Bacterial Viruses Subcommittee and Archaeal Viruses Subcommittee of the ICTV : update of taxonomy changes in 2021

This is a self-archived version of an original article. This version may differ from the original in pagination and typographic details. Author(s): Title: Year: Version: Copyright: Rights: Rights url: Please cite the original version: CC BY 4.0 https://creativecommons.org/licenses/by/4.0/ Bacterial Viruses Subcommittee and Archaeal Viruses Subcommittee of the ICTV : update of taxonomy changes in 2021 © The Author(s) 2021 Published version Krupovic, Mart; Turner, Dann; Morozova, Vera; Dyall-Smith, Mike; Oksanen, Hanna M.; Edwards, Rob; Dutilh, Bas E.; Lehman, Susan M.; Reyes, Alejandro; Baquero, Diana P.; Sullivan, Matthew B.; Uchiyama, Jumpei; Nakavuma, Jesca; Barylski, Jakub; Young, Mark J.; Du, Shishen; Alfenas-Zerbini, Poliane; Kushkina, Alla; Kropinski, Andrew M.; Kurtböke, Ipek; Brister, J. Rodney; Lood, Cédric; Sarkar, B. L.; Yigang, Tong; Liu, Ying; Huang, Li; Wittmann, Johannes; Chanishvili, Nina; van Zyl, Leonardo J.; Rumnieks, Janis; Mochizuki, Tomohiro; Jalasvuori, Matti; Aziz, Ramy K.; Łobocka, Małgorzata; Stedman, Kenneth M.; Shkoporov, Andrey N.; Gillis, Annika; Peng, Xu; Enault, François; Knezevic, Petar; Lavigne, Rob; Rhee, Sung-Keun; Cvirkaite-Krupovic, Virginija; Moraru, Cristina; Moreno Switt, Andrea I.; Poranen, Minna M.; Millard, Andrew; Prangishvili, David; Adriaenssens, Evelien M. Krupovic, M., Turner, D., Morozova, V., Dyall-Smith, M., Oksanen, H. M., Edwards, R., Dutilh, B. E., Lehman, S. M., Reyes, A., Baquero, D. P., Sullivan, M. B., Uchiyama, J., Nakavuma, J., Barylski, J., Young, M. J., Du, S., Alfenas-Zerbini, P., Kushkina, A., Kropinski, A. M., . . . Adriaenssens, E. M. (2021). Bacterial Viruses Subcommittee and Archaeal Viruses Subcommittee of the ICTV : update of taxonomy changes in 2021. Archives of Virology, 166(11), 3239-3244. https://doi.org/10.1007/s00705-021-05205-9 2021 Vol.:(0123456789) 1 3 Archives of Virology https://doi.org/10.1007/s00705-021-05205-9 VIROLOGY DIVISION NEWS Bacterial Viruses Subcommittee andArchaeal Viruses Subcommittee oftheICTV: update oftaxonomy changes in2021 MartKrupovic1 · DannTurner2 · VeraMorozova3 · MikeDyall‑Smith4 · HannaM.Oksanen5 · RobEdwards6 · BasE.Dutilh7 · SusanM.Lehman8 · AlejandroReyes9 · DianaP.Baquero1 · MatthewB.Sullivan10,11 · JumpeiUchiyama12 · JescaNakavuma13 · JakubBarylski14· MarkJ.Young15 · ShishenDu16 · PolianeAlfenas‑Zerbini17 · AllaKushkina18 · AndrewM.Kropinski19,20 · IpekKurtböke21 · J.RodneyBrister22· CédricLood23 · B.L.Sarkar24· TongYigang25 · YingLiu1· LiHuang26 · JohannesWittmann27· NinaChanishvili28· LeonardoJ.vanZyl29 · JanisRumnieks30 · TomohiroMochizuki31 · MattiJalasvuori32 · RamyK.Aziz33 · MałgorzataŁobocka34 · KennethM.Stedman35 · AndreyN.Shkoporov36 · AnnikaGillis37 · XuPeng38 · FrançoisEnault39 · PetarKnezevic40 · RobLavigne23 · Sung‑KeunRhee41 · VirginijaCvirkaite‑Krupovic1 · CristinaMoraru42 · AndreaI.MorenoSwitt43 · MinnaM.Poranen5· AndrewMillard44 · DavidPrangishvili45· EvelienM.Adriaenssens46 © The Author(s) 2021 Abstract In this article, we – the Bacterial Viruses Subcommittee and the Archaeal Viruses Subcommittee of the International Committee on Taxonomy of Viruses (ICTV) – summarise the results of our activities for the period March 2020 – March 2021. We report the division of the former Bacterial and Archaeal Viruses Subcommittee in two separate Subcommittees, welcome new members, a new Subcommittee Chair and Vice Chair, and give an overview of the new taxa that were proposed in 2020, approved by the Executive Committee and ratified by vote in 2021. In particular, a new realm, three orders, 15 families, 31 subfamilies, 734 genera and 1845 species were newly created or redefined (moved/promoted). Changes inSubcommittee structure andmembership Bacteriophage and archaeal virus taxonomy has been formally under the auspices of the ICTV Bacterial and Archaeal Viruses Subcommittee, which, at its inception