Consensus statement: Virus taxonomy in the age of metagenomics

Journal name:
Nature Reviews Microbiology
Volume:
15,
Pages:
161–168
Year published:
DOI:
doi:10.1038/nrmicro.2016.177
Published online

Abstract

The number and diversity of viral sequences that are identified in metagenomic data far exceeds that of experimentally characterized virus isolates. In a recent workshop, a panel of experts discussed the proposal that, with appropriate quality control, viruses that are known only from metagenomic data can, and should be, incorporated into the official classification scheme of the International Committee on Taxonomy of Viruses (ICTV). Although a taxonomy that is based on metagenomic sequence data alone represents a substantial departure from the traditional reliance on phenotypic properties, the development of a robust framework for sequence-based virus taxonomy is indispensable for the comprehensive characterization of the global virome. In this Consensus Statement article, we consider the rationale for why metagenomic sequence data should, and how it can, be incorporated into the ICTV taxonomy, and present proposals that have been endorsed by the Executive Committee of the ICTV.

At a glance

Figures

  1. Prevalence, abundance and affiliation of marine viruses.
    Figure 1: Prevalence, abundance and affiliation of marine viruses.

    The 15,222 virus populations that were identified across the Global Ocean Viromes (GOV) dataset69 are shown according to their prevalence (x-axis, number of sampling stations in which the population was detected), average abundance (y-axis, log10 scale, average of normalized coverage across all samples in which the population was detected), and are coloured by the taxonomic affiliation of their host (affiliation is based on best basic local alignment search tool (BLAST) hit of predicted genes; a population was associated to a virus isolate and its host when ≥50% of predicted genes were affiliated to this virus isolate; 512 of the 15,222 populations could be affiliated). Figure courtesy of S. Roux and M.B.S., The Ohio State University, USA.

  2. Genetic diversity of CRESS-DNA viruses.
    Figure 2: Genetic diversity of CRESS-DNA viruses.

    The replication-associated protein (Rep) sequences of 659 circular Rep-encoding single-stranded DNA (ssDNA) viruses (CRESS-DNA viruses) were compared with 10 representative Repsequences from viruses classified in the families Geminiviridae, Nanoviridae, Circoviridae and Genomoviridae, and a group of alpha satellites that are associated with geminiviruses or nanoviruses. Amino acid sequences were aligned using Multiple Alignment using Fast Fourier Transform (MAFFT; G-INS-i option)70, and a maximum likelihood phylogenetic tree was constructed using Fasttree71. Branches with less than 50% SH (Shimodaira–Hasegawa)-like support were collapsed.

  3. Summary of the proposed classification pipeline.
    Figure 3: Summary of the proposed classification pipeline.

    The proposed classification pipeline (red arrows) enables both metagenomic sequence data and conventionally derived virus sequences to be classified. Inferred biological properties that are obtained by bioinformatic analysis of virus sequences together with information on sequence relatedness and gene content, and, optionally, any observed biological properties (dotted line), may all be used as defining criteria for species and higher rank taxonomic assignment in the International Committee on Taxonomy of Viruses (ICTV) taxonomy. This procedure differs from current (green arrows) and previous practice (blue arrows), in which biological data and/or host information and sequence data (current), or biological data alone (1970s–1990s), were required for classification.

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Author information

Affiliations

  1. Nuffield Department of Medicine, University of Oxford, South Parks Road, Oxford OX1 3SY, UK.

    • Peter Simmonds
  2. 24 Woodland Way, Stevenage, Hertfordshire SG2 8BT, UK.

    • Mike J. Adams
  3. Institute for Veterinary Medical Research, Centre for Agricultural Research, Hungarian Academy of Sciences, 21 Hungária krt., Budapest H-1143, Hungary.

    • Mária Benkő &
    • Balázs Harrach
  4. University of South Florida, College of Marine Science, 140 7th Avenue South, Saint Petersburg, Florida 33701, USA.

    • Mya Breitbart
  5. National Center for Biotechnology Information, National Library of Medicine, National Institutes of Health, Bethesda, Maryland 20894, USA.

    • J. Rodney Brister &
    • Eugene V. Koonin
  6. Department of Biomedical and Molecular Sciences, Queen's University, 18 Stuart Street, Kingston, Ontario K7L 3N6, Canada.

