{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T16:28:06Z","timestamp":1790094486923,"version":"4.0.1"},"reference-count":82,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2025,9,17]],"date-time":"2025-09-17T00:00:00Z","timestamp":1758067200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"the FFS Greub grant","award":["IMUR 31932"],"award-info":[{"award-number":["IMUR 31932"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Aquaculture plays a vital role in meeting the global demand for high-quality protein. However, the fish industry is challenged by infectious diseases, including gill conditions such as epitheliocystis. Epitheliocystis is characterized by cyst-like epithelial lesions, which occur in the gills of fish, and is associated with intracellular bacteria including Chlamydia-related bacteria. Although epitheliocystis was initially regarded as of low significance, attention is increasing due to its impact on commercially important fish species in intense farming conditions. This review evaluates the roles of aquatic chlamydiae as pathogens contributing to fish morbidity and mortality, and as members of fish microbiota. Additionally, Chlamydia-related bacteria are thought to be involved in complex gill disease (CGD), characterized by lamellar fusion, epithelial hyperplasia, and inflammation. Recent discoveries have expanded the diversity of Chlamydiota isolated from fish, with novel species such as Candidatus (Ca.) Panilichlamydia rohitae, Ca. Piscichlamydia trichopodus, and Chlamydia vaughanii identified in different fish hosts. Most causative agents of epitheliocystis have not yet been cultured in vitro, although C. vaughanii, the first Chlamydiaceae member isolated from fish, was successfully cultured. As C. vaughanii was recently shown to be able to propagate in mammalian cells, it raises concerns about its zoonotic potential, although a pathogenic role has yet to be described.<\/jats:p>","DOI":"10.3390\/microorganisms13092166","type":"journal-article","created":{"date-parts":[[2025,9,17]],"date-time":"2025-09-17T11:00:00Z","timestamp":1758106800000},"page":"2166","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Aquatic Chlamydiae: A Review of Their Roles in Fish Health"],"prefix":"10.3390","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-1794-2210","authenticated-orcid":false,"given":"Basma","family":"Mahmoud-Elkamouny","sequence":"first","affiliation":[{"name":"Institute of Microbiology, Lausanne University Hospital, University of Lausanne, 1011 Lausanne, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1158-6361","authenticated-orcid":false,"given":"Carole","family":"Kebbi-Beghdadi","sequence":"additional","affiliation":[{"name":"Institute of Microbiology, Lausanne University Hospital, University of Lausanne, 1011 Lausanne, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9529-3317","authenticated-orcid":false,"given":"Gilbert","family":"Greub","sequence":"additional","affiliation":[{"name":"Institute of Microbiology, Lausanne University Hospital, University of Lausanne, 1011 Lausanne, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,9,17]]},"reference":[{"key":"ref_1","unstructured":"FAO (2022). The State of World Fisheries and Aquaculture. Towards Blue Transformation, FAO."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1007\/s12639-017-0938-y","article-title":"An overall estimation of losses caused by diseases in the Brazilian fish farms","volume":"41","author":"Martins","year":"2017","journal-title":"J. Parasit. Dis."},{"key":"ref_3","first-page":"58","article-title":"Economic impacts of aquatic parasites on global finfish production","volume":"2015","author":"Shinn","year":"2015","journal-title":"Glob. Aquac. Advocate"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1152\/physrev.00050.2003","article-title":"The Multifunctional Fish Gill: Dominant Site of Gas Exchange, Osmoregulation, Acid-Base Regulation, and Excretion of Nitrogenous Waste","volume":"85","author":"Evans","year":"2005","journal-title":"Physiol. Rev."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.jcpa.2018.07.004","article-title":"Complex Gill Disease: An Emerging Syndrome in Farmed Atlantic Salmon (Salmo salar L.)","volume":"163","author":"Herrero","year":"2018","journal-title":"J. Comp. Pathol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"2140","DOI":"10.1111\/raq.12426","article-title":"Epidemiology of marine gill diseases in Atlantic salmon (Salmo salar) aquaculture: A review","volume":"12","author":"Boerlage","year":"2020","journal-title":"Rev. Aquac."