"},{"key":"2026102209164141700_bib67","doi-asserted-by":"publisher","first-page":"1403","DOI":"10.1111\/1462-2920.13023","article-title":"Every base matters: assessing small subunit rRNA primers for marine microbiomes with mock communities, time series and global field samples","volume":"18","author":"Parada","year":"2016","journal-title":"Environ Microbiol"},{"key":"2026102209164141700_bib68","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1007\/s10482-014-0254-9","article-title":"Granulosicoccus undariae sp. nov., a member of the family Granulosicoccaceae isolated from a brown algae reservoir and emended description of the genus Granulosicoccus","volume":"106","author":"Park","year":"2014","journal-title":"Antonie Van Leeuwenhoek"},{"key":"2026102209164141700_bib69","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1186\/s12866-023-03025-2","article-title":"Host individual and gut location are more important in gut microbiota community composition than temporal variation in the marine herbivorous fish Kyphosus sydneyanus","volume":"23","author":"Pisaniello","year":"2023","journal-title":"BMC Microbiol"},{"key":"2026102209164141700_bib70","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1007\/s00442-016-3593-6","article-title":"Land colonisation by fish is associated with predictable changes in life history","volume":"181","author":"Platt","year":"2016","journal-title":"Oecologia"},{"key":"2026102209164141700_bib71","doi-asserted-by":"publisher","first-page":"e00063","DOI":"10.1128\/AEM.00063-18","article-title":"The gills of reef fish support a distinct microbiome influenced by host-specific factors","volume":"84","author":"Pratte","year":"2018","journal-title":"Appl Environ Microb"},{"key":"2026102209164141700_bib72","doi-asserted-by":"publisher","first-page":"D590","DOI":"10.1093\/nar\/gks1219","article-title":"The SILVA ribosomal RNA gene database project: improved data processing and web-based tools","volume":"41","author":"Quast","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"2026102209164141700_bib73","doi-asserted-by":"publisher","first-page":"189","DOI":"10.3390\/ani8110189","article-title":"Cetobacterium is a major component of the microbiome of giant Amazonian fish (Arapaima gigas) in Ecuador","volume":"8","author":"Ram\u00edrez","year":"2018","journal-title":"Animals"},{"key":"2026102209164141700_bib74","doi-asserted-by":"publisher","first-page":"41","DOI":"10.3390\/fishes3040041","article-title":"Biological and ecological roles of external fish mucus: a review","volume":"3","author":"Reverter","year":"2018","journal-title":"Fishes"},{"key":"2026102209164141700_bib75","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1186\/s40168-019-0694-6","article-title":"The skin microbiome of vertebrates","volume":"7","author":"Ross","year":"2019","journal-title":"Microbiome"},{"key":"2026102209164141700_bib76","doi-asserted-by":"publisher","first-page":"2119","DOI":"10.1099\/ijs.0.64360-0","article-title":"Rubritalea marina gen. nov., sp. nov., a marine representative of the phylum \u201cVerrucomicrobia\u201d, isolated from a sponge (Porifera)","volume":"56","author":"Scheuermayer","year":"2006","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026102209164141700_bib77","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1515\/bot-2020-0023","article-title":"Sulfoquinovose metabolism in marine algae","volume":"64","author":"Scholz","year":"2021","journal-title":"Bot Mar"},{"key":"2026102209164141700_bib78","doi-asserted-by":"publisher","first-page":"567408","DOI":"10.3389\/fmicb.2021.567408","article-title":"Microbiome composition and function in aquatic vertebrates: small organisms making big impacts on aquatic animal health","volume":"12","author":"Sehnal","year":"2021","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib79","doi-asserted-by":"publisher","first-page":"1690","DOI":"10.3390\/md12031690","article-title":"New