in 1966, was named the Viruses of Prokaryotes Subcommittee, led by David E. Bradley (https:// t alk. ictvo nline. org/ infor mation/ w/ ictvhisto r y). Given the revived interest in bacterial and archaeal viruses and recent enormous increase in the number of characterized isolates and need for creation of numerous taxa to classify them, the Executive Committee (EC) voted on the creation of two separate Subcommittees (EC51, July 2019), formally starting their mandates after EC52 (October 2020). The new Bacterial Viruses Subcommittee is chaired by Evelien Adriaenssens, supported by Dann Turner as the Vice Chair, and the new Archaeal Viruses Subcommittee is chaired by Mart Krupovic. Both Chairs were elected for three-year terms ending in 2023. As a result, this taxonomy update summarises both bacterial and archaeal virus proposals for the last time, reflecting the fact that proposals were submitted prior to the formal reorganisation of the original Subcommittee. In the new Bacterial Viruses Subcommittee, we continue our structure of Study Groups (SGs), regional representatives and general members. We would like to welcome new members Jesca Nakavuma (Uganda), Alejandro Reyes (Colombia), Cristina Moraru (Germany), Susan Lehman (USA), Cédric Lood (Belgium) and Andrey Shkoporov (Ireland) and would like to thank those who have left the Subcommittee for their service. In the framework of the Bacterial and Archaeal Viruses Subcommittee, all taxonomic proposals dealing with archaeal viruses were handled by a single SG. The new Handling Editor: Sead Sabanadzovic. * Evelien M. Adriaenssens ev[email protected] Extended author information available on the last page of the article M. Krupovic etal. 1 3 Archaeal Viruses Subcommittee currently includes 11 SGs (Table1), each created for a three-year term. Of note, the Halopanivirales SG, chaired by Hanna M. Oksanen (Finland), oversees the taxonomy of evolutionarily related viruses of the order Halopanivirales, which includes two families of viruses infecting archaea (Sphaerolipoviridae and Simuloviridae) and one family of bacteriophages (Matsushitaviridae). Thus, the latter SG bridges the two Subcommittees. Additional SGs will be created in the near future, in particular, to develop the taxonomy of archaeal members of the class Caudoviricetes; only a handful of isolated representatives of this highly diverse and expansive group of archaeal viruses are currently classified. Taxonomy update We had a record year of submissions of taxonomy proposals in 2020, with 188 proposals submitted, of which all but one were approved and ratified (Supplementary TableS1). The changes at each of the taxonomic ranks in use for bacterial and archaeal viruses are summarised in Table2. We Table 1 Composition of the archaeal viruses subcommittee *Study Group Chair Study Group Member Country Bicaudaviridae SG Mart Krupovic* France Li Huang China Mark J. Young USA Virginija Cvirkaite-Krupovic France Desulfurococcales viruses SG Tomohiro Mochizuki* Japan Mart Krupovic France Fuselloviridae SG Kenneth M. Stedman* USA Mart Krupovic France Halopanivirales SG Hanna M. Oksanen* Finland Mike Dyall-Smith Australia Shishen Du China Matti Jalasvuori Finland Halspiviridae SG Mike Dyall-Smith* Australia Hanna M. Oksanen Finland Ovaliviridae SG Li Huang* China Pleolipoviridae SG Hanna M. Oksanen* Finland Mike Dyall-Smith Australia Ying Liu France Portogloboviridae SG Mart Krupovic* France Ying Liu France Thaspiviridae SG Sung-Keun Rhee* Republic of Korea Mart Krupovic France Tokiviricetes SG Mart Krupovic* France Tomohiro Mochizuki Japan Xu Peng Denmark Diana P. Baquero France Turriviridae SG Mark J. Young* USA Table 2 Summary of taxonomic changes for bacterial and archaeal viruses for Master Species List 36, ratified March 2021 Species Genus Subfamily Family Order Class Phylum Kingdom Realm Abolished 20 2 0 0 0 0 0 0 0 New 1775 700 28 14 2 1 1 1 1 Moved or promoted (and/or renamed) 70 34 3 1 1 0 0 0 0 Renamed 33 8 0 1 1 1 0 0 0 