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Plehn, M. (1924). Praktikum der fischkrankheiten, Schweizerbart (E. N\u00e4gele) gmbh.","DOI":"10.5962\/bhl.title.140778"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1007\/BF02219125","article-title":"Epitheliocystis, a new infectious disease of the bluegill (Lepomis macrochirus)","volume":"35","author":"Hoffman","year":"1969","journal-title":"Antonie Van Leeuwenhoek"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1111\/j.1365-2761.2006.00747.x","article-title":"Epitheliocystis in fish","volume":"29","author":"Nowak","year":"2006","journal-title":"J. Fish. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"61","DOI":"10.3354\/dao037061","article-title":"Epitheliocystis agents in sea bream Sparus aurata: Morphological evidence for two distinct chlamydia-like developmental cycles","volume":"37","author":"Crespo","year":"1999","journal-title":"Dis. Aquat. Org."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"27","DOI":"10.3354\/dao076027","article-title":"Characterization of a Neochlamydia-like bacterium associated with epitheliocystis in cultured Arctic charr Salvelinus alpinus","volume":"76","author":"Bebak","year":"2007","journal-title":"Dis. Aquat. Org."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"55","DOI":"10.3354\/dao01968","article-title":"Severe mortality in mesocosm-reared sharpsnout sea bream Diplodus puntazzo larvae due to epitheliocystis infection","volume":"82","author":"Katharios","year":"2008","journal-title":"Dis. Aquat. Org."},{"key":"ref_13","first-page":"105","article-title":"Gill Tissue Reactions to an Epitheliocystis Infection in Cultured Red Seabream, Pagrus major","volume":"17","author":"Syasina","year":"2004","journal-title":"J. Fish. Pathol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0044-8486(91)90067-H","article-title":"Epitheliocystis in the wild and cultured amberjack, Seriola dumerili Risso: Ultrastructural observations","volume":"95","author":"Grau","year":"1991","journal-title":"Aquaculture"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"5286","DOI":"10.1128\/JCM.42.11.5286-5297.2004","article-title":"Characterization of \u201cCandidatus Piscichlamydia salmonis\u201d (Order Chlamydiales), a Chlamydia-like Bacterium Associated with Epitheliocystis in Farmed Atlantic Salmon (Salmo salar)","volume":"42","author":"Draghi","year":"2004","journal-title":"J. Clin. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1111\/j.1365-2761.2011.01251.x","article-title":"A review of infectious gill disease in marine salmonid fish: Infectious gill disease in salmonids","volume":"34","author":"Mitchell","year":"2011","journal-title":"J. Fish. Dis."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1080\/713610404","article-title":"Increasing Diversity within Chlamydiae","volume":"29","author":"Corsaro","year":"2003","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1099\/00207713-49-2-415","article-title":"Emended description of the order Chlamydiales, proposal of Parachlamydiaceae fam. nov. and Simkaniaceae fam. nov., each containing one monotypic genus, revised taxonomy of the family Chlamydiaceae, including a new genus and five new species, and standards for the identification of organisms","volume":"49","author":"Everett","year":"1999","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1128\/CMR.19.2.283-297.2006","article-title":"Pathogenic potential of novel Chlamydiae and diagnostic approaches to infections due to these obligate intracellular bacteria","volume":"19","author":"Corsaro","year":"2006","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_20","first-page":"1","article-title":"Twenty years of research into Chlamydia-like organisms: A revolution in our understanding of the biology and pathogenicity of members of the phylum Chlamydiae","volume":"73","author":"Vaughan","year":"2015","journal-title":"FEMS Pathog. Dis."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1232","DOI":"10.1016\/j.micinf.2011.07.003","article-title":"Estrella lausannensis, a new star in the Chlamydiales order","volume":"13","author":"Lienard","year":"2011","journal-title":"Microbes Infect."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2125","DOI":"10.1111\/j.1462-2920.2006.01094.x","article-title":"Criblamydia sequanensis, a new intracellular Chlamydiales isolated from Seine river water using amoebal co-culture","volume":"8","author":"Thomas","year":"2006","journal-title":"Environ. Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1128\/aem.63.1.115-121.1997","article-title":"Obligate intracellular bacterial parasites of acanthamoebae related to Chlamydia spp. Appl","volume":"63","author":"Amann","year":"1997","journal-title":"Environ. Microbiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2613","DOI":"10.1128\/AEM.66.6.2613-2619.2000","article-title":"Phylogenetic diversity among geographically dispersed Chlamydiales endosymbionts recovered from clinical and environmental isolates of Acanthamoeba spp. Appl","volume":"66","author":"Fritsche","year":"2000","journal-title":"Environ. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/j.syapm.2006.06.001","article-title":"\u2018Candidatus Rhabdochlamydia crassificans\u2019, an intracellular bacterial pathogen of the cockroach Blatta orientalis (Insecta: Blattodea)","volume":"30","author":"Corsaro","year":"2007","journal-title":"Syst. Appl. Microbiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/j.ttbdis.2013.11.009","article-title":"Presence of Chlamydiales DNA in ticks and fleas suggests that ticks are carriers of Chlamydiae","volume":"5","author":"Croxatto","year":"2014","journal-title":"Ticks Tick-Borne Dis."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1581","DOI":"10.1099\/ijs.0.63454-0","article-title":"Novel chlamydiae in whiteflies and scale insects: Endosymbionts \u2018Candidatus Fritschea bemisiae\u2019 strain Falk and \u2018Candidatus Fritschea eriococci\u2019 strain Elm","volume":"55","author":"Everett","year":"2005","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1099\/ijs.0.02802-0","article-title":"\u2018Candidatus Rhabdochlamydia porcellionis\u2019, an intracellular bacterium from the hepatopancreas of the terrestrial isopod Porcellio scaber (Crustacea: Isopoda)","volume":"54","author":"Drobne","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"8177","DOI":"10.1128\/AEM.02183-15","article-title":"The high prevalence and diversity of Chlamydiales DNA within Ixodes ricinus ticks suggest a role for ticks as reservoirs and vectors of Chlamydia-related bacteria","volume":"81","author":"Pilloux","year":"2015","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1436","DOI":"10.1111\/tbed.12908","article-title":"Epitheliocystis in fish: An emerging aquaculture disease with a global impact","volume":"65","author":"Blandford","year":"2018","journal-title":"Transbound. Emerg. Dis."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1016\/j.aquaculture.2019.03.007","article-title":"Epitheliocystis in rohu Labeo rohita (Hamilton, 1822) is caused by novel Chlamydiales","volume":"505","author":"Sood","year":"2019","journal-title":"Aquaculture"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"847977","DOI":"10.3389\/fvets.2022.847977","article-title":"Co-infection of Candidatus Piscichlamydia Trichopodus (Order Chlamydiales) and Henneguya sp. (Myxosporea, Myxobolidae) in Snakeskin Gourami Trichopodus pectoralis (Regan 1910)","volume":"9","author":"Dong","year":"2022","journal-title":"Front. Vet. Sci."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1099\/ijs.0.000090","article-title":"Taxogenomics of the order Chlamydiales","volume":"65","author":"Pillonel","year":"2015","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"31","DOI":"10.3354\/dao03672","article-title":"Epitheliocystis caused by a novel Chlamydia emerging in Spinibarbus denticulatus in China","volume":"150","author":"Liu","year":"2022","journal-title":"Dis. Aquat. Org."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"006753","DOI":"10.1099\/ijsem.0.006753","article-title":"Chlamydia vaughanii sp. nov., a novel Chlamydia isolated from a tropical fish (bushymouth catfish)","volume":"75","author":"Marquis","year":"2025","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1177\/0300985817751218","article-title":"A Review on Chlamydial Diseases in Animals: Still a Challenge for Pathologists?","volume":"55","author":"Borel","year":"2018","journal-title":"Vet. Pathol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1128\/AEM.72.1.284-290.2006","article-title":"Molecular Evidence for Association of Chlamydiales Bacteria with Epitheliocystis in Leafy Seadragon (Phycodurus eques), Silver Perch (Bidyanus bidyanus), and Barramundi (Lates calcarifer)","volume":"72","author":"Meijer","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1016\/j.vetmic.2014.03.022","article-title":"Correction: Chlamydial infections of fish: Diverse pathogens and emerging causes of disease in aquaculture species","volume":"171","author":"Stride","year":"2014","journal-title":"Vet. Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e66840","DOI":"10.1371\/journal.pone.0066840","article-title":"\u2018Cand. Actinochlamydia clariae\u2019 gen. nov., sp. nov., a Unique Intracellular Bacterium Causing Epitheliocystis in Catfish (Clarias gariepinus) in Uganda","volume":"8","author":"Steigen","year":"2013","journal-title":"PLoS ONE"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"39","DOI":"10.3354\/dao02171","article-title":"Identification of \u2018Candidatus Piscichlamydia salmonis\u2019 in Arctic charr Salvelinus alpinus during a survey of charr production facilities in North America","volume":"89","author":"Draghi","year":"2010","journal-title":"Dis. Aquat. Org."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1111\/j.1462-2920.2007.01445.x","article-title":"Characterization of \u2018Candidatus Clavochlamydia salmonicola\u2019: An intracellular bacterium infecting salmonid fish","volume":"10","author":"Karlsen","year":"2008","journal-title":"Environ. Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1111\/j.1365-2761.2010.01171.x","article-title":"Epitheliocystis in Atlantic salmon, Salmo salar L., farmed in fresh water in Ireland is associated with \u2018Candidatus Clavochlamydia salmonicola\u2019 infection","volume":"33","author":"Mitchell","year":"2010","journal-title":"J. Fish. Dis."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2048","DOI":"10.1111\/j.1462-2920.2011.02670.x","article-title":"A natural freshwater origin for two chlamydial species, Candidatus Piscichlamydia salmonis and Candidatus Clavochlamydia salmonicola, causing mixed infections in wild brown trout (Salmo trutta)","volume":"14","author":"Polkinghorne","year":"2012","journal-title":"Environ. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1007\/s00203-014-1038-3","article-title":"Characterization of \u2018Candidatus Syngnamydia salmonis\u2019 (Chlamydiales, Simkaniaceae), a bacterium associated with epitheliocystis in Atlantic salmon (Salmo salar L.)","volume":"197","author":"Nylund","year":"2015","journal-title":"Arch. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"41","DOI":"10.3354\/dao02313","article-title":"Diseases of farmed Atlantic salmon Salmo salar associated with infections by the microsporidian Paranucleospora theridion","volume":"94","author":"Nylund","year":"2011","journal-title":"Dis. Aquat. Org."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"201","DOI":"10.3354\/dao02266","article-title":"Microbial and pathological findings in farmed Atlantic salmon Salmo salar with proliferative gill inflammation","volume":"91","author":"Steinum","year":"2010","journal-title":"Dis. Aquat. Org."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1007\/s00203-014-1061-4","article-title":"A new intracellular bacterium, Candidatus Similichlamydia labri sp. nov. (Chlamydiaceae) producing epitheliocysts in ballan wrasse, Labrus bergylta (Pisces, Labridae)","volume":"197","author":"Steigen","year":"2015","journal-title":"Arch. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"e82889","DOI":"10.1371\/journal.pone.0082889","article-title":"\u201cCandidatus Similichlamydia laticola\u201d, a Novel Chlamydia-like Agent of epitheliocystis in Seven Consecutive Cohorts of Farmed Australian Barramundi, Lates calcarifer (Bloch)","volume":"8","author":"Stride","year":"2013","journal-title":"PLoS ONE"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"249","DOI":"10.3354\/dao02441","article-title":"Candidatus Renichlamydia lutjani, a Gram-negative bacterium in internal organs of blue-striped snapper Lutjanus kasmira from Hawaii","volume":"98","author":"Corsaro","year":"2012","journal-title":"Dis. Aquat. Org."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"e70853","DOI":"10.1371\/journal.pone.0070853","article-title":"Candidatus Syngnamydia Venezia, a Novel Member of the Phylum Chlamydiae from the Broad Nosed Pipefish, Syngnathus typhle","volume":"8","author":"Fehr","year":"2013","journal-title":"PLoS ONE"},{"key":"ref_51","first-page":"131","article-title":"Epitheliocystis Distribution and Characterization in Brown Trout (Salmo trutta) from the Headwaters of Two Major European Rivers, the Rhine and Rhone","volume":"7","author":"Vaughan","year":"2016","journal-title":"Front. Physiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"35","DOI":"10.3354\/dao02563","article-title":"\u2018Candidatus Branchiomonas cysticola\u2019 is a common agent of epitheliocysts in seawater-farmed Atlantic salmon Salmo salar in Norway and Ireland","volume":"103","author":"Mitchell","year":"2013","journal-title":"Dis. Aquat. Org."