PROF. SINA ADL (Orcid ID : 0000-0001-6324-6065) PROF. DAVID BASS (Orcid ID : 0000-0002-9883-7823) DR. CÉDRIC BERNEY (Orcid ID : 0000-0001-8689-9907) DR. PACO CÁRDENAS (Orcid ID : 0000-0003-4045-6718) DR. IVAN CEPICKA (Orcid ID : 0000-0002-4322-0754) DR. MICAH DUNTHORN (Orcid ID : 0000-0003-1376-4109) PROF. BENTE EDVARDSEN (Orcid ID : 0000-0002-6806-4807) DR. DENIS H. LYNN (Orcid ID : 0000-0002-1554-7792) DR. EDWARD A.D MITCHELL (Orcid ID : 0000-0003-0358-506X) PROF. JONG SOO PARK (Orcid ID : 0000-0001-6253-5199) DR. GUIFRÉ TORRUELLA (Orcid ID : 0000-0002-6534-4758) DR. VASILY V. ZLATOGURSKY (Orcid ID : 0000-0002-2688-3900) Article type : Original Article Corresponding author mail id: [email protected] Adl et al.---Classification of Eukaryotes Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes Sina M. Adla, David Bassb,c, Christopher E. Laned, Julius Lukeše,f, Conrad L. Schochg, Alexey Smirnovh, Sabine Agathai, Cedric Berneyj, Matthew W. Brownk,l, Fabien Burkim, Paco Cárdenasn, o p q r,s t Ivan Čepička , Ludmila Chistyakova , Javier del Campo , Micah Dunthorn , Bente Edvardsen , Yana u v w x y z Eglit , Laure Guillou , VladimírAuthor Manuscript Hampl , Aaron A. Heiss , Mona Hoppenrath , Timothy Y. James , Sergey Karpovh, Eunsoo Kimx, Martin Koliskoe, Alexander Kudryavtsevh,aa, Daniel J. G. Lahrab, Enrique Laraac,ad, Line Le Gallae, Denis H. Lynnaf,ag, David G. Mannah, Ramon Massana i Moleraq, This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as doi: 10.1111/jeu.12691-5149 This article is protected by copyright. All rights reserved Edward A. D. Mitchellac,ai , Christine Morrowaj, Jong Soo Parkak, Jan W. Pawlowskial, Martha J. Powellam, Daniel J. Richteran, Sonja Rueckertao, Lora Shadwickap, Satoshi Shimanoaq, Frederick W. Spiegelap, Guifré Torruella i Cortesar, Noha Youssefas, Vasily Zlatogurskyh,at, Qianqian Zhangau,av. a Department of Soil Sciences, College of Agriculture and Bioresources, University of Saskatchewan, Saskatoon, SK Canada, and b Dept of Life Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, and c Centre for Environment, Fisheries and Aquaculture Science (CEFAS), Barrack Road, The Nothe, Weymouth, Dorset DT4 8UB, and d Department of Biological Sciences, University of Rhode Island, Kingston, Rhode Island, 02881 USA, and e Institute of Parasitology, Biology Centre, Czech Academy of Sciences, 37005 České Budějovice, Czechia, and f Faculty of Science, University of South Bohemia, 37005 České Budějovice, Czechia, and gNational Institute for Biotechnology Information, National Library of Medicine, National Institutes of Health, Bethesda, MD 20892, USA, and hDepartment of Invertebrate Zoology, Faculty of Biology, Saint Petersburg State University, 199034 Saint Petersburg, Russia, and iDepartment of Biosciences, University of Salzburg, Hellbrunnerstrasse 34, A-5020 Salzburg, Austria, and jCNRS, UMR 7144 (AD2M), Groupe Evolution des Protistes et Ecosystèmes Pélagiques, Station Biologique de Roscoff, Place Georges Teissier, Roscoff 29680, France, and kDepartment of Biological Sciences, Mississippi State University, MS USA, and lInstitute for Genomics, Biocomputing & Biotechnology, Mississippi State University, MS USA, and mDepartment of Organismal Biology, Program in Systematic Biology, Science for Life Laboratory, Uppsala University, Uppsala 75236, Sweden, and nPharmacognosy, Department of Medicinal Chemistry, Uppsala University, BMC Box 574, SE-75123 Uppsala, Sweden, and oDepartment of Zoology, Faculty of Science, Charles University, Vinicna 7, 128 44 Prague, Czechia, and pCore Facility Centre for Culture Collection of Microorganisms, Saint Petersburg State University, 198504 Saint Petersburg, Russia, and q Institut de Ciències del Mar, CSIC, Passeig Marítim de la Barceloneta, 37-49 08003 Barcelona, Catalonia, Spain, and rDepartment of Ecology, University of Kaiserslautern, Erwin-Schroedinger Street, D-67663 Kaiserslautern, Germany, and sDepartment of Eukaryotic Microbiology, University of Duisburg-Essen, Universitätsstrasse 5, D-45141 Essen, Germany, and tDepartment of Biosciences, University of Oslo, P.O. Box 1066 Blindern, 0316 Oslo, Norway, and uDepartment of Biology, Dalhousie University, Halifax, NS, Canada, and vSorbonne Université, Université Pierre et Marie Curie - Paris 6, CNRS, UMR 7144 (AD2M), Station Biologique de Roscoff, Place Georges Teissier, CS90074, 29688 Roscoff, France, and w Author Manuscript Department of Parasitology, Faculty of Science, Charles University, BIOCEV, Průmyslová 595, 252 42 Vestec, Czechia, and xDepartment of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024 USA, and ySenckenberg am Meer, DZMB – German Centre for Marine Biodiversity Research, D-26382 Wilhelmshaven, Germany, and z Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI 48109 USA, and This article is protected by copyright. All rights reserved aaLaboratory of Parasitic Worms and Protistology, Zoological Institute RAS, 199034 Saint Petersburg, Russia, and abDepartment of Zoology, Institute of Biosciences, University of Sao Paulo, Matao Travessa 14 Cidade Universitaria, Sao Paulo, 05508-090 SP, Brazil, and acLaboratory of Soil Biodiversity, University of Neuchâtel, Rue Emile-Argand 11, Neuchâtel, Switzerland, and adReal Jardín Botánico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain, and aeInstitut de Systématique, Évolution, Biodiversité, Muséum National d’Histoire Naturelle, Sorbonne Universités, 57 rue Cuvier, CP 39 75005, Paris, France, and afDepartment of Integrative Biology, University of Guelph, Summerlee Science Complex, Guelph, Ontario N1G 2W1 Canada, and agDepartment of Zoology, University of British Columbia, 4200-6270 University Blvd., Vancouver, British Columbia V6T 1Z4 Canada, and ahRoyal Botanic Garden, Edinburgh, EH3 5LR, United Kingdom and Institute for Agrifood Research and Technology, C/ Poble Nou km 5.5, Sant Carles de La Ràpita, E-43540 Spain , and aiJardin Botanique de Neuchâtel, Chemin du Perthuis-du-Sault 58, CH-2000 Neuchâtel, Switzerland, and ajDepartment of Natural Sciences, National Museums Northern Ireland, 153 Bangor Road, Holywood BT18 OEU, Northern Ireland, UK, and akDepartment of Oceanography and Kyungpook Institute of Oceanography, School of Earth System Sciences, Kyungpook National University, Daegu, Republic of Korea, and alDepartment of Genetics and Evolution, University of Geneva 1211, Geneva 4, Switzerland, and amDepartment of Biological Sciences, The University of Alabama, Tuscaloosa, AL 35487 USA, and anInstitut de Biologia Evolutiva (CSIC-Univeritat Pompeu Fabra), Passeig Marítim de la Barceloneta 37-49, 08003 Barcelona, Catalonia, Spain, and aoSchool of Applied Sciences, Edinburgh Napier University, Edinburgh, EH11 4BN, United Kingdom, and apDepartment of Biological Sciences, University of Arkansas, Fayetteville, Arkansas, USA, and aqScience Research Centre, Hosei University, 2-17-1 Fujimi, Chiyoda-ku, Tokyo, 102-8160, Japan, and arLaboratoire Evolution et Systématique, Université Paris-XI, 91405 Orsay, France, and asDepartment of Microbiology and Molecular Genetics, Oklahoma State University, Stillwater, OK 74074 USA, and atDepartment of Organismal Biology, Systematic Biology Program, Uppsala university, Uppsala, Sweden, and au Yantai Institute of Coastal Zone Research, Chinese Academy of Science, Yantai 264003, China, and avInstitute of Evolution & Marine Biodiversity, Ocean University of China, Qingdao 266003, China. Abstract: This revision of the classification of eukaryotes follows that of Adl et al., 2012 [J. Euk. Microbiol. 59(5)] and retains an emphasis on protists. Changes since have improved the resolution of many nodes in phylogenetic analyses. For some clades even families are being clearly resolved. As we had predicted, environmental sampling in the intervening years has massively increased the genetic information at hand. Consequently, we have discoveredAuthor Manuscript novel clades, exciting new genera, and uncovered a massive species level diversity beyond the morphological species descriptions. Several clades known from environmental samples only have found their home. Sampling soils, deeper marine waters, and the deep sea will continue to fill us with surprises. The main changes in this revision are the confirmation that eukaryotes form at least two domains, the loss of monophyly in the Exavata, robust support for the Haptista and Cryptista. We provide suggested primer sets for DNA sequences from environmental samples that are effective for each This article is protected by copyright. All rights reserved clade. We have provided a guide to trophic functional guilds in an appendix, to facilitate the interpretation of environmental samples. THIS revision of the classification of eukaryotes updates that of the International Society of Protistologists (Adl et al., 2012). Since then, there has been a massive increase in DNA sequence information of phylogenetic relevance from environmental samples. We now have a much better sense of the undescribed biodiversity in our environment (Pawlowski et al., 2012; de Vargas et al., 2015). While significant, it still remains a partial estimation as several continents and soils in general are poorly sampled, and the deeper ocean is hard to reach. This new data clarified phylogenetic relationships