Archives of Virology

, Volume 161, Issue 8, pp 2351–2360 | Cite as

Taxonomy of the order Mononegavirales: update 2016

  • Claudio L. Afonso
  • Gaya K. Amarasinghe
  • Krisztián Bányai
  • Yīmíng Bào
  • Christopher F. Basler
  • Sina Bavari
  • Nicolás Bejerman
  • Kim R. Blasdell
  • François-Xavier Briand
  • Thomas Briese
  • Alexander Bukreyev
  • Charles H. Calisher
  • Kartik Chandran
  • Jiāsēn Chéng
  • Anna N. Clawson
  • Peter L. Collins
  • Ralf G. Dietzgen
  • Olga Dolnik
  • Leslie L. Domier
  • Ralf Dürrwald
  • John M. Dye
  • Andrew J. Easton
  • Hideki Ebihara
  • Szilvia L. Farkas
  • Juliana Freitas-Astúa
  • Pierre Formenty
  • Ron A. M. Fouchier
  • Yànpíng Fù
  • Elodie Ghedin
  • Michael M. Goodin
  • Roger Hewson
  • Masayuki Horie
  • Timothy H. Hyndman
  • Dàohóng Jiāng
  • Elliot W. Kitajima
  • Gary P. Kobinger
  • Hideki Kondo
  • Gael Kurath
  • Robert A. Lamb
  • Sergio Lenardon
  • Eric M. Leroy
  • Ci-Xiu Li
  • Xian-Dan Lin
  • Lìjiāng Liú
  • Ben Longdon
  • Szilvia Marton
  • Andrea Maisner
  • Elke Mühlberger
  • Sergey V. Netesov
  • Norbert Nowotny
  • Jean L. Patterson
  • Susan L. Payne
  • Janusz T. Paweska
  • Rick E. Randall
  • Bertus K. Rima
  • Paul Rota
  • Dennis Rubbenstroth
  • Martin Schwemmle
  • Mang Shi
  • Sophie J. Smither
  • Mark D. Stenglein
  • David M. Stone
  • Ayato Takada
  • Calogero Terregino
  • Robert B. Tesh
  • Jun-Hua Tian
  • Keizo Tomonaga
  • Noël Tordo
  • Jonathan S. Towner
  • Nikos Vasilakis
  • Martin Verbeek
  • Viktor E. Volchkov
  • Victoria Wahl-Jensen
  • John A. Walsh
  • Peter J. Walker
  • David Wang
  • Lin-Fa Wang
  • Thierry Wetzel
  • Anna E. Whitfield
  • Jiǎtāo Xiè
  • Kwok-Yung Yuen
  • Yong-Zhen Zhang
  • Jens H. Kuhn
Virology Division News


In 2016, the order Mononegavirales was emended through the addition of two new families (Mymonaviridae and Sunviridae), the elevation of the paramyxoviral subfamily Pneumovirinae to family status (Pneumoviridae), the addition of five free-floating genera (Anphevirus, Arlivirus, Chengtivirus, Crustavirus, and Wastrivirus), and several other changes at the genus and species levels. This article presents the updated taxonomy of the order Mononegavirales as now accepted by the International Committee on Taxonomy of Viruses (ICTV).


Soybean Cyst Nematode Emend Garter Snake Taxonomic Change Orchid Fleck Virus 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


The viral order Mononegavirales was established in 1991 to accommodate related viruses with nonsegmented, linear, single-stranded negative-sense RNA genomes. These viruses were initially assigned to three mononegaviral families: Filoviridae, Paramyxoviridae, and Rhabdoviridae [20, 21]. In subsequent years, these families continued to grow through the inclusion of numerous novel species and genera, and the order was therefore emended in 1995 [4], 1997 [23], 2000 [24], 2005 [25], and 2011 [8]. The families Bornaviridae and Nyamiviridae joined the other three mononegaviral families in 1996 [22] and 2014 [1, 11], respectively. In 2015, the Study Groups of the International Committee on Taxonomy of Viruses (ICTV) responsible for the taxonomy of the order and its five families embarked on a joint effort to assign unclassified mononegaviruses to existing or novel taxa and to streamline order nomenclature. Here we present a brief overview of the first round of these efforts, which by now is accepted by the ICTV Executive Committee and, thereby, is official taxonomy.