and rare carotenoids isolated from marine bacteria and their antioxidant activities","volume":"12","author":"Shindo","year":"2014","journal-title":"Mar Drugs"},{"key":"2026102209164141700_bib80","doi-asserted-by":"publisher","first-page":"1465","DOI":"10.1099\/mic.0.001244","article-title":"Vitamin B12 biosynthesis of Cetobacterium ceti isolated from the intestinal content of captive common bottlenose dolphins (Tursiops truncatus)","volume":"168","author":"Suzuki","year":"2022","journal-title":"Microbiology"},{"key":"2026102209164141700_bib81","doi-asserted-by":"publisher","first-page":"e00789","DOI":"10.1128\/AEM.00789-20","article-title":"Fish skin and gut microbiomes show contrasting signatures of host species and habitat","volume":"86","author":"Sylvain","year":"2020","journal-title":"Appl Environ Microb"},{"key":"2026102209164141700_bib82","volume-title":"Fantaxtic\u2014Nested bar plots for Phyloseq data","author":"Teunisse","year":"2022"},{"key":"2026102209164141700_bib83","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1186\/s43008-020-00048-6","article-title":"Setting scientific names at all taxonomic ranks in italics facilitates their quick recognition in scientific papers","volume":"11","author":"Thines","year":"2020","journal-title":"IMA Fungus"},{"key":"2026102209164141700_bib84","doi-asserted-by":"publisher","first-page":"541853","DOI":"10.3389\/fmars.2020.541853","article-title":"Skin mucus of marine fish as a source for the development of antimicrobial agents","volume":"7","author":"Tiralongo","year":"2020","journal-title":"Front Mar Sci"},{"key":"2026102209164141700_bib85","first-page":"43","article-title":"Novel ecological niche of Cetobacterium somerae, an anaerobic bacterium in the intestinal tracts of freshwater fish","volume":"46","author":"Tsuchiya","year":"2008","journal-title":"Lett Appl Microbiol"},{"key":"2026102209164141700_bib86","first-page":"301","article-title":"Oxygen tolerance of Cetobacterium somerae isolated from the gut of freshwater fish and their environments","volume":"11","author":"Tsuchiya","year":"2023","journal-title":"Int J Aquat Biol"},{"key":"2026102209164141700_bib87","doi-asserted-by":"crossref","first-page":"4257","DOI":"10.1038\/s41598-022-07980-0","article-title":"DNA- and RNA-based bacterial communities and geochemical zonation under changing sediment porewater dynamics on the Aldabra Atoll","volume":"12","author":"von\u00a0Hoyningen-Huene","year":"2022","journal-title":"Sci Rep"},{"key":"2026102209164141700_bib88","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1186\/s40168-025-02152-4","article-title":"Cetobacterium somerae-derived argininosuccinic acid promotes intestinal and liver ureagenesis to alleviate ammonia intoxication","volume":"13","author":"Wang","year":"2025","journal-title":"Microbiome"},{"key":"2026102209164141700_bib89","doi-asserted-by":"publisher","first-page":"102583","DOI":"10.1016\/j.aqrep.2024.102583","article-title":"Effect of the gut core microbiota Cetobacterium on the growth, physiology, and nutritional metabolism of Nile tilapia (Oreochromis niloticus)","volume":"40","author":"Wang","year":"2025","journal-title":"Aquac Rep"},{"key":"2026102209164141700_bib90","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1111\/j.1095-8649.1984.tb04879.x","article-title":"Surface secretions of the skin of Blennius (Lipophrys) pholis L","volume":"25","author":"Whitear","year":"1984","journal-title":"J Fish Biol"},{"key":"2026102209164141700_bib91","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","volume-title":"Ggplot2: Elegant graphics for data analysis","author":"Wickham","year":"2016"},{"key":"2026102209164141700_bib92","doi-asserted-by":"publisher","first-page":"101046","DOI":"10.1016\/j.aqrep.2022.101046","article-title":"Growth performance, hepatic enzymes, and gut health status of common carp (Cyprinus carpio) in response to dietary Cetobacterium somerae fermentation product","volume":"23","author":"Xie","year":"2022","journal-title":"Aquac Rep"},{"key":"2026102209164141700_bib93","doi-asserted-by":"publisher","first-page":"1886","DOI":"10.3389\/fmicb.2020.01886","article-title":"Crawling motility on the host tissue surfaces is associated with the pathogenicity of the zoonotic spirochete Leptospira","volume":"11","author":"Xu","year":"2020","journal-title":"Front Microbiol"},{"key":"2026102209164141700_bib94","doi-asserted-by":"publisher","first-page":"103078","DOI":"10.1016\/j.aqrep.2025.103078","article-title":"Effects of butyric acid on the inhibition of Vibrio parahaemolyticus and its impacts on growth performance, intestinal microbiota, and water microbiota in Litopenaeus vannamei","volume":"45","author":"Yin","year":"2025","journal-title":"Aquac Rep"},{"key":"2026102209164141700_bib95","doi-asserted-by":"publisher","first-page":"1515","DOI":"10.1099\/ijs.0.025031-0","article-title":"Rubritalea halochordaticola sp. nov., a carotenoid-producing verrucomicrobial species isolated from a marine chordate","volume":"61","author":"Yoon","year":"2011","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026102209164141700_bib96","doi-asserted-by":"publisher","first-page":"2337","DOI":"10.1099\/ijs.0.65243-0","article-title":"Rubritalea spongiae sp. nov. and Rubritalea tangerina sp. nov., two carotenoid- and squalene-producing marine bacteria of the family Verrucomicrobiaceae within the phylum \u201cVerrucomicrobia\u201d, isolated from marine animals","volume":"57","author":"Yoon","year":"2007","journal-title":"Int J Syst Evol Microbiol"},{"key":"2026102209164141700_bib97","doi-asserted-by":"publisher","first-page":"4642","DOI":"10.1038\/s41598-022-08511-7","article-title":"Gut microbiota analysis of Blenniidae fishes including an algae-eating fish and clear boundary formation among isolated Vibrio strains","volume":"12","author":"Yoshida","year":"2022","journal-title":"Sci Rep"},{"key":"2026102209164141700_bib98","doi-asserted-by":"publisher","first-page":"106148","DOI":"10.1016\/j.micpath.2023.106148","article-title":"Characterization of Cetobacterium somerae CPU-CS01 isolated from the intestine of healthy crucian carp (Carassius auratus) as potential probiotics against Aeromonas hydrophila infection","volume":"180","author":"Zhao","year":"2023","journal-title":"Microb Pathog"},{"key":"2026102209164141700_bib99","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1016\/j.fsi.2022.04.019","article-title":"Effect of dietary supplementation of Cetobacterium somerae XMX-1 fermentation product on gut and liver health and resistance against bacterial infection of the genetically improved farmed tilapia (GIFT, Oreochromis niloticus)","volume":"124","author":"Zhou","year":"2022","journal-title":"Fish Shellfish Immunol"}],"container-title":["FEMS Microbiology Letters"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/femsle\/advance-article-pdf\/doi\/10.1093\/femsle\/fnag123\/71665376\/fnag123.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/femsle\/article-pdf\/doi\/10.1093\/femsle\/fnag123\/71665376\/fnag123.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/femsle\/article-pdf\/doi\/10.1093\/femsle\/fnag123\/71665376\/fnag123.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T13:16:50Z","timestamp":1792675010000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/femsle\/article\/doi\/10.1093\/femsle\/fnag123\/8875694"}},"subtitle":[],"editor":[{"given":"Guerrino","family":"Macori","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026]]},"references-count":99,"URL":"https:\/\/doi.org\/10.1093\/femsle\/fnag123","relation":{},"ISSN":["1574-6968"],"issn-type":[{"value":"1574-6968","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026]]},"published":{"date-parts":[[2026]]},"article-number":"fnag123"}}