{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T01:43:15Z","timestamp":1792806195946,"version":"4.5.0"},"reference-count":107,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,3,11]],"date-time":"2026-03-11T00:00:00Z","timestamp":1773187200000},"content-version":"vor","delay-in-days":69,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000015","name":"US Department of Energy","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000015","id-type":"DOI","asserted-by":"publisher"}]},{"name":"University of Arizona Technology and Research Initiative Fund"},{"name":"National Science Foundation Atmospheric and Geospace Science (AGS) Atmospheric Chemistry Program","award":["1933280"],"award-info":[{"award-number":["1933280"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,14]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Carbonic anhydrase (CA) enzymes catalyze the interconversion of carbon dioxide and bicarbonate with an efficiency exceeded only by superoxide dismutase. CA enzymes have evolved convergently in phylogenetically distant organisms, forming eight structurally unrelated classes that share physiological functions involved in photosynthesis, respiration, pH homeostasis, CO2 transport, and carbonyl sulfide hydrolysis that play central roles in medicine and the environment. Here, we leverage the recent surge in publicly available genomes and metagenomes to re-examine our understanding of the abundance, diversity, and phylogenetic relationships of the three major CA classes in Bacteria\/Archaea and microbial Eukaryotes (Fungi, algae). We recovered a total of 57\u2009218 \u03b1-, \u03b2-, and \u03b3-CA sequences from 24\u2009184 metagenomes and genomes, including the first putative \u03b1-CA from an archaeal species. CA sequences formed 3859 protein clusters (1188 with three or more sequences). Sequences within a cluster were typically taxonomically conserved only at higher levels (i.e. Superkingdom, Phylum). When viewed within a phylogenetic framework, the majority of subclades for each CA class contained CAs representing multiple Superkingdoms, although numerous novel \u03b2-CA clades appear unique to Fungi. Queries of CA Hidden Markov models against all public metagenome and metatranscriptome datasets revealed that CA is a ubiquitous enzyme present in virtually all sampled environments. However, CA clusters that were taxonomically conserved also appeared more environment-specific, which may explain high CA diversity. This work represents an important contribution to our understanding of the evolution, diversity, and environmental distribution of an enzyme that is key to life and has broad environmental and industrial applications.<\/jats:p>","DOI":"10.1093\/ismeco\/ycag054","type":"journal-article","created":{"date-parts":[[2026,3,10]],"date-time":"2026-03-10T12:36:18Z","timestamp":1773146178000},"source":"Crossref","is-referenced-by-count":7,"title":["Genomic and metagenomic survey of microbial carbonic anhydrase genes reveals novel clades, high diversity, and biome specificity"],"prefix":"10.1093","volume":"6","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4959-1257","authenticated-orcid":false,"given":"Mario E E","family":"Franco","sequence":"first","affiliation":[{"name":"BIO5 Institute, University of Arizona , Tucson, AZ 85721 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3126-2199","authenticated-orcid":false,"given":"Esther","family":"Singer","sequence":"additional","affiliation":[{"name":"DOE Joint Genome Institute, Lawrence Berkeley National Laboratory , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5831-5895","authenticated-orcid":false,"given":"Simon","family":"Roux","sequence":"additional","affiliation":[{"name":"DOE Joint Genome Institute, Lawrence Berkeley National Laboratory , Berkeley, CA 94720 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4244-4366","authenticated-orcid":false,"given":"Laura K","family":"Meredith","sequence":"additional","affiliation":[{"name":"BIO5 Institute, University of Arizona , Tucson, AZ 85721 ,","place":["United States"]},{"name":"School of Natural Resources and the Environment, University of Arizona , Tucson, AZ 85721 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7608-5029","authenticated-orcid":false,"given":"Jana M","family":"U\u2019Ren","sequence":"additional","affiliation":[{"name":"Department of Plant Pathology, Washington State University , Pullman, WA 99164, United States"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,3,11]]},"reference":[{"key":"2026042109463890200_ref1","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1146\/annurev.ph.58.030196.002515","article-title":"Multiple