Taxonomic changes at the order level

In recent years, several mononegaviruses have been described that are only distantly related to the members of the families Bornaviridae, Filoviridae, Nyamiviridae, Paramyxoviridae, and Rhabdoviridae. These viruses include Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1) found in an ascomycete in China [16]; Sunshine Coast virus (SunCV; previously called Sunshine virus) isolated from Australian carpet pythons [10]; and Líshí spider virus 2 (LsSV-2), Sānxiá water strider virus 4 (SxWSV-4), Tǎchéng tick virus 6 (TcTV-6), Wēnzhōu crab virus 1 (WzCV-1), and Xīnchéng mosquito virus (XcMV) detected in Chinese arthropods [15]. To accommodate these viruses in the order and to appropriately reflect their phylogenetic relationships to other mononegaviral taxa, two new families and four free-floating genera were established: Mymonaviridae (accommodating SsNSRV-1), Sunviridae (SunCV), Anphevirus (XcMV), Arlivirus (LsSV-2), Chengtivirus (TcTV-6), Crustavirus (WzCV-1), and Wastrivirus (SxWSV-4). In addition, the paramyxoviral subfamily Pneumovirinae was elevated to family status (Pneumoviridae) because the members of this taxon are as closely related to filoviruses as to the members of the paramyxoviral subfamily Paramxyovirinae (now dissolved) (Table 1).
Table 1

Taxonomy of the order Mononegavirales as of 2016



Virus (abbreviation)

Family Bornaviridae


Elapid 1 bornavirus

Loveridge’s garter snake virus 1 (LGSV-1)

Mammalian 1 bornavirus*

Borna disease virus 1 (BoDV-1)

Borna disease virus 2 (BoDV-2)

Passeriform 1 bornavirus

canary bornavirus 1 (CnBV-1)

canary bornavirus 2 (CnBV-2)

canary bornavirus 3 (CnBV-3)

Passeriform 2 bornavirus

estrildid finch bornavirus 1 (EsBV-1)

Psittaciform 1 bornavirus

parrot bornavirus 1 (PaBV-1)

parrot bornavirus 2 (PaBV-2)

parrot bornavirus 3 (PaBV-3)

parrot bornavirus 4 (PaBV-4)

parrot bornavirus 7 (PaBV-7)

Psittaciform 2 bornavirus

parrot bornavirus 5 (PaBV-5)

Waterbird 1 bornavirus

aquatic bird bornavirus 1 (ABBV-1)

aquatic bird bornavirus 2 (ABBV-2)

Family Filoviridae


Lloviu cuevavirus*

Lloviu virus (LLOV)


Bundibugyo ebolavirus

Bundibugyo virus (BDBV)

Reston ebolavirus

Reston virus (RESTV)

Sudan ebolavirus

Sudan virus (SUDV)

Taï Forest ebolavirus

Taï Forest virus (TAFV)

Zaire ebolavirus*

Ebola virus (EBOV)


Marburg marburgvirus*

Marburg virus (MARV)

Ravn virus (RAVV)

Family Mymonaviridae


Sclerotinia sclerotimonavirus*

Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1)

Family Nyamiviridae


Midway nyavirus

Midway virus (MIDWV)

Nyamanini nyavirus*

Nyamanini virus (NYMV)

Sierra Nevada nyavirus

Sierra Nevada virus (SNVV)


Soybean cyst nematode socyvirus*

soybean cyst nematode virus 1 (SbCNV-1)

Family Paramyxoviridae


Atlantic salmon paramyxovirus*

Atlantic salmon paramyxovirus (AsaPV)


Avian paramyxovirus 2

avian paramyxovirus 2 (APMV-2)

Avian paramyxovirus 3

avian paramyxovirus 3 (APMV-3)

Avian paramyxovirus 4

avian paramyxovirus 4 (APMV-4)

Avian paramyxovirus 5

avian paramyxovirus 5 (APMV-5)

Avian paramyxovirus 6

avian paramyxovirus 6 (APMV-6)