    • Eric B. Carstens
  7. UK Medical Research Council (MRC)–University of Glasgow Centre for Virus Research, Sir Michael Stoker Building, 464 Bearsden Road, Glasgow G61 1QH, UK.

    • Andrew J. Davison &
    • Richard Orton
  8. Blood Systems Research Institute, 270 Masonic Avenue, San Francisco, California 94118, USA.

    • Eric Delwart
  9. Department of Laboratory Medicine, 521 Parnassus Avenue, University of California, San Francisco, California 94118, USA.

    • Eric Delwart
  10. Department of Medical Microbiology, Leiden University Medical Center, PO Box 9600, E4-P, room E4-72, 2300 RC Leiden, The Netherlands.

    • Alexander E. Gorbalenya
  11. Faculty of Bioengineering and Bioinformatics, Lomonosov Moscow State University, 119899 Moscow, Russia.

    • Alexander E. Gorbalenya
  12. 3 Portman Drive, Child Okeford, Blandford Forum, Dorset DT11 8HU, UK.

    • Roger Hull
  13. The Pirbright Institute, Ash Road, Pirbright, Woking, Surrey GU24 0NF, UK.

    • Andrew M.Q. King
  14. Department of Microbiology, Institut Pasteur, 25 Rue du Dr Roux, 75015 Paris, France.

    • Mart Krupovic
  15. National Institute of Allergy and Infectious Diseases (NIAID), Integrated Research Facility at Fort Detrick (IRF–Frederick), B-8200 Research Plaza, Fort Detrick, Frederick, Maryland 21702, USA.

    • Jens H. Kuhn
  16. Department of Microbiology, University of Alabama at Birmingham (UAB), Bevill Biomedical Research Building (BBRB) Suite 276, 845 19th Street South, Birmingham, Alabama 35294–2170, USA.

    • Elliot J. Lefkowitz
  17. Department of Microbiology and Immunobiology, Harvard Medical School, 77 Avenue Louis Pasteur, Boston, Massachusetts 02115, USA.

    • Max L. Nibert
  18. Department of Plant Pathology and Environmental Microbiology, Center for Infectious Disease Dynamics, Pennsylvania State University, University Park, Pennsylvania 16802, USA.

    • Marilyn J. Roossinck
  19. Department of Biochemistry, Molecular Biology, Entomology and Plant Pathology, Mississippi State University, 100 Old Highway 12 Mail Stop 9775, Mississippi State, Mississippi 39762, USA.

    • Sead Sabanadzovic
  20. Departments of Microbiology and Civil, Environmental and Geodetic Engineering, Ohio State University, Columbus, Ohio 43210, USA.

    • Matthew B. Sullivan
  21. Departments of Earth, Ocean and Atmospheric Sciences, Microbiology and Immunology, and Botany, University of British Columbia, Vancouver, V6T 1Z4, Canada.

    • Curtis A. Suttle
  22. Canadian Institute for Advanced Research (CIFAR), 180 Dundas Street West, Toronto, Ontario M5G 1Z8, Canada.

    • Curtis A. Suttle
  23. Department of Pathology and Center for Biodefense and Emerging Infectious Diseases, University of Texas Medical Branch, 301 University Boulevard, Galveston, Texas 77555–0609, USA.

    • Robert B. Tesh
  24. Wageningen Plant Research, Wageningen University and Research Centre (WUR–PRI), Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands.

    • René A. van der Vlugt
  25. The Center for Fundamental and Applied Microbiomics, The Biodesign Institute and School of Life Sciences, Arizona State University, 1001 South McAllister Avenue, Tempe, Arizona 85281–2115, USA.

    • Arvind Varsani
  26. Departamento de Fitopatologia/BIOAGRO, Universidade Federal de Viçosa, Viçosa, Minas Gerais 36570–900, Brazil.

    • F. Murilo Zerbini
  27. Retired from The John Innes Centre, Norwich, Norfolk, UK.

    • Roger Hull

Competing interests statement

The authors declare no competing interests.

Corresponding author

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Author details

  • Peter Simmonds

    Peter Simmonds studies the effects and disease mechanisms of human and veterinary viruses. Following his graduation in medicine, he pursued postgraduate medical training (MRCPath 1995) and became Professor of Virology at the University of Edinburgh, UK, in 2001. He has pursued a research programme that investigates the mechanisms of viral pathogenesis, disease associations of emerging and newly discovered viruses, and, more recently, investigations of viral and human genetic determinants of viral persistence and immune responses. He has been an elected member of the International Committee on Taxonomy of Viruses (ICTV) Executive Committee since 2011. He is currently Professor of Virology in the Nuffield Department of Medicine at the University of Oxford, UK.