roles of carbonic anhydrase in cellular transport and metabolism","volume":"58","author":"Henry","year":"1996","journal-title":"Annu Rev Physiol"},{"key":"2026042109463890200_ref2","doi-asserted-by":"publisher","first-page":"15184","DOI":"10.1073\/pnas.96.26.15184","article-title":"Carbonic anhydrase is an ancient enzyme widespread in prokaryotes","volume":"96","author":"Smith","year":"1999","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026042109463890200_ref3","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1038\/nrd2467","article-title":"Carbonic anhydrases: novel therapeutic applications for inhibitors and activators","volume":"7","author":"Supuran","year":"2008","journal-title":"Nat Rev Drug Discov"},{"key":"2026042109463890200_ref4","doi-asserted-by":"publisher","first-page":"325","DOI":"10.3109\/14756366.2014.910202","article-title":"An overview of the alpha-, beta- and gamma-carbonic anhydrases from bacteria: can bacterial carbonic anhydrases shed new light on evolution of bacteria?","volume":"30","author":"Capasso","year":"2015","journal-title":"J Enzyme Inhib Med Chem"},{"key":"2026042109463890200_ref5","doi-asserted-by":"publisher","first-page":"9828","DOI":"10.1073\/pnas.1603112113","article-title":"Thylakoid luminal \u03b8-carbonic anhydrase critical for growth and photosynthesis in the marine diatom Phaeodactylum tricornutum","volume":"113","author":"Kikutani","year":"2016","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026042109463890200_ref6","first-page":"165","volume-title":"Advances in Agronomy","author":"Sharififar","year":"2023"},{"key":"2026042109463890200_ref7","doi-asserted-by":"publisher","first-page":"114104","DOI":"10.1016\/j.geoderma.2019.114104","article-title":"Desert soil sequesters atmospheric CO2 by microbial mineral formation","volume":"361","author":"Liu","year":"2020","journal-title":"Geoderma"},{"key":"2026042109463890200_ref8","doi-asserted-by":"publisher","first-page":"100008","DOI":"10.1016\/j.bgtech.2023.100008","article-title":"Applications of microbial-induced carbonate precipitation: a state-of-the-art review","volume":"1","author":"Wang","year":"2023","journal-title":"Biogeotechnics"},{"key":"2026042109463890200_ref9","doi-asserted-by":"publisher","first-page":"2427","DOI":"10.5194\/bg-19-2427-2022","article-title":"Global modelling of soil carbonyl sulfide exchanges","volume":"19","author":"Abadie","year":"2022","journal-title":"Biogeosciences"},{"key":"2026042109463890200_ref10","doi-asserted-by":"publisher","first-page":"D09302","DOI":"10.1029\/2006JD007665","article-title":"On the global distribution, seasonality, and budget of atmospheric carbonyl sulfide (COS) and some similarities to CO2","volume":"112","author":"Montzka","year":"2007","journal-title":"J Geophys Res"},{"key":"2026042109463890200_ref11","doi-asserted-by":"publisher","first-page":"e0201523","DOI":"10.1128\/aem.02015-23","article-title":"Sulfur assimilation using gaseous carbonyl sulfide by the soil fungus Trichoderma harzianum","volume":"90","author":"Iizuka","year":"2024","journal-title":"Appl Environ Microbiol"},{"key":"2026042109463890200_ref12","doi-asserted-by":"publisher","first-page":"106527","DOI":"10.1016\/j.ympev.2019.106527","article-title":"Beneath the surface: evolution of methane activity in the bacterial multicomponent monooxygenases","volume":"139","author":"Osborne","year":"2019","journal-title":"Mol Phylogenet Evol"},{"key":"2026042109463890200_ref13","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1111\/j.1574-6976.2000.tb00546.x","article-title":"Prokaryotic carbonic anhydrases","volume":"24","author":"Smith","year":"2000","journal-title":"FEMS Microbiol Rev"},{"key":"2026042109463890200_ref14","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1016\/j.bmc.2013.10.045","article-title":"Cloning, characterization and anion inhibition study of the \u03b4-class carbonic anhydrase (TweCA) from the marine diatom Thalassiosira weissflogii","volume":"22","author":"Del Prete","year":"2014","journal-title":"Bioorg Med Chem"},{"key":"2026042109463890200_ref15","doi-asserted-by":"publisher","first-page":"4389","DOI":"10.1016\/j.bmcl.2014.08.015","article-title":"Discovery of a new family of carbonic anhydrases in the malaria pathogen Plasmodium falciparum\u2014the \u03b7-carbonic anhydrases","volume":"24","author":"Del Prete","year":"2014","journal-title":"Bioorg Med Chem Lett"},{"key":"2026042109463890200_ref16","doi-asserted-by":"publisher","first-page":"1060","DOI":"10.1080\/14756366.2020.1755852","article-title":"Bacterial \u03b9-carbonic anhydrase: a new active class of carbonic anhydrase identified in the genome of the gram-negative bacterium Burkholderia territorii","volume":"35","author":"Del Prete","year":"2020","journal-title":"J Enzyme Inhib Med Chem"},{"key":"2026042109463890200_ref17","doi-asserted-by":"publisher","first-page":"129411","DOI":"10.1016\/j.bmcl.2023.129411","article-title":"A simple yet multifaceted 90 years old, evergreen enzyme: carbonic anhydrase, its inhibition and activation","volume":"93","author":"Supuran","year":"2023","journal-title":"Bioorg Med Chem Lett"},{"key":"2026042109463890200_ref18","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1016\/j.compbiolchem.2016.01.003","article-title":"On origin and evolution of carbonic anhydrase isozymes: a phylogenetic analysis from whole-enzyme to active site","volume":"61","author":"Banerjee","year":"2016","journal-title":"Comput Biol Chem"},{"key":"2026042109463890200_ref19","doi-asserted-by":"publisher","first-page":"8466","DOI":"10.1128\/AEM.05702-11","article-title":"Carboxylases in natural and synthetic microbial pathways","volume":"77","author":"Erb","year":"2011","journal-title":"Appl Environ Microbiol"},{"key":"2026042109463890200_ref20","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1186\/s13567-018-0583-1","article-title":"The CO2-dependence of Brucella ovis and Brucella abortus biovars is caused by defective carbonic anhydrases","volume":"49","author":"P\u00e9rez-Etayo","year":"2018","journal-title":"Vet Res"},{"key":"2026042109463890200_ref21","doi-asserted-by":"publisher","first-page":"324549","DOI":"10.1155\/2012\/324549","article-title":"Dispensabilities of carbonic anhydrase in proteobacteria","volume":"2012","author":"Ueda","year":"2012","journal-title":"Int J Evol Biol"},{"key":"2026042109463890200_ref22","doi-asserted-by":"publisher","DOI":"10.3390\/biology12060770","article-title":"Genome study of \u03b1-, \u03b2-, and \u03b3-carbonic anhydrases from the thermophilic microbiome of marine hydrothermal vent ecosystems","volume":"12","author":"Gheibzadeh","year":"2023","journal-title":"Biology"},{"key":"2026042109463890200_ref23","doi-asserted-by":"publisher","first-page":"737","DOI":"10.1517\/13543776.2013.798648","article-title":"Anticancer carbonic anhydrase inhibitors: a patent review (2008 \u2013 2013)","volume":"23","author":"Monti","year":"2013","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref24","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1517\/13543776.2013.780598","article-title":"Diuretics with carbonic anhydrase inhibitory action: a patent and literature review (2005\u20132013)","volume":"23","author":"Carta","year":"2013","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref25","doi-asserted-by":"publisher","first-page":"705","DOI":"10.1517\/13543776.2013.794788","article-title":"Antiglaucoma carbonic anhydrase inhibitors: a patent review","volume":"23","author":"Masini","year":"2013","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref26","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1517\/13543776.2013.782394","article-title":"Anticonvulsant\/antiepileptic carbonic anhydrase inhibitors: a patent review","volume":"23","author":"Aggarwal","year":"2013","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref27","doi-asserted-by":"publisher","first-page":"855","DOI":"10.2174\/156802607780636726","article-title":"Carbonic anhydrase inhibitors as anticonvulsant agents","volume":"7","author":"Thiry","year":"2007","journal-title":"Curr Top Med Chem"},{"key":"2026042109463890200_ref28","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1517\/13543776.2013.790957","article-title":"Antiobesity carbonic anhydrase inhibitors: a literature and patent review","volume":"23","author":"Scozzafava","year":"2013","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref29","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1080\/13543776.2018.1523897","article-title":"Carbonic anhydrase inhibitors and their potential in a range of therapeutic areas","volume":"28","author":"Supuran","year":"2018","journal-title":"Expert Opin Ther Pat"},{"key":"2026042109463890200_ref30","doi-asserted-by":"publisher","first-page":"1233","DOI":"10.1042\/CS20210040","article-title":"Emerging role of carbonic anhydrase inhibitors","volume":"135","author":"Supuran","year":"2021","journal-title":"Clin Sci"},{"key":"2026042109463890200_ref31","doi-asserted-by":"publisher","first-page":"4324","DOI":"10.3390\/ijms22094324","article-title":"A highlight on the inhibition of fungal carbonic anhydrases as drug targets for the antifungal