Avian paramyxovirus 7

avian paramyxovirus 7 (APMV-7)

Avian paramyxovirus 8

avian paramyxovirus 8 (APMV-8)

Avian paramyxovirus 9

avian paramyxovirus 9 (APMV-9)

Avian paramyxovirus 10

avian paramyxovirus 10 (APMV-10)

Avian paramyxovirus 11

avian paramyxovirus 11 (APMV-11)

Avian paramyxovirus 12

avian paramyxovirus 12 (APMV-12)

Newcastle disease virus*

avian paramyxovirus 1 (APMV-1)a


Fer-de-Lance paramyxovirus*

Fer-de-Lance virus (FDLV)b


Cedar henipavirus

Cedar virus (CedV)

Ghanaian bat henipavirus

Kumasi virus (KV)c

Hendra virus*

Hendra virus (HeV)

Mojiang henipavirus

Mòjiāng virus (MojV)

Nipah virus

Nipah virus (NiV)


Canine distemper virus

canine distemper virus (CDV)

Cetacean morbillivirus

cetacean morbillivirus (CeMV)

Feline morbillivirus

feline morbillivirus (FeMV)d

Measles virus*

measles virus (MeV)

Peste-des-petits-ruminants virus

peste-des-petits-ruminants virus (PPRV)

Phocine distemper virus

phocine distemper virus (PDV)

Rinderpest virus

rinderpest virus (RPV)


Bovine parainfluenza virus 3

bovine parainfluenza virus 3 (BPIV-3)

Human parainfluenza virus 1

human parainfluenza virus 1 (HPIV-1)

Human parainfluenza virus 3

human parainfluenza virus 3 (HPIV-3)e

Porcine parainfluenza virus 1

porcine parainfluenza virus 1 (PPIV-1)

Sendai virus*

Sendai virus (SeV)f


Human parainfluenza virus 2

human parainfluenza virus 2 (HPIV-2)

Human parainfluenza virus 4

human parainfluenza virus 4a (HPIV-4a)

human parainfluenza virus 4b (HPIV-4b)

Mapuera virus

Mapuera virus (MapV)

Mumps virus*

mumps virus (MuV)

bat mumps virus (BMV)g

Parainfluenza virus 5

parainfluenza virus 5 (PIV-5)h

Porcine rubulavirus

La Piedad Michoacán Mexico virus (LPMV)i

Simian virus 41

simian virus 41 (SV-41)

Family Pneumoviridae


Avian metapneumovirus*

avian metapneumovirus (AMPV)j

Human metapneumovirus

human metapneumovirus (HMPV)


Bovine respiratory syncytial virus

bovine respiratory syncytial virus (BRSV)

Human respiratory syncytial virus*

human respiratory syncytial virus A2 (HRSV-A2)

human respiratory syncytial virus B1 (HRSV-B1)

human respiratory syncytial virus S2 (HRSV-S2)

Murine pneumonia virus

murine pneumonia virus (MPV)

Family Rhabdoviridae


Alfalfa dwarf cytorhabdovirus

alfalfa dwarf virus (ADV)

Barley yellow striate mosaic cytorhabdovirus

barley yellow striate mosaic virus (BYSMV)

Broccoli necrotic yellows cytorhabdovirus

broccoli necrotic yellows virus (BNYV)

Festuca leaf streak cytorhabdovirus

festuca leaf streak virus (FLSV)

Lettuce necrotic yellows cytorhabdovirus*

lettuce necrotic yellows virus (LNYV)

Lettuce yellow mottle cytorhabdovirus

lettuce yellow mottle virus (LYMoV)

Northern cereal mosaic cytorhabdovirus

northern cereal mosaic virus (NCMV)

Sonchus cytorhabdovirus 1

sonchus virus (SonV)

Strawberry crinkle cytorhabdovirus

strawberry crinkle virus (SCV)

Wheat American striate mosaic cytorhabdovirus

wheat American striate mosaic virus (WASMV)


Coffee ringspot dichorhavirus

coffee ringspot virus (CoRSV)

Orchid fleck dichorhavirus*

orchid fleck virus (OFV)k


Adelaide River ephemerovirus

Adelaide River virus (ARV)