  • Mike J. Adams

    Mike J. Adams holds a Ph.D. in plant pathology from the University of London, UK. He has held research posts at Rothamsted Research, Harpenden, UK, where he was most recently Principal Investigator involved in the characterization, epidemiology and control of viruses that are transmitted by plasmodiophorid vectors. He has now retired but retains an interest in the taxonomy of viruses, especially those that infect plants. He has been a member of the International Committee on Taxonomy of Viruses (ICTV) Executive Committee since 2005.

  • Mária Benkő

    Mária Benkő is a private professor at the University of Veterinary Medicine, Budapest, Hungary. Her research activity is focused on different DNA viruses of veterinary importance. She has described novel adenoviruses, herpesviruses, parvoviruses and circoviruses in different domesticated and wild animals, including lower vertebrates, namely fish, amphibians and reptiles. She is Chair of the Adenoviridae Study Group of International Committee on Taxonomy of Viruses (ICTV). She also chairs the Advisory Committee on Gene Technology Procedures for the Department of Agriculture of the Hungarian Government.

  • Mya Breitbart

    Mya Breitbart is a professor in the College of Marine Science at the University of South Florida, St. Petersburg, USA. She earned her B.Sc. in biology from Florida Institute of Technology, Melbourne, USA, in 2000, and her Ph.D. from the joint doctoral programme in cellular and molecular biology at San Diego State University, California, USA, and the University of California, San Diego, USA, in 2006. Her research uses metagenomic sequencing to explore the identity, diversity and distribution of viruses in a wide range of hosts and environments.

  • J. Rodney Brister

    J. Rodney Brister is a staff scientist and Viral Genomes Unit Chief at the National Center for Biotechnology Information (NCBI), National Library of Medicine, National Institutes of Health, Bethesda, Maryland, USA. He received his M.Sc. in biology from Emory University, Atlanta, Georgia, USA, and his Ph.D. in Molecular Genetics and Microbiology from the University of Florida, Gainesville, USA. His research interests include viral genome structure, the evolution of viruses and the development of computational frameworks for the storage, analysis and retrieval of viral sequence data.

  • Eric B. Carstens

    Eric B. Carstens is a professor in the Department of Biomedical and Molecular Sciences at Queen's University, Kingston, Ontario, Canada. He received his M.Sc. from the University of Alberta, Edmonton, Canada, where he carried out electron microscopy studies of adenovirus structural proteins, and his Ph.D. in adenovirus molecular genetics from the University of Sherbrooke, Quebec, Canada. He was a postdoctoral fellow at the Institute of Genetics at the University of Cologne, Germany, where he studied adenovirus and baculovirus replication. His research has focused on the molecular analysis of baculovirus genes that are involved in the regulation of virus DNA replication and transcription. He has served several roles on the Executive Committee of the International Committee on Taxonomy of Viruses (ICTV), including Subcommittee Chair of Invertebrate Viruses, Vice President, and President and Treasurer.

  • Andrew J. Davison

    Andrew J. Davison studies herpesvirus genomics. He graduated with a B.A. in biochemistry, in 1976, from the University of Cambridge, UK, and obtained his Ph.D. in virology, in 1981, from the University of Glasgow, UK, where he has worked since, apart from two years spent at the National Institutes of Health, Bethesda, Maryland, USA. He leads research groups at the UK Medical Research Council (MRC)–University of Glasgow Centre for Virus Research (CVR), UK, on human cytomegalovirus and viral genomics and bioinformatics. He is the current President of the International Committee on Taxonomy of Viruses (ICTV).