armamentarium","volume":"22","author":"Supuran","year":"2021","journal-title":"Int J Mol Sci"},{"key":"2026042109463890200_ref32","doi-asserted-by":"publisher","first-page":"1689","DOI":"10.1517\/14728222.2015.1067685","article-title":"Bacterial, fungal and protozoan carbonic anhydrases as drug targets","volume":"19","author":"Capasso","year":"2015","journal-title":"Expert Opin Ther Targets"},{"key":"2026042109463890200_ref33","doi-asserted-by":"publisher","first-page":"60","DOI":"10.1016\/j.hal.2014.12.006","article-title":"Metatranscriptome profiling reveals versatile N-nutrient utilization, CO2 limitation, oxidative stress, and active toxin production in an Alexandrium fundyense bloom","volume":"42","author":"Zhuang","year":"2015","journal-title":"Harmful Algae"},{"key":"2026042109463890200_ref34","doi-asserted-by":"publisher","first-page":"2094","DOI":"10.1038\/s41396-019-0426-8","article-title":"A new widespread subclass of carbonic anhydrase in marine phytoplankton","volume":"13","author":"Jensen","year":"2019","journal-title":"ISME J"},{"key":"2026042109463890200_ref35","doi-asserted-by":"publisher","first-page":"3196","DOI":"10.1111\/1462-2920.15545","article-title":"Initiation of efficient C4 pathway in response to low ambient CO2 during the bloom period of a marine dinoflagellate","volume":"23","author":"Zhang","year":"2021","journal-title":"Environ Microbiol"},{"key":"2026042109463890200_ref36","doi-asserted-by":"publisher","volume-title":"Appl Environ Microbiol","author":"Gr\u00fcterich","DOI":"10.1128\/aem.02177-24"},{"key":"2026042109463890200_ref37","doi-asserted-by":"publisher","first-page":"n\/a","DOI":"10.1264\/jsme2.ME20058","article-title":"Fungal carbonyl sulfide hydrolase of Trichoderma harzianum strain THIF08 and its relationship with clade D \u03b2-carbonic anhydrases","volume":"36","author":"Masaki","year":"2021","journal-title":"Microbes Environ"},{"key":"2026042109463890200_ref38","doi-asserted-by":"publisher","first-page":"3818","DOI":"10.1021\/ja307735e","article-title":"Carbonyl sulfide hydrolase from Thiobacillus thioparus strain THI115 is one of the \u03b2-carbonic anhydrase family enzymes","volume":"135","author":"Ogawa","year":"2013","journal-title":"J Am Chem Soc"},{"key":"2026042109463890200_ref39","doi-asserted-by":"publisher","first-page":"842","DOI":"10.1002\/jgrg.20068","article-title":"A coupled model of the global cycles of carbonyl sulfide and CO2: a possible new window on the carbon cycle","volume":"118","author":"Berry","year":"2013","journal-title":"J Geophys Res Biogeosci"},{"key":"2026042109463890200_ref40","doi-asserted-by":"publisher","first-page":"11577","DOI":"10.1029\/1999JD900090","article-title":"Controlling variables for the uptake of atmospheric carbonyl sulfide by soil","volume":"104","author":"Kesselmeier","year":"1999","journal-title":"J Geophys Res"},{"key":"2026042109463890200_ref41","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1016\/j.soilbio.2017.09.009","article-title":"The interaction of soil phototrophs and fungi with pH and their impact on soil CO2, CO18O and OCS exchange","volume":"115","author":"Sauze","year":"2017","journal-title":"Soil Biol Biochem"},{"key":"2026042109463890200_ref42","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1038\/s41396-018-0270-2","article-title":"Soil exchange rates of COS and CO18O differ with the diversity of microbial communities and their carbonic anhydrase enzymes","volume":"13","author":"Meredith","year":"2019","journal-title":"ISME J"},{"key":"2026042109463890200_ref43","doi-asserted-by":"publisher","first-page":"121","DOI":"10.5194\/soil-5-121-2019","article-title":"Microbial community responses determine how soil\u2013atmosphere exchange of carbonyl sulfide, carbon monoxide, and nitric oxide responds to soil moisture","volume":"5","author":"Behrendt","year":"2019","journal-title":"Soil"},{"key":"2026042109463890200_ref44","doi-asserted-by":"publisher","DOI":"10.1093\/femsle\/fnw197","article-title":"Degradation and emission of carbonyl sulfide, an atmospheric trace gas, by fungi isolated from forest soil","volume":"363","author":"Masaki","year":"2016","journal-title":"FEMS Microbiol Lett"},{"key":"2026042109463890200_ref45","doi-asserted-by":"publisher","DOI":"10.1093\/femsle\/fnw223","article-title":"Degradation of carbonyl sulfide by Actinomycetes and detection of clade D of \u03b2-class carbonic anhydrase","volume":"363","author":"Ogawa","year":"2016","journal-title":"FEMS Microbiol Lett"},{"key":"2026042109463890200_ref46","doi-asserted-by":"publisher","first-page":"1343","DOI":"10.1002\/2016JG003678","article-title":"Exchange