Taxonomy of bacterial and archaeal viruses 1 3 created a record 1775 new species, 700 new genera, 28 new subfamilies, 14 new families and two new orders and also created one new realm containing one new kingdom, one new phylum and one new class. Given the large numbers of proposals and taxa that were created, moved or renamed, it becomes unfeasible to describe all the changes in detail; however, we urge interested readers to consult Supplementary TableS1 and the associated proposals from the ICTV website (https:// talk. ictvo nline. org/ files/ ictv_ offic ial_ taxon omy_ updat es_ since_ the_ 8th_ report/ m/ proka ryoteoffic ial). Instead, below we provide a brief overview of the most notable changes. The new realm Adnaviria, anew megataxonomy ofarchaeal filamentous viruses Recently, the taxonomic framework of the ICTV has been expanded from five to 15 ranks, with the highest-level rank, realm, being equivalent to the domain rank used for cellular organisms [1]. Until recently, four such realms had been established for classification of viruses infecting hosts from different domains of life [2, 3]. This year, a new realm, Adnaviria, was created for classification of archaeal filamentous viruses with dsDNA genomes that adopt the A-form conformation within their virions [4, 5]. The realm includes three families, Tristromaviridae, Rudiviridae and Lipothrixviridae, which are evolutionarily related to each other, but not to any other known group of viruses. The families Rudiviridae and Lipothrixviridae were already grouped together in the order Ligamenvirales [6], and a single-family order, Primavirales, has now been created for the family Tristromaviridae. The two orders are unified in the class Tokiviricetes, kingdom Zilligvirae and realm Adnaviria. Class Caudoviricetes, order Caudovirales We delineated three new families of short-tailed phages with small genomes that were formerly assigned to the family Podoviridae. The family Salasmaviridae, named in honour of Margarita Salas Falgueras, comprises the existing subfamily Picovirinae, which includes the classical bacillus phage φ29 and a range of related Bacillus-infecting phages with genomes between 18 and 27 kb in size. The family Rountreeviridae, named in honour of Phyllis Margaret Rountree, groups Enterococcus-infecting phages with genomes between 17 and 19 kb in size, whereas the family Guelinviridae, named after Antonina Guelin, groups Clostridiuminfecting phages with genome sizes between 16 and 19 kb. The new family Schitoviridae, named after Giancarlo Schito, is the formalisation of the group of N4-like phages defined by the presence of a large virion-associated RNA polymerase, described in more detail in reference [7]. The new family Zobellviridae, named after Claude Zobell, groups a set of globally distributed podoviruses associated with marine ecosystems first proposed by Bischoff and colleagues [8]. Class Leviviricetes Based on the investigation by Callanan and colleagues on the expansion of known ssRNA virus genomes [9], the family Leviviridae was elevated to the rank of class, named Leviviricetes (replacing the class Allassoviricetes), and expanded to include two orders (Norzivirales and Timlovirales) and six new families: Fiersviridae (renamed from the original family Leviviridae), Atkinsviridae, Duinviridae, Solspiviridae, Blumeviridae and Steitzviridae. A detailed description of the new taxa will be published separately. Class Tectiliviricetes, new family Autolykiviridae The new family Autolykiviridae formalises the group of nontailed dsDNA bacteriophages discovered and described by Kauffmann and colleagues [10], combining features of both corticoviruses and tectiviruses. This new family contains two new genera and five new species. Order Halopanivirales, new families Matsushitaviridae andSimuloviridae Until recently, the family Sphaerolipoviridae, which includes icosahedral tailless viruses with internal membranes, consisted of three genera, Alphasphaerolipovirus, Betasphaerolipovirus