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1556\/004.2017.003","article-title":"Diverse Chlamydia-like agents associated with epitheliocystis infection in two cyprinid fish species, the common carp (Cyprinus carpio L.) and the gibel carp (Carassius auratus gibelio L.)","volume":"65","author":"Sellyei","year":"2017","journal-title":"Acta Vet. Hung."},{"key":"ref_54","first-page":"508","article-title":"Ca. Similichlamydia in Epitheliocystis Co-infection of Gilthead Seabream Gills: Unique Morphological Features of a Deep Branching Chlamydial Family","volume":"8","author":"Katharios","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1136\/vr.101193","article-title":"Novel Chlamydiales associated with epitheliocystis in grass carp (Ctenopharyngodon idella)","volume":"172","author":"Kumar","year":"2013","journal-title":"Vet. Rec."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"75","DOI":"10.3354\/dao02255","article-title":"Novel Chlamydiales associated with epitheliocystis in a leopard shark Triakis semifasciata","volume":"91","author":"Polkinghorne","year":"2010","journal-title":"Dis. Aquat. Org."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1899","DOI":"10.1111\/1462-2920.13694","article-title":"Culture-independent genomics of a novel chlamydial pathogen of fish provides new insight into host-specific adaptations utilized by these intracellular bacteria","volume":"19","author":"Pillonel","year":"2017","journal-title":"Environ. Microbiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"13","DOI":"10.3354\/dao02586","article-title":"Epitheliocystis hyperinfection in captive spotted eagle rays Aetobatus narinari associated with a novel Chlamydiales 16S rDNA signature sequence","volume":"104","author":"Camus","year":"2013","journal-title":"Dis. Aquat. Org."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1111\/jfd.12711","article-title":"Candidatus Actinochlamydia pangasiae sp. nov. (Chlamydiales, Actinochlamydiaceae), a bacterium associated with epitheliocystis in Pangasianodon hypophthalmus","volume":"41","author":"Sood","year":"2018","journal-title":"J. Fish. Dis."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"4914","DOI":"10.1128\/AEM.00746-13","article-title":"Molecular Characterization of \u201cCandidatus Similichlamydia latridicola\u201d gen. nov., sp. nov. (Chlamydiales: \u201cCandidatus Parilichlamydiaceae\u201d), a Novel Chlamydia-Like Epitheliocystis Agent in the Striped Trumpeter, Latris lineata (Forster)","volume":"79","author":"Stride","year":"2013","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1590","DOI":"10.1128\/AEM.02899-12","article-title":"Molecular characterization of \u201cCandidatus Parilichlamydia carangidicola,\u201d a novel Chlamydia-like epitheliocystis agent in yellowtail kingfish, Seriola lalandi (Valenciennes), and the proposal of a new family \u201cCandidatus Parilichlamydiaceae\u201d fam. nov. (order Chlamydiales)","volume":"79","author":"Stride","year":"2013","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1217","DOI":"10.1111\/jfd.12814","article-title":"Investigation of co-infections with pathogens associated with gill disease in Atlantic salmon during an amoebic gill disease outbreak","volume":"41","author":"Downes","year":"2018","journal-title":"J. Fish. Dis."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1007\/s00705-008-0117-7","article-title":"Morphogenesis of salmonid gill poxvirus associated with proliferative gill disease in farmed Atlantic salmon (Salmo salar) in Norway","volume":"153","author":"Nylund","year":"2008","journal-title":"Arch. Virol."},{"key":"ref_64","doi-asserted-by":"crossref","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":"ref_65","doi-asserted-by":"crossref","first-page":"104079","DOI":"10.1016\/j.dci.2021.104079","article-title":"Mucosal immunoglobulins of teleost fish: A decade of advances","volume":"121","author":"Salinas","year":"2021","journal-title":"Dev. Comp. Immunol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Slinger, J., Adams, M.B., and Wynne, J.W. (2020). Bacteriomic Profiling of Branchial Lesions Induced by Neoparamoeba perurans Challenge Reveals Commensal Dysbiosis and an Association with Tenacibaculum dicentrarchi in AGD-Affected Atlantic Salmon (Salmo salar L.). Microorganisms, 8.","DOI":"10.3390\/microorganisms8081189"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1729","DOI":"10.1016\/j.fsi.2013.09.032","article-title":"The mucosal immune system of fish: The evolution of tolerating commensals while fighting