of carbonyl sulfide (OCS) between soils and atmosphere under various CO2 concentrations","volume":"122","author":"Bunk","year":"2017","journal-title":"J Geophys Res Biogeosci"},{"key":"2026042109463890200_ref47","doi-asserted-by":"publisher","first-page":"62","DOI":"10.3390\/soilsystems2040062","article-title":"Nitrogen fertilization reduces the capacity of soils to take up atmospheric carbonyl sulphide","volume":"2","author":"Kaisermann","year":"2018","journal-title":"Soil Syst"},{"key":"2026042109463890200_ref48","doi-asserted-by":"publisher","first-page":"3093","DOI":"10.1111\/j.1462-2920.2008.01735.x","article-title":"The effects of chronic nitrogen fertilization on alpine tundra soil microbial communities: implications for carbon and nitrogen cycling","volume":"10","author":"Nemergut","year":"2008","journal-title":"Environ Microbiol"},{"key":"2026042109463890200_ref49","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1002\/2015JG003149","article-title":"Litter dominates surface fluxes of carbonyl sulfide in a Californian oak woodland: surface COS fluxes in an oak woodland","volume":"121","author":"Sun","year":"2016","journal-title":"J Geophys Res Biogeosci"},{"key":"2026042109463890200_ref50","doi-asserted-by":"publisher","first-page":"6363","DOI":"10.5194\/hess-21-6363-2017","article-title":"Non-destructive estimates of soil carbonic anhydrase activity and associated soil water oxygen isotope composition","volume":"21","author":"Jones","year":"2017","journal-title":"Hydrol Earth Syst Sci"},{"key":"2026042109463890200_ref51","doi-asserted-by":"publisher","first-page":"145","DOI":"10.5194\/soil-7-145-2021","article-title":"Oxygen isotope exchange between water and carbon dioxide in soils is controlled by pH, nitrate and microbial biomass through links to carbonic anhydrase activity","volume":"7","author":"Jones","year":"2021","journal-title":"Soil"},{"key":"2026042109463890200_ref52","doi-asserted-by":"publisher","first-page":"597","DOI":"10.5194\/bg-15-597-2018","article-title":"The role of soil pH on soil carbonic anhydrase activity","volume":"15","author":"Sauze","year":"2018","journal-title":"Biogeosciences"},{"key":"2026042109463890200_ref53","doi-asserted-by":"publisher","DOI":"10.1029\/2006JG000177","article-title":"Diurnally variable \u03b418O signatures of soil CO2 fluxes indicate carbonic anhydrase activity in a forest soil","volume":"111","author":"Seibt","year":"2006","journal-title":"J Geophys Res"},{"key":"2026042109463890200_ref54","doi-asserted-by":"publisher","first-page":"2178","DOI":"10.1111\/j.1365-2486.2008.01635.x","article-title":"Evaporation and carbonic anhydrase activity recorded in oxygen isotope signatures of net CO2 fluxes from a Mediterranean soil","volume":"14","author":"Wingate","year":"2008","journal-title":"Glob Chang Biol"},{"key":"2026042109463890200_ref55","doi-asserted-by":"publisher","first-page":"22411","DOI":"10.1073\/pnas.0905210106","article-title":"The impact of soil microorganisms on the global budget of delta18O in atmospheric CO2","volume":"106","author":"Wingate","year":"2009","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026042109463890200_ref56","doi-asserted-by":"publisher","first-page":"566","DOI":"10.1111\/ejss.12241","article-title":"The effect of soil moisture, soil particle size, litter layer and carbonic anhydrase on the oxygen isotopic composition of soil-released CO2: oxygen isotopic composition of soil-released CO2","volume":"66","author":"Von Sperber","year":"2015","journal-title":"Eur J Soil Sci"},{"key":"2026042109463890200_ref57","doi-asserted-by":"publisher","DOI":"10.1016\/j.tim.2025.08.001","article-title":"Volatile traits expand the microbial playbook","volume":"34","author":"Ledford","journal-title":"Trends Microbiol"},{"key":"2026042109463890200_ref58","doi-asserted-by":"publisher","first-page":"D261","DOI":"10.1093\/nar\/gku1223","article-title":"Expanded microbial genome coverage and improved protein family annotation in the COG database","volume":"43","author":"Galperin","year":"2015","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref59","doi-asserted-by":"publisher","first-page":"D115","DOI":"10.1093\/nar\/gkr1044","article-title":"IMG: the integrated microbial genomes database and comparative analysis system","volume":"40","author":"Markowitz","year":"2012","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref60","doi-asserted-by":"publisher","first-page":"D699","DOI":"10.1093\/nar\/gkt1183","article-title":"MycoCosm portal: gearing up for 1000 fungal