Berrimah ephemerovirus

Berrimah virus (BRMV)

Bovine fever ephemerovirus*

bovine ephemeral fever virus (BEFV)l

Kotonkan ephemerovirus

kotonkan virus (KOTV)

Obodhiang ephemerovirus

Obodhiang virus (OBOV)


Aravan lyssavirus

Aravan virus (ARAV)

Australian bat lyssavirus

Australian bat lyssavirus (ABLV)

Bokeloh bat lyssavirus

Bokeloh bat lyssavirus (BBLV)

Duvenhage lyssavirus

Duvenhage virus (DUVV)

European bat 1 lyssavirus

European bat lyssavirus 1 (EBLV-1)

European bat 2 lyssavirus

European bat lyssavirus 2 (EBLV-2)

Ikoma lyssavirus

Ikoma lyssavirus (IKOV)

Irkut lyssavirus

Irkut virus (IRKV)

Khujand lyssavirus

Khujand virus (KHUV)

Lagos bat lyssavirus

Lagos bat virus (LBV)

Mokola lyssavirus

Mokola virus (MOKV)

Rabies lyssavirus*

rabies virus (RABV)

Shimoni bat lyssavirus

Shimoni bat virus (SHIBV)

West Caucasian bat lyssavirus

West Caucasian bat virus (WCBV)


Hirame novirhabdovirus

Hirame rhabdovirus (HIRV)

Oncorhynchus 1 novirhabdovirus*

infectious hematopoietic necrosis virus (IHNV)

Oncorhynchus 2 novirhabdovirus

viral hemorrhagic septicemia virus (VHSV)m

Snakehead novirhabdovirus

snakehead rhabdovirus (SHRV)


Datura yellow vein nucleorhabdovirus

datura yellow vein virus (DYVV)

Eggplant mottled dwarf nucleorhabdovirus

eggplant mottled dwarf virus (EMDV)

Maize fine streak nucleorhabdovirus

maize fine streak virus (MSFV)

Maize Iranian mosaic nucleorhabdovirus

maize Iranian mosaic virus (MIMV)

Maize mosaic nucleorhabdovirus

maize mosaic virus (MMV)

Potato yellow dwarf nucleorhabdovirus*

potato yellow dwarf virus (PYDV)

Rice yellow stunt nucleorhabdovirus

rice yellow stunt virus (RYSV)

rice transitory yellowing virus (RTYV)

Sonchus yellow net nucleorhabdovirus

sonchus yellow net virus (SYNV)

Sowthistle yellow vein nucleorhabdovirus

sowthistle yellow vein virus (SYVV)

Taro vein chlorosis nucleorhabdovirus

taro vein chlorosis virus (TaVCV)


Anguillid perhabdovirus

eel virus European X (EVEX)

Perch perhabdovirus*

perch rhabdovirus (PRV)

Sea trout perhabdovirus

lake trout rhabdovirus (LTRV)


Drosophila affinis sigmavirus

Drosophila affinis sigmavirus (DAffSV)

Drosophila ananassae sigmavirus

Drosophila ananassae sigmavirus (DAnaSV)

Drosophila immigrans sigmavirus

Drosophila immigrans sigmavirus (DImmSV)

Drosophila melanogaster sigmavirus*

Drosophila melanogaster sigmavirus (DMelSV)

Drosophila obscura sigmavirus

Drosophila obscura sigmavirus (DObsSV)

Drosophila tristis sigmavirus

Drosophila tristis sigmavirus (DTriSV

Muscina stabulans sigmavirus

Muscina stabulans sigmavirus (MStaSV)


Carp sprivivirus*

spring viremia of carp virus (SVCV)

Pike fry sprivivirus

grass carp rhabdovirus (GrCRV)

pike fry rhabdovirus (PFRV)

Tench rhabdovirus (TenRV)


Coastal Plains tibrovirus

Coastal Plains virus (CPV)

Tibrogargan tibrovirus*

Bivens Arm virus (BAV)

Tibrogargan virus (TIBV)


Durham tupavirus*

Durham virus (DURV)