and Gammasphaerolipovirus. The first two of these genera included viruses infecting halophilic archaea, whereas the last one included phages infecting thermophilic bacteria [11]. Although viruses from the three genera are evolutionarily related [12], they display considerable sequence divergence. Thus, the genera Betasphaerolipovirus and Gammasphaerolipovirus have been renamed and moved from the Sphaerolipoviridae into new families, Simuloviridae and Matsushitaviridae (named after Isao Matsushita), respectively. The order Halopanivirales now contains a family of bacterial viruses, Matsushitaviridae, and two families of archaeal viruses, family Sphaerolipoviridae and Simuloviridae. As mentioned above, the order is under purview of a single Study Group, which is part of both the Archaeal Viruses SC and the Bacterial Viruses SC. M. Krupovic etal. 1 3 Order Tubulavirales, new family Paulinoviridae The new family Paulinoviridae addresses the challenge of defining family demarcation criteria for phages with small genomes, where members of the same family would share at least two orthologous proteins. Informed by prior work on a gene-content network of predicted filamentous prophage sequences [13], we moved the genera Bifilivirus and Thomixvirus from the family Inoviridae to the family Paulinoviridae. Online (10th) Report oftheICTV Virus Taxonomy: The Classification and Nomenclature of Viruses - The Online (10th) Report of the ICTV is freely accessible at http:// ictv. global/ report, and summaries of the chapters on each virus family are published in the Journal of General Virology. In 2020, four new chapters on bacterial and archaeal viruses were produced by members of the Archaeal Viruses SC and Bacterial Viruses SC, namely, on the families Herelleviridae [14], Spiraviridae [15], Ovaliviridae [16] and Finnlakeviridae [17]. Conclusion This past year has been extremely productive in terms of new bacterial and archaeal virus taxa described. It would not have been possible without an active pool of scientists, both within and outside the subcommittee and its study groups, who scour databases, perform analyses and submit proposals. We continue to encourage people to contact us to formalise new discoveries into the taxonomic framework and will keep reaching out to interested parties. Finally, we would like to acknowledge one person in particular, Prof Andrew Kropinski, the Subcommittee Chair from 2014-2020, who authored, co-authored and/or assisted with the majority of proposals that have been approved in the last decade. Supplementary Information The online version contains supplementary material available at https:// doi. or g/ 10. 1007/ s0070502105205-9. Acknowledgements We would like to acknowledge the invaluable help from colleagues at NCBI, in particular Igor Tolstoy. Author contributions All authors contributed to the creation or assessment of taxonomy proposals; M.K. and E.M.A wrote the manuscript with input from D.T. and A.M.K; all authors evaluated and approved the final version. Funding V.M. was supported by Russian Ministry of Education and Science Project No. 0245-2021-0008; H.M.O. was supported by University of Helsinki funding for FINStruct and Instruct-ERIC research infrastructure; R.A.E. was supported by the National Institute Of Diabetes And Digestive And Kidney Diseases of the National Institutes of Health under Award Number RC2DK116713; B.E.D. was supported by the European Research Council (ERC) Consolidator grant 865694: DiversiPHI; M.B.S. was supported by National Science Foundation Advances in Biological Infrastructure award #1758974; M.Y. acknowledges funding by the Gordon and Betty Moore Foundation grant GBMF9195; S.D. was supported by National Natural Science Foundation of of China grant 32070032; C.L. is supported by the Fonds Wetenschappelijk Onderzoek (1S64718N); L.H. was funded by the Strategic Priority Research Program of the Chinese Academy of Sciences Award XDB42000000,National