genomes","volume":"42","author":"Grigoriev","year":"2014","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref61","doi-asserted-by":"publisher","first-page":"D1004","DOI":"10.1093\/nar\/gkaa898","article-title":"PhycoCosm, a comparative algal genomics resource","volume":"49","author":"Grigoriev","year":"2021","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref62","doi-asserted-by":"publisher","first-page":"D427","DOI":"10.1093\/nar\/gky995","article-title":"The Pfam protein families database in 2019","volume":"47","author":"El-Gebali","year":"2019","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref63","doi-asserted-by":"publisher","first-page":"1026","DOI":"10.1038\/nbt.3988","article-title":"MMseqs2 enables sensitive protein sequence searching for the analysis of massive data sets","volume":"35","author":"Steinegger","year":"2017","journal-title":"Nat Biotechnol"},{"key":"2026042109463890200_ref64","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","article-title":"MAFFT multiple sequence alignment software version 7: improvements in performance and usability","volume":"30","author":"Katoh","year":"2013","journal-title":"Mol Biol Evol"},{"key":"2026042109463890200_ref65","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1101\/gr.5969107","article-title":"MEGAN analysis of metagenomic data","volume":"17","author":"Huson","year":"2007","journal-title":"Genome Res"},{"issue":"49","key":"2026042109463890200_ref66","doi-asserted-by":"publisher","first-page":"D480","DOI":"10.1093\/nar\/gkaa1100","article-title":"UniProt: the universal protein knowledgebase in 2021","volume":"2021","author":"The UniProt Consortium","journal-title":"Nucleic Acids Res"},{"key":"2026042109463890200_ref67","doi-asserted-by":"publisher","first-page":"1972","DOI":"10.1093\/bioinformatics\/btp348","article-title":"trimAl: a tool for automated alignment trimming in large-scale phylogenetic analyses","volume":"25","author":"Capella-Guti\u00e9rrez","year":"2009","journal-title":"Bioinformatics"},{"key":"2026042109463890200_ref68","doi-asserted-by":"publisher","first-page":"1647","DOI":"10.1093\/bioinformatics\/bts199","article-title":"Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data","volume":"28","author":"Kearse","year":"2012","journal-title":"Bioinformatics"},{"key":"2026042109463890200_ref69","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1093\/embo-reports\/kvd016","article-title":"A wheel invented three times","volume":"1","author":"Liljas","year":"2000","journal-title":"EMBO Rep"},{"key":"2026042109463890200_ref70","doi-asserted-by":"publisher","first-page":"2023","DOI":"10.1042\/BCJ20160115","article-title":"Structure and function of carbonic anhydrases","volume":"473","author":"Supuran","year":"2016","journal-title":"Biochem J"},{"key":"2026042109463890200_ref71","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1006\/mpev.1996.0006","article-title":"Functional diversity, conservation, and convergence in the evolution of the alpha-, beta-, and gamma-carbonic anhydrase gene families","volume":"5","author":"Hewett-Emmett","year":"1996","journal-title":"Mol Phylogenet Evol"},{"key":"2026042109463890200_ref72","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/molbev\/msu300","article-title":"IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies","volume":"32","author":"Nguyen","year":"2015","journal-title":"Mol Biol Evol"},{"key":"2026042109463890200_ref73","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1038\/nmeth.4285","article-title":"ModelFinder: fast model selection for accurate phylogenetic estimates","volume":"14","author":"Kalyaanamoorthy","year":"2017","journal-title":"Nat Methods"},{"key":"2026042109463890200_ref74","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1093\/molbev\/msx281","article-title":"UFBoot2: improving the ultrafast bootstrap approximation","volume":"35","author":"Hoang","year":"2018","journal-title":"Mol Biol Evol"},{"key":"2026042109463890200_ref75","doi-asserted-by":"publisher","first-page":"1188","DOI":"10.1093\/molbev\/mst024","article-title":"Ultrafast approximation for phylogenetic bootstrap","volume":"30","author":"Minh","year":"2013","journal-title":"Mol Biol Evol"},{"key":"2026042109463890200_ref76","article-title":"IQ-TREE version 2.1.2: tutorials and manual, phylogenomic software by maximum likelihood","author":"Minh"},{"key":"2026042109463890200_ref77","volume-title":"caper: Comparative Analyses of Phylogenetics and Evolution in R","author":"Orme"},{"key":"2026042109463890200_ref78","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/j.ympev.2016.02.010","article-title":"Contributions of North American endophytes