  • Eric Delwart

    Eric Delwart is an investigator at the Blood Systems Research Institute, San Francisco, California, USA, and Professor in the Department of Laboratory Medicine at the University of California, San Francisco (UCSF), USA. He received his Ph.D. from the University of Wisconsin–Madison, USA, where he studied cell entry of an avian retrovirus and HIV. During his postdoctoral research at Stanford University, California, USA, he developed a simple DNA heteroduplex mobility assay to measure genetic relationships between re-annealed DNA strands and HIV evolution. At the Aaron Diamond AIDS Research Center, New York, USA, he studied drug resistance in HIV and T cell receptor re-arrangements. He is now investigating transfusion-transmitted viruses (HIV, hepatitis B virus (HBV), hepatitis C virus (HCV), West Nile virus (WNV) and Zika virus (ZIKV)), and uses viral metagenomics to identify novel pathogens and to characterize the viromes of different human and animal populations.

  • Alexander E. Gorbalenya

    Alexander Gorbalenya received his M.Sc. in genetics from Novosibirsk State University, Russia. During the final years of his Ph.D. training in virus biochemistry at the Institute of Poliomyelitis, Moscow, Russia, and the Institute of Molecular Biology, Koltsovo, Russia, he started using nascent comparative sequence analysis to study the structure, function and evolution of viruses, which became his calling in virology. He was awarded a D.Sci. in molecular biology from Lomonosov Moscow State University, Russia, in 1991, and was Lead Research Scientist at the Institute of Poliomyelitis, in 1998, before moving to the US National Cancer Institute, Frederick, Maryland, USA, as a visiting scientist, and, since 2001, to Leiden University Medical Center, The Netherlands. Currently he is the Leiden University Fund Professor of Applied Bioinformatics in Virology and Professor at the Faculty of Bioengineering and Bioinformatics, Lomonosov Moscow State University, Russia. In 2005, he became a member of the Executive Committee of the International Committee on Taxonomy of Viruses (ICTV), and has been Vice President since 2011.

  • Balázs Harrach

    Balázs Harrach is a corresponding member of the Hungarian Academy of Sciences, an honorary professor at the University of Veterinary Medicine, Budapest, Hungary. His main interest is the study of the diversity and evolution of animal adenoviruses. Based on the discovery and complete genomic characterization of numerous novel adenoviruses, the taxonomy of the family Adenoviridae has been radically altered so that, currently, it contains five established and one proposed genera. He is Chair of the Animal DNA Viruses and Retroviruses Subcommittee of the International Committee on Taxonomy of Viruses (ICTV), the Editor-in-Chief of Acta Veterinaria Hungarica, and the Vice President of the Hungarian Society for Bioinformatics.

  • Roger Hull

    Roger Hull has studied plant viruses for more than 40 years. He obtained his Ph.D. at the University of London, UK, and became a research scientist at the Agricultural Research Council Virus Research Unit, Cambridge, UK, and then the John Innes Institute (now known as the John Innes Centre), Norfolk, UK (retired). He studied a wide range of viruses, focusing mainly on non-conventional approaches to resistance, diagnostics, epidemiology and evolution. He has published more than 250 peer-reviewed papers and several books, including the last two editions of Plant Virology. His awards include a D.Sc. (the University of London), several Professorships, Fellowship of the American Phytopathological Society, life membership of the American Society of Virology, the British Society of Plant Pathology and the International Committee on Taxonomy of Viruses (ICTV). Since his retirement he has maintained his interests in virology and has written books and served on various teaching and research initiatives.

  • Andrew M.Q. King

    Andrew M. Q. King worked, for many years, at Pirbright Laboratory, Institute for Animal Health (now known as the Pirbright Institute), Surrey, UK, before retiring in 2004. His main interests were in understanding the molecular and structural interactions between foot-and-mouth disease virus and antibodies and cellular receptors. His involvement with the International Committee on Taxonomy of Viruses (ICTV) pre-dates 1993, when he took over as Chair of the Picornaviridae Study Group. In 2001, he joined the Executive Committee, to represent the eight study groups that covered positive-strand RNA viruses of vertebrates. Since 2007, he has served as Proposals Secretary, piloting taxonomic proposals through the approval process of the ICTV and publicising proposals online. More recently, he was head of the editorial team that produced the Ninth Report of the ICTV.