Tupaia tupavirus

tupaia virus (TUPV)


Lettuce big-vein associated varicosavirus*

lettuce big-vein associated virus (LBVaV)n


Alagoas vesiculovirus

vesicular stomatitis Alagoas virus (VSAV)

Carajas vesiculovirus

Carajás virus (CJSV)

Chandipura vesiculovirus

Chandipura virus (CHPV)

Cocal vesiculovirus

Cocal virus (COCV)

Indiana vesiculovirus*

vesicular stomatitis Indiana virus (VSIV)

Isfahan vesiculovirus

Isfahan virus (ISFV)

Maraba vesiculovirus

Maraba virus (MARAV)

New Jersey vesiculovirus

vesicular stomatitis New Jersey virus (VSNJV)

Piry vesiculovirus

Piry virus (OIRYV)


Flanders virus

Flanders virus (FLAV)

Ngaingan virus

Ngaingan virus (NGAV)

Wongabel virus

Wongabel virus (WONV)

Family Sunviridae


Reptile sunshinevirus 1*

Sunshine Coast virus (SunCV)



Xincheng anphevirus*

Xīnchéng mosquito virus (XcMV)


Lishi arlivirus*

Líshí spider virus 2 (LsSV-2)


Tacheng chengtivirus*

Tǎchéng tick virus 6 (TcTV-6)


Wenzhou crustavirus*

Wēnzhōu crab virus 1 (WzCV-1)


Sanxia wastrivirus*

Sānxiá water strider virus 4 (SxWSV-4)

Listed are all mononegaviruses that have been classified into species. Asterisks denote type species

aIncludes: Newcastle disease virus (NDV) and pigeon paramyxovirus

bSynonym: anaconda paramyxovirus

cSynonym: GH-M74a virus

dAbbreviation as recently introduced in [26]

eHistorically, an isolate from a samango monkey (Cercopithecus mitis) in 1963 was long classified as a distinct species called simian agent 10 (SA-10), but was shown later to be HPIV-3. SA-10 was sometimes called SV-10

fSynonym: murine parainfluenza virus 1

gSynonym: bat paramyxovirus

hSynonym: simian virus 5

iSynonym: porcine rubulavirus

jSynonyms: avian pneumovirus, turkey rhinotracheitis virus

kSynonyms: citrus leprosis virus nuclear type, citrus necrotic spot virus

lSynonym Tzipori virus

mSynonyms: Egtved virus, Paralichthys olivaceus rhabdovirus

nSynonym: tobacco stunt virus

Taxonomic changes at the family level

The monogeneric family Bornaviridae was reorganized in 2015 by establishing five distinct species in the genus Bornavirus [2, 12] following a non-Latinized binomial species name format [29]. These efforts were continued in 2016 by expanding the genus by an additional two species (Elapid 1 bornavirus for Loveridge’s garter snake virus 1 [27] and Psittaciform 2 bornavirus for parrot bornavirus 5 [9, 18]) (Table 1).

The monogeneric family Nyamiviridae was expanded to include a second genus (Socyvirus) for the until-then free-floating nyamivirus species Soybean cyst nematode virus. This species name was changed to Soybean cyst nematode socyvirus to adhere to the non-Latinized binomial species name format [29] (Table 1).

The family Paramyxoviridae was emended by expanding the genus Avulavirus by three species (Avian paramyxovirus 10-12 for avian paramyxoviruses 10-12, respectively [5, 19, 28]), the genus Henipavirus by three species (Cedar henipavirus for Cedar virus [17], Ghanaian bat henipavirus for Kumasi virus [GH-M74a] [7], and Mojiang henipavirus for Mòjiāng virus [31]), the genus Morbillivirus by one species (Feline morbillivirus for feline morbillivirus [30]) and the genus Respirovirus by one species (Porcine parainfluenza virus 1 for porcine parainfluenza virus 1 [14]). The species Simian Virus 10 was dissolved on the evidence that simian virus 10 is an isolate of human parainfluenzavirus 3 rather than a distinct virus [13]. The genus Pneumovirus, now included in the new family Pneumoviridae, was renamed Orthopneumovirus to avoid confusion between family and genus members (Table 1).