to the phylogeny, ecology, and taxonomy of Xylariaceae (Sordariomycetes, Ascomycota)","volume":"98","author":"U\u2019Ren","year":"2016","journal-title":"Mol Phylogenet Evol"},{"key":"2026042109463890200_ref79","volume-title":"pheatmap: Pretty Heatmaps","author":"Kolde"},{"key":"2026042109463890200_ref80","doi-asserted-by":"publisher","first-page":"7","DOI":"10.3390\/metabo7040056","article-title":"An overview of the bacterial carbonic anhydrases","volume":"7","author":"Supuran","year":"2017","journal-title":"Metabolites"},{"key":"2026042109463890200_ref81","doi-asserted-by":"publisher","first-page":"170037","DOI":"10.1038\/sdata.2017.37","article-title":"Metagenome sequencing and 98 microbial genomes from Juan de Fuca ridge flank subsurface fluids","volume":"4","author":"Jungbluth","year":"2017","journal-title":"Sci Data"},{"key":"2026042109463890200_ref82","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/s00294-009-0238-x","article-title":"Evolution of carbonic anhydrases in fungi","volume":"55","author":"Elleuche","year":"2009","journal-title":"Curr Genet"},{"key":"2026042109463890200_ref83","doi-asserted-by":"publisher","first-page":"20160352","DOI":"10.1098\/rsta.2016.0352","article-title":"Resurrecting ancestral genes in bacteria to interpret ancient biosignatures","volume":"375","author":"Kacar","year":"2017","journal-title":"Philos Trans A Math Phys Eng Sci"},{"key":"2026042109463890200_ref84","doi-asserted-by":"publisher","first-page":"1254","DOI":"10.1080\/14756366.2016.1201479","article-title":"New light on bacterial carbonic anhydrases phylogeny based on the analysis of signal peptide sequences","volume":"31","author":"Supuran","year":"2016","journal-title":"J Enzyme Inhib Med Chem"},{"key":"2026042109463890200_ref85","doi-asserted-by":"publisher","first-page":"4065","DOI":"10.1007\/s00253-022-11990-3","article-title":"The production and biochemical characterization of \u03b1-carbonic anhydrase from Lactobacillus rhamnosus GG","volume":"106","author":"Urba\u0144ski","year":"2022","journal-title":"Appl Microbiol Biotechnol"},{"key":"2026042109463890200_ref86","doi-asserted-by":"publisher","volume-title":"Bergey's Manual of Systematics of Archaea and Bacteria","author":"Huber","DOI":"10.1002\/9781118960608.gbm00479"},{"key":"2026042109463890200_ref87","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1016\/S0723-2020(11)80176-1","article-title":"Archaeoglobus profundus sp. nov., represents a new species within the sulfate-reducing Archaebacteria","volume":"13","author":"Burggraf","year":"1990","journal-title":"Syst Appl Microbiol"},{"key":"2026042109463890200_ref88","doi-asserted-by":"publisher","first-page":"1401","DOI":"10.1126\/science.1089370","article-title":"The origins of genome complexity","volume":"302","author":"Lynch","year":"2003","journal-title":"Science"},{"key":"2026042109463890200_ref89","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/j.tig.2008.08.005","article-title":"Widespread conservation of genetic redundancy during a billion years of eukaryotic evolution","volume":"24","author":"Vavouri","year":"2008","journal-title":"Trends Genet"},{"key":"2026042109463890200_ref90","doi-asserted-by":"publisher","first-page":"1016","DOI":"10.1016\/j.cub.2005.04.035","article-title":"Gene complexity and gene duplicability","volume":"15","author":"He","year":"2005","journal-title":"Curr Biol"},{"key":"2026042109463890200_ref91","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1007\/s00239-008-9191-4","article-title":"Symbiobacterium lost carbonic anhydrase in the course of evolution","volume":"68","author":"Nishida","year":"2009","journal-title":"J Mol Evol"},{"key":"2026042109463890200_ref92","author":"Lehneck"},{"key":"2026042109463890200_ref93","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1111\/mmi.12607","article-title":"The filamentous ascomycete Sordaria macrospora can survive in ambient air without carbonic anhydrases","volume":"92","author":"Lehneck","year":"2014","journal-title":"Mol Microbiol"},{"key":"2026042109463890200_ref94","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1128\/MMBR.00015-07","article-title":"Function, structure, and evolution of the RubisCO-like proteins and their RubisCO homologs","volume":"71","author":"Tabita","year":"2007","journal-title":"Microbiol Mol Biol Rev"},{"key":"2026042109463890200_ref95","doi-asserted-by":"publisher","first-page":"2223","DOI":"10.1073\/pnas.1310811111","article-title":"Stability-activity tradeoffs constrain the adaptive evolution of