  • Eugene V. Koonin

    Eugene V. Koonin is the leader of the Evolutionary Genomics Group in the National Center for Biotechnology Information (NCBI) at the National institutes of Health (NIH), Bethesda, Maryland, USA. He is known primarily for his research on genome evolution, especially in microorganisms and viruses, host–parasite co-evolution, and, more specifically, functions and the evolution of CRISPR–Cas systems. He received his Ph.D. in molecular biology, in 1983, from the Department of Biology, Moscow State University, Russia. He was a research scientist at the Institute of Poliomyelitis, and subsequently, a senior research scientist and Laboratory Chief at the Institute of Microbiology of the Russian Academy of Sciences. He joined the NCBI, in 1991, as a visiting scientist and became a senior investigator in 1996. He is the author of 'The Logic of Chance: The Nature and Origin of Biological Evolution' and is the founder and Editor-in-Chief of Biology Direct, an open access, open peer-review journal.

  • Mart Krupovic

    Mart Krupovic is an assistant professor in the Department of Microbiology at the Institut Pasteur, Paris, France. He received his Ph.D. in general microbiology, in 2010, from the University of Helsinki, Finland. He has a profound interest in the origin and evolution of the global virosphere, and the evolutionary connections between viruses and other types of mobile genetic elements, including plasmids and transposons. He also develops an experimental line of research on different aspects of virus–host interactions in hyperthermophilic archaea. He is a member of the Executive Committee of the International Committee on Taxonomy of Viruses (ICTV).

  • Jens H. Kuhn

    Jens H. Kuhn is the Virology Lead (contractor) at the National Institute of Allergy and Infectious Diseases (NIAID) Integrated Research Facility at Fort Detrick, Frederick, Maryland, USA. He specializes in the development and evaluation of medical countermeasures against highly virulent viral human and animal pathogens that require biosafety level 4 containment, emerging virus discovery and characterization, and RNA virus diversity. He is a member of the Executive Committee of the International Committee on Taxonomy of Viruses (ICTV).

  • Elliot J. Lefkowitz

    Elliot J. Lefkowitz is Professor of Microbiology at the University of Alabama at Birmingham, USA. His research interests are directed at contributing to the understanding of microbial genomics and evolution by developing and using computational tools and bioinformatics techniques to mine sequence and other data for significant patterns that are characteristic of function and/or evolution. In particular, he has investigated the evolution of poxviruses, tracing the effect of molecular events, such as horizontal gene transfer between host and virus, on virus evolution. He serves as Data Secretary for the International Committee on Taxonomy of Viruses (ICTV).

  • Max L. Nibert

    Max L. Nibert is Professor of Microbiology and Immunobiology at Harvard Medical School, Boston, Massachusetts, USA. His research efforts over many years have concerned various different double-stranded RNA (dsRNA) viruses. He currently serves as Chair of the Fungal and Protist Viruses Subcommittee of the International Committee on Taxonomy of Viruses (ICTV).

  • Richard Orton

    Richard Orton is a postdoctoral research associate at the UK Medical Research Council (MRC)–University of Glasgow Centre for Virus Research (CVR), UK. He received his Ph.D. in bioinformatics from the University of Edinburgh, UK, in 2006. His research focuses on the development of bioinformatics methods for the characterization of viral quasispecies, molecular epidemiology and metagenomics. He is currently employed on a Wellcome Trust Bioresource Grant (WT108418AIA), which aims to drive substantial changes in the operations and public contributions of the International Committee on Taxonomy of Viruses (ICTV).

  • Marilyn J. Roossinck

    Marilyn J. Roossinck is a professor of virus ecology in the Departments of Plant Pathology and Environmental Microbiology, and Biology, at Pennsylvania State University, University Park, USA. She works in the Center for Infectious Disease Dynamics at the Huck Institutes of Life Sciences, Pennsylvania State University. She received her Ph.D. in microbiology and immunology from the University of Colorado School of Medicine, Denver, USA, in 1986. She and her team have been studying virus ecology and experimental evolution for 25 years, using plant and fungal viruses as models, and have published several seminal papers in this area. She is an expert in virus diversity and biodiversity, and has conducted extensive work on complex interactions between beneficial viruses and their hosts that are involved in the adaptation of plants and fungi to extreme environments. She is the United States representative to the International Committee on Taxonomy of Viruses (ICTV).

  • Sead Sabanadzovic

    Sead Sabanadzovic is a professor at Mississippi State University (MSU), USA. He received his M.Sc. and Ph.D. in plant virology from the Mediterranean Agronomic Institute of Bari (MAIB), Italy, and from the University of Bari Aldo Moro, Italy, in 1993 and 1997, respectively. Before joining MSU in 2004, he worked at the MAIB and briefly at the University of California, Davis (UC Davis), USA. His current research focuses on the multifaceted characterization of viromes of cultivated and non-cultivated plants and fungi. For more than a decade, he has been actively involved in virus taxonomy. Currently, he is an elected member of the International Committee on Taxonomy of Viruses (ICTV) Executive Committee.