The family Rhabdoviridae was expanded by two genera: Dichorhavirus (new; [6]) and Varicosavirus (previously free-floating outside of the order) to accommodate bisegmented plant viruses (coffee ringspot virus and orchid fleck virus; lettuce big-vein associated virus). The species Alfalfa dwarf cytorhabdovirus (for alfalfa dwarf virus [3]) was added to the genus Cytorhabdovirus. Finally, the non-Latinized binomial species name format [29] was applied throughout the family (Table 1).

A summary of the current, ICTV-accepted taxonomy of the order Mononegavirales is presented in Table 1.



We thank Laura Bollinger (NIH/NIAID Integrated Research Facility at Fort Detrick, Frederick, MD, USA) for critically editing the manuscript.

Thomas Briese, Ralf Dürrwald, Masayuki Horie, Jens H. Kuhn, Norbert Nowotny, Susan L. Payne, Dennis Rubbenstroth, Martin Schwemmle, Keizo Tomonaga: The members of the International Committee on Taxonomy of Viruses (ICTV) Bornaviridae Study Group; Gaya K. Amarasinghe, Christopher F. Basler, Sina Bavari, Alexander Bukreyev, Kartik Chandran, Olga Dolnik, John M. Dye, Hideki Ebihara, Pierre Formenty, Roger Hewson, Gary P. Kobinger, Jens H. Kuhn, Eric M. Leroy, Elke Mühlberger, Sergey V. Netesov, Jean L. Patterson, Janusz T. Paweska, Sophie J. Smither, Ayato Takada, Jonathan S. Towner, Viktor E. Volchkov, Victoria Wahl-Jensen: The members of the ICTV Filoviridae Study Group; Ralf G. Dietzgen, Andrew J. Easton, Jens H. Kuhn, Gael Kurath, Norbert Nowotny, Bertus K. Rima, Dennis Rubbenstroth, Nikos Vasilakis, Peter J. Walker: The members of the ICTV Mononegavirales Study Group; Ralf G. Dietzgen, Leslie L. Domier, Elodie Ghedin, Dàohóng Jiāng, Jens H. Kuhn, Nikos Vasilakis, David Wang: The members of the ICTV Nyamiviridae Study Group; Peter L. Collins, Andrew J. Easton, Ron A. M. Fouchier, Gael Kurath, Robert A. Lamb, Andrea Maisner, Rick E. Randall, Bertus K. Rima, Paul Rota, Lin-Fa Wang: The members of the ICTV Paramyxoviridae Study Group; Kim R. Blasdell, Charles H. Calisher, Ralf G. Dietzgen, Hideki Kondo, Gael Kurath, Ben Longdon, David M. Stone, Robert B. Tesh, Noël Tordo, Nikos Vasilakis, Peter J. Walker, Anna E. Whitfield: The members of the ICTV Rhabdoviridae Study Group.

Compliance with ethical standards

The views and conclusions contained in this document are those of the authors and should not be interpreted as necessarily representing the official policies, either expressed or implied, of the US Department of the Army, the US Department of Defense, the US Department of Health and Human Services, the Department of Homeland Security (DHS) Science and Technology Directorate (S&T) or of the institutions and companies affiliated with the authors. In no event shall any of these entities have any responsibility or liability for any use, misuse, inability to use, or reliance upon the information contained herein. The US departments do not endorse any products or commercial services mentioned in this publication.


This work was supported in part through Battelle Memorial Institute’s prime contract with the US National Institute of Allergy and Infectious Diseases (NIAID) under Contract No. HHSN272200700016I. A subcontractor to Battelle Memorial Institute who performed this work is: J. H. K., an employee of Tunnell Government Services, Inc. This work was also funded in part under Contract No. HSHQDC-07-C-00020 awarded by DHS S&T for the management and operation of the National Biodefense Analysis and Countermeasures Center (NBACC), a Federally Funded Research and Development Center (V. W.-J.); and National Institutes of Health (NIH) contract HHSN272201000040I/HHSN27200004/D04 (N. V., R. B. T.). Y. B. was supported by the Intramural Research Program of the NIH, National Library of Medicine.

Conflict of interest

The authors have no conflicts of interest.

Ethical approval

This article does not contain any studies with human participants or animals performed by any of the authors.


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Copyright information

© Springer-Verlag Wien (Outside the USA) 2016

Authors and Affiliations

  1. 1.Southeast Poultry Research Laboratory, Agricultural Research ServiceUS Department of AgricultureAthensUSA
  2. 2.Department of Pathology and ImmunologyWashington University School of MedicineSt. LouisUSA
  3. 3.Institute for Veterinary Medical Research, Centre for Agricultural ResearchHungarian Academy of SciencesBudapestHungary
  4. 4.Information Engineering BranchNational Center for Biotechnology Information, National Library of Medicine, National Institutes of HealthBethesdaUSA
  5. 5.Center for Microbial Pathogenesis, Institute for Biomedical SciencesGeorgia State UniversityAtlantaUSA
  6. 6.United States Army Medical Research Institute of Infectious Diseases, Fort DetrickFrederickUSA
  7. 7.Instituto de Patología Vegetal, Centro de Investigaciones Agropecuarias, Instituto Nacional de Tecnología AgropecuariaCórdobaArgentina
  8. 8.Consejo Nacional de Investigaciones Científicas y TécnicasBuenos AiresArgentina
  9. 9.CSIRO Health and BiosecurityAustralian Animal Health LaboratoryGeelongAustralia
  10. 10.Avian and Rabbit Virology Immunology and Parasitology UnitFrench Agency for Food, Environmental and Occupational Health and SafetyPloufraganFrance
  11. 11.Center for Infection and Immunity, Mailman School of Public HealthColumbia UniversityNew YorkUSA
  12. 12.Departments of Pathology and Microbiology & Immunology, Galveston National LaboratoryThe University of Texas Medical BranchGalvestonUSA
  13. 13.Arthropod-Borne and Infectious Diseases Laboratory, College of Veterinary Medicine and Biomedical SciencesColorado State UniversityFort CollinsUSA
  14. 14.Department of Microbiology and ImmunologyAlbert Einstein College of MedicineBronxUSA
  15. 15.State Key Laboratory of Agricultural Microbiology, The Provincial Key Lab of Plant Pathology of Húběi Province, College of Plant Science and TechnologyHuázhōng Agricultural UniversityWuhanChina
  16. 16.Integrated Research Facility at Fort Detrick (IRF-Frederick), Division of Clinical Research (DCR)National Institute of Allergy and Infectious Diseases (NIAID), National Institutes of Health (NIH)FrederickUSA
  17. 17.Respiratory Viruses Section, Laboratory of Infectious DiseasesNational Institute of Allergy and Infectious Diseases, National Institutes of HealthBethesdaUSA
  18. 18.Queensland Alliance for Agriculture and Food InnovationThe University of QueenslandSt. LuciaAustralia
  19. 19.Institute of VirologyPhilipps University MarburgMarburgGermany
  20. 20.Department of Crop SciencesUniversity of IllinoisChampaignUSA
  21. 21.IDT BiologikaDessau-RosslauGermany
  22. 22.School of Life SciencesUniversity of WarwickCoventryUK
  23. 23.Rocky Mountain Laboratories Integrated Research FacilityNational Institute of Allergy and Infectious Diseases, National Institutes of HealthHamiltonUSA
  24. 24.Embrapa Cassava and FruitsCruz das AlmasBrazil
  25. 25.World Health OrganizationGenevaSwitzerland
  26. 26.Department of Viroscience, Postgraduate School Molecular MedicineErasmus University Medical CenterRotterdamThe Netherlands
  27. 27.Department of Biology, Center for Genomics and Systems BiologyNew York UniversityNew YorkUSA
  28. 28.Plant PathologyUniversity of KentuckyLexingtonUSA
  29. 29.Public Health England, Porton DownWiltshireUK
  30. 30.Joint Faculty of Veterinary Medicine, Transboundary Animal Diseases Research CenterKagoshima UniversityKagoshimaJapan
  31. 31.School of Veterinary and Life SciencesMurdoch UniversityMurdochAustralia
  32. 