RubisCO","volume":"111","author":"Studer","year":"2014","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026042109463890200_ref96","doi-asserted-by":"publisher","first-page":"1988","DOI":"10.1080\/14756366.2021.1972995","article-title":"An overview on the recently discovered iota-carbonic anhydrases","volume":"36","author":"Nocentini","year":"2021","journal-title":"J Enzyme Inhib Med Chem"},{"key":"2026042109463890200_ref97","doi-asserted-by":"publisher","first-page":"830","DOI":"10.1038\/ismej.2012.160","article-title":"Phylogenetic conservatism of functional traits in microorganisms","volume":"7","author":"Martiny","year":"2013","journal-title":"ISME J"},{"key":"2026042109463890200_ref98","doi-asserted-by":"publisher","first-page":"965","DOI":"10.1007\/s00294-017-0700-0","article-title":"CO2 sensing in fungi: at the heart of metabolic signaling","volume":"63","author":"Martin","year":"2017","journal-title":"Curr Genet"},{"key":"2026042109463890200_ref99","doi-asserted-by":"publisher","first-page":"935","DOI":"10.1080\/21505594.2019.1688753","article-title":"Antivirulence and avirulence genes in human pathogenic fungi","volume":"10","author":"Siscar-Lewin","year":"2019","journal-title":"Virulence"},{"key":"2026042109463890200_ref100","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.jsb.2019.07.011","article-title":"The structural basis of the low catalytic activities of the two minor \u03b2-carbonic anhydrases of the filamentous fungus Aspergillus fumigatus","volume":"208","author":"Kim","year":"2019","journal-title":"J Struct Biol"},{"key":"2026042109463890200_ref101","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1007\/978-3-642-19974-5_11","volume-title":"Evolution of Fungi and Fungal-Like Organisms","author":"Elleuche","year":"2011"},{"key":"2026042109463890200_ref102","doi-asserted-by":"publisher","first-page":"8433","DOI":"10.1007\/s00253-014-5993-z","article-title":"A matter of structure: structural comparison of fungal carbonic anhydrases","volume":"98","author":"Lehneck","year":"2014","journal-title":"Appl Microbiol Biotechnol"},{"key":"2026042109463890200_ref103","doi-asserted-by":"publisher","first-page":"3041","DOI":"10.1099\/00221287-134-11-3041","article-title":"Oxidation of carbon disulphide as the sole source of energy for the autotrophic growth of Thiobacillus thioparus strain TK-m","volume":"134","author":"Smith","year":"1988","journal-title":"Microbiology"},{"key":"2026042109463890200_ref104","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1099\/mic.0.2007\/011213-0","article-title":"Degradation of ambient carbonyl sulfide by Mycobacterium spp. in soil","volume":"154","author":"Kato","year":"2008","journal-title":"Microbiology"},{"key":"2026042109463890200_ref105","doi-asserted-by":"publisher","first-page":"104","DOI":"10.3389\/fmicb.2011.00104","article-title":"Mycobacteria isolated from angkor monument sandstones grow chemolithoautotrophically by oxidizing elemental sulfur","volume":"2","author":"Kusumi","year":"2011","journal-title":"Front Microbiol"},{"key":"2026042109463890200_ref106","doi-asserted-by":"publisher","first-page":"4046","DOI":"10.1128\/JB.00627-13","article-title":"Bacterial CS2 hydrolases from Acidithiobacillus thiooxidans strains are homologous to the archaeal catenane CS2 hydrolase","volume":"195","author":"Smeulders","year":"2013","journal-title":"J Bacteriol"},{"key":"2026042109463890200_ref107","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1007\/s00248-010-9699-1","article-title":"Oxidation of elemental sulfur by Fusarium solani strain THIF01 harboring endobacterium Bradyrhizobium sp","volume":"60","author":"Li","year":"2010","journal-title":"Microb Ecol"}],"container-title":["ISME Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/ismecommun\/advance-article-pdf\/doi\/10.1093\/ismeco\/ycag054\/67298640\/ycag054.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismecommun\/article-pdf\/6\/1\/ycag054\/67298640\/ycag054.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismecommun\/article-pdf\/6\/1\/ycag054\/67298640\/ycag054.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,21]],"date-time":"2026-04-21T13:46:45Z","timestamp":1776779205000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/ismecommun\/article\/doi\/10.1093\/ismeco\/ycag054\/8514074"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1]]},"references-count":107,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,1,14]]}},"URL":"https:\/\/doi.org\/10.1093\/ismeco\/ycag054","relation":{},"ISSN":["2730-6151"],"issn-type":[{"value":"2730-6151","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1]]},"published":{"date-parts":[[2026,1]]},"article-number":"ycag054"}}