  • Matthew B. Sullivan

    Matthew B. Sullivan is an associate professor at Ohio State University, Columbus, USA. He received his Ph.D., in 2004, in biological oceanography from the Woods Hole Oceanographic Institution, Massachusetts, USA, and the Massachusetts Institute of Technology (MIT), Cambridge, USA. His research explores the roles that microbial viruses have in oceans, soils and humans through the use of existing and novel quantitative approaches. As a Gordon and Betty Moore Foundation Investigator, his efforts have helped map and understand the global virosphere, and establish experimental and informatic approaches to link and explore virus–host interactions in nature and the laboratory.

  • Curtis A. Suttle

    Curtis A. Suttle is Professor of Earth, Ocean and Atmospheric Sciences, Microbiology and Immunology, and Botany, University of British Columbia, Vancouver, Canada. His research focuses on viruses and their diversity and roles in the global system. His work emphasizes that viruses are the most abundant and genetically diverse life forms in the oceans, and, as major agents of mortality, are crucial for global nutrient and energy cycling. He has been appointed as a fellow of the Royal Society of Canada, the Canadian Institute for Advanced Research, the American Academy of Microbiology and the Institute of Advanced Studies at Hong Kong University of Science and Technology (HKUST), and has been awarded the A.G. Huntsman, Timothy R. Parsons and G. Evelyn Hutchinson Medals for research excellence.

  • Robert B. Tesh

    Robert B. Tesh is Professor of Pathology, and of Microbiology and Immunology at the University of Texas Medical Branch (UTMB), Galveston, USA. He is currently the Director of the World Reference Center for Emerging Viruses and Arboviruses at UTMB. He spent 12 years as a research scientist with the National Institute of Allergy and Infectious Diseases (NIAID; part of the National Institutes of Health (NIH)), Bethesda, Maryland, USA, 15 years on the faculty of Yale University School of Medicine, New Haven, Connecticut, USA, and for the past 21 years he has worked at UTMB. He is author or co-author of more than 450 publications in the areas of arbovirology, medical entomology, virus ecology, epidemiology of zoonotic diseases, virus discovery and the characterization of novel viruses.

  • René A. van der Vlugt

    René A. van der Vlugt is Professor of Ecological Plant Virology in the Laboratory of Virology at Wageningen University, The Netherlands. He received his Ph.D. from Wageningen University, in 1993. Apart from the development and applications of plant virus diagnostics, his main interests are in virus identification and characterization, and the interactions between viruses, plants and their vectors. Currently, he and his team are using metagenomic approaches to study the diversity of plant viruses, and the evolution and adaptation of viral communities to different hosts and environmental conditions. He is currently a member of the Secoviridae Study Group of the International Committee on Taxonomy of Viruses (ICTV).

  • Arvind Varsani

    Arvind Varsani is a molecular virologist at The Biodesign Center for Fundamental and Applied Microbiomics and the School of Life Sciences at Arizona State University, USA. He works across ecosystems to study viral dynamics. His research group has a strong focus on the diversity, evolution and phylogeography of single-stranded DNA viruses. He is a member of the International Committee on Taxonomy of Viruses (ICTV) Executive and serves on two ICTV sub-committees (Geminiviridae and Circoviridae).

  • F. Murilo Zerbini

    F. Murilo Zerbini is a professor in the Department of Plant Pathology at the Federal University of Viçosa, Brazil. He received his Ph.D. from the University of California, Davis (UC Davis), in 1996. His main area of interest is the variability and evolution of geminiviruses (plant DNA viruses). Since the late 1990s, his group has been involved in the characterization of geminivirus populations that infect cultivated and non-cultivated plants in Brazil and neighbouring countries in South America, with the description of several new species. He has been a member of the Geminiviridae Study Group of the International Committee on Taxonomy of Viruses (ICTV) since 2002, and is currently a member of the ICTV Executive Committee.

Supplementary information

Word documents

  1. Supplementary information S1 (box) (48 KB)

    Classification of viruses derived from metagenomic sequences

Additional data