Natural Science Foundation of China (NSFC) grant 31970170; T.M. was supported by JSPS KAKENHI Grant Numbers 21H02100, 19H04827, 18K14372, and JST JPMJJR2005, and SUMITOMO Fund 200673; MJ is supported by Academy of Finland grants #336518 and #297049 and Jane and Aatos Erkko Foundation; R.K.A. is supported by the Egyptian Academy of Scientific Research and Technology (ASRT) project #3046 (JESOR); M.L. is supported by the statutory funds for the Institute of Biochemistry and Biophysics of the Polish Academy of Sciences; K.M.S. is supported by the US. National Science Foundation (MCB1929273 and MCB2025305); A.N.S. is supported by Wellcome Trust Research Career Development Fellowship (220646/Z/20/Z) and by European Research Council (ERC) grant under the European Union’s Horizon 2020 research and innovation programme (grant agreement 101001684); X.P. is supported by Novo Nordisk Foundation/Hallas-Møller Ascending Investigator Grant (grant number NNF17OC0031154) and the Danish Council for Independent Research/ FNU (grant number DFF–0135-00402); S.K.R. was supported by the National Research Foundation of Korea (NRF) grants funded by the Ministry of Science, ICT & Future Planning (2021R1A2C3004015); C.M. acknowledges funding by the Deutsche Forschungsgemeinschaft within the Collaborative Research Center TRR51 Roseobacter (INST 184/170-1); M.M.P. acknowledges funding from the Academy of Finland (grant 331627) and the Sigrid Juselius Foundation; E.M.A. gratefully acknowledges funding by the Biotechnology and Biological Sciences Research Council (BBSRC); this research was funded by the BBSRC Institute Strategic Programme Gut Microbes and Health BB/R012490/1 and its constituent projects BBS/E/F/000PR10353 and BBS/E/F/000PR10356. Declarations Conflict of interest All authors are current or former members of the Bacterial and Archaeal Viruses Subcommittee, Bacterial Viruses Subcommittee and/or Archaeal Viruses Subcommittee of the International Committee on Taxonomy of Viruses (ICTV). Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http:// creat iveco mmons. org/ licen ses/ b y/4. 0/. Taxonomy of bacterial and archaeal viruses 1 3 References 1. Gorbalenya AE, Krupovic M, Mushegian A etal (2020) The new scope of virus taxonomy: partitioning the virosphere into 15 hierarchical ranks. Nat Microbiol 5:668–674. https:// doi. org/ 10. 1038/ s415640200709-x 2. Koonin EV, Dolja VV, Krupovic M etal (2020) Global organization and proposed megataxonomy of the virus world. Microbiol Mol Biol Rev 84:1–33. https:// doi. org/ 10. 1128/ MMBR. 0006119 3. Walker PJ, Siddell SG, Lefkowitz EJ etal (2020) Changes to virus taxonomy and the Statutes ratified by the International Committee on Taxonomy of Viruses (2020). 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Authors and Affiliations MartKrupovic1 · DannTurner2 · VeraMorozova3 · MikeDyall‑Smith4 · HannaM.Oksanen5 · RobEdwards6 · BasE.Dutilh7 · SusanM.Lehman8 · AlejandroReyes9 · DianaP.Baquero1 · MatthewB.Sullivan10,11 · JumpeiUchiyama12 · JescaNakavuma13 · JakubBarylski14· MarkJ.Young15 · ShishenDu16 · PolianeAlfenas‑Zerbini17 · AllaKushkina18 · AndrewM.Kropinski19,20 · IpekKurtböke21 · J.RodneyBrister22· CédricLood23 · B.L.Sarkar24· TongYigang25 · YingLiu1· LiHuang26 · JohannesWittmann27· NinaChanishvili28· LeonardoJ.vanZyl29 · JanisRumnieks30 · TomohiroMochizuki31 · MattiJalasvuori32 · RamyK.Aziz33 · MałgorzataŁobocka34 · KennethM.Stedman35 · AndreyN.Shkoporov36 · AnnikaGillis37 · XuPeng38 · FrançoisEnault39 · PetarKnezevic40 · RobLavigne23 · Sung‑KeunRhee41 · VirginijaCvirkaite‑Krupovic1 · CristinaMoraru42 · AndreaI.MorenoSwitt43 · MinnaM.Poranen5· AndrewMillard44 · DavidPrangishvili45· EvelienM.Adriaenssens46 1 Archaeal Virology Unit, Institut Pasteur, Paris, France 2 Department ofApplied Sciences, Faculty ofHealth andApplied Sciences, University oftheWest ofEngland, Bristol, UK 3 Laboratory ofMolecular Microbiology, Institute ofChemical Biology andFundamental Medicine SB RAS, Novosibirsk, Russia 