pathogens","volume":"35","author":"Gomez","year":"2013","journal-title":"Fish Shellfish Immunol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"10728","DOI":"10.1038\/ncomms10728","article-title":"Mucosal immunoglobulins at respiratory surfaces mark an ancient association that predates the emergence of tetrapods","volume":"7","author":"Xu","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1146\/annurev.micro.62.081307.162818","article-title":"Chlamydiae as Symbionts in Eukaryotes","volume":"62","author":"Horn","year":"2008","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1111\/fwb.13437","article-title":"Microbiota associated with the skin, gills, and gut of the fish Parachondrostoma toxostoma from the Rh\u00f4ne basin","volume":"65","author":"Guivier","year":"2020","journal-title":"Freshw. Biol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"739211","DOI":"10.1016\/j.aquaculture.2022.739211","article-title":"Time is a stronger predictor of microbiome community composition than tissue in external mucosal surfaces of Atlantic salmon (Salmo salar) reared in a semi-natural freshwater environment","volume":"566","author":"Chalmers","year":"2023","journal-title":"Aquaculture"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1111\/j.1574-695X.2011.00856.x","article-title":"Permissivity of fish cell lines to three Chlamydia-related bacteria: Waddlia chondrophila, Estrella lausannensis and Parachlamydia acanthamoebae","volume":"63","author":"Batista","year":"2011","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1038\/nrg2091","article-title":"Animal models of human disease: Zebrafish swim into view","volume":"8","author":"Lieschke","year":"2007","journal-title":"Nat. Rev. Genet."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"498","DOI":"10.1038\/nature12111","article-title":"The zebrafish reference genome sequence and its relationship to the human genome","volume":"496","author":"Howe","year":"2013","journal-title":"Nature"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1038\/291293a0","article-title":"Production of clones of homozygous diploid zebra fish (Brachydanio rerio)","volume":"291","author":"Streisinger","year":"1981","journal-title":"Nature"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/j.dci.2007.11.011","article-title":"Immunology and zebrafish: Spawning new models of human disease","volume":"32","author":"Meeker","year":"2008","journal-title":"Dev. Comp. Immunol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1007\/s00248-005-0005-6","article-title":"Bacterial Community Composition in Different Sediments from the Eastern Mediterranean Sea: A Comparison of Four 16S Ribosomal DNA Clone Libraries","volume":"50","author":"Polymenakou","year":"2005","journal-title":"Microb. Ecol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"44","DOI":"10.2166\/wh.2017.106","article-title":"Presence of Waddlia chondrophila in hot water systems from non-domestic buildings in France","volume":"16","author":"Humeau","year":"2018","journal-title":"J. Water Health"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1264\/jsme2.ME12048","article-title":"Well Water as a Possible Source of Waddlia chondrophila Infections","volume":"27","author":"Codony","year":"2012","journal-title":"Microbes Environ."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"100635","DOI":"10.1016\/j.nmni.2019.100635","article-title":"Limited presence of Waddlia chondrophila in drinking water systems in the Netherlands","volume":"34","author":"Learbuch","year":"2020","journal-title":"New Microbes New Infect."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1829","DOI":"10.3389\/fmicb.2016.01829","article-title":"A Zebrafish Model for Chlamydia Infection with the Obligate Intracellular Pathogen Waddlia chondrophila","volume":"7","author":"Fehr","year":"2016","journal-title":"Front. Microbiol."},{"key":"ref_82","unstructured":"Morin, M. (2023). Evaluating a Zebrafish Model to Determine The Etiology of Epitheliocystis, University of Massachusetts Amherst."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/13\/9\/2166\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:46:58Z","timestamp":1760035618000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/13\/9\/2166"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,9,17]]},"references-count":82,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2025,9]]}},"alternative-id":["microorganisms13092166"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms13092166","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,9,17]]}}}