4 Veterinary Biosciences, Faculty ofVeterinary andAgricultural Sciences, University ofMelbourne, Parkville, Australia 5 Molecular andIntegrative Biosciences Research Programme, Faculty ofBiological andEnvironmental Sciences, University ofHelsinki, Helsinki, Finland 6 Flinders Accelerator forMicrobiome Exploration, Adelaide, Australia 7 Theoretical Biology andBioinformatics, Department ofBiology, Utrecht University, Utrecht, Netherlands 8 Center forBiologics Evaluation andResearch, US Food andDrug Administration, 10903 New Hampshire Ave, SilverSpring, MD20903, USA M. Krupovic etal. 1 3 9 Max Planck Tandem Group inComputational Biology, Department ofBiological Sciences, Universidad de Los Andes, Bogotá, Colombia 10 Department ofMicrobiology, The Ohio State University, Columbus, OH43210, USA 11 Department Civil, Environmental, andGeodetic Engineering, The Ohio State University, Columbus, OH43210, USA 12 School ofVeterinary Medicine, Azabu University, Sagamihara, Japan 13 College ofVeterinary Medicine, Animal Resources andBiosecurity, Makerere University, Kampala, Uganda 14 Department ofMolecular Virology, Faculty ofBiology, Institute ofExperimental Biology, Adam Mickiewicz University inPoznan, Collegium Biologicum-Umultowska 89, 61-614Poznan, Poland 15 Department ofPlant Sciences andPlant Pathology, Montana State University, Bozeman, MT, USA 16 Department ofMicrobiology, College ofLife Sciences, Wuhan University, Wuhan, China 17 Department ofMicrobiology, Universidade Federal de Viçosa, Viçosa, Brazil 18 Zabolotny Institute ofMicrobiology andVirology, NAS ofUkraine, Kyiv, Ukraine 19 Department ofFood Science, University ofGuelph, Guelph, Canada 20 Department ofPathobiology, University ofGuelph, Guelph, Canada 21 University oftheSunshine Coast, SippyDowns, Australia 22 National Center forBiotechnology Information, National Library ofMedicine, National Institutes ofHealth, Bethesda, USA 23 KU Leuven, Leuven, Belgium 24 Emeritus ICMR-National Institute ofCholera andEnteric Diseases (NICED), Kolkata, India 25 Beijing University ofChemical Technology, Beijing, China 26 State Key Laboratory ofMicrobial Resources, Institute ofMicrobiology, Chinese Academy ofSciences, Beijing, China 27 Leibniz Institute DSMZ-German Collection ofMicroorganisms andCell Cultures GmbH, Berlin, Germany 28 The Eliava Institute ofBacteriophage, MIcrobiology andVirology, Tbilisi, Georgia 29 University oftheWestern Cape, WesternCape, SouthAfrica 30 Latvian Biomedical Research andStudy Center, Riga, Latvia 31 Earth-Life Science Institute, Tokyo Institute ofTechnology, Tokyo, Japan 32 University ofJyvaskyla, Jyvaskyla, Finland 33 Department ofMicrobiology andImmunology, Faculty ofPharmacy, Cairo University andMicrobiology andImmunology Research Program, Children’s Cancer Hospital Egypt, Cairo57357, Egypt 34 Laboratory ofBacteriophage Biology, Institute ofBiochemistry andBiophysics ofthePolish Academy ofSciences, Warsaw, Poland 35 Biology Department andCenter forLife inExtreme Environments, Portland State University, Portland, USA 36 University College Cork, Cork, Ireland 37 Laboratory ofFood andEnvironmental Microbiology, Earth andLife Institute, UC Louvain, Louvain-la-Neuve, Belgium 38 Microbial Immunity Group, Department ofBiology, University ofCopenhagen, Copenhagen, Denmark 39 Université Clermont Auvergne, CNRS, LMGE, 63000Clermont-Ferrand, France 40 Faculty ofSciences Department ofBiology andEcology, University ofNovi Sad, NoviSad, Serbia 41 Department ofBiological Sciences andBiotechnology, Chungbuk National University, Cheongju, SouthKorea 42 Institute forChemistry andBiology oftheMarine Environment, University ofOldenburg, Oldenburg, Germany 43 Escuela de Medicina Veterinaria, Pontificia Universidad Católica de Chile, Santiago, Chile 44 Department ofGenetics andGenome Biology, University ofLeicester, Leicester, UK 45 Ivane Javakhishvili Tbilisi State University, Tbilisi, Georgia 46 Quadram Institute Biosciences, Norwich Research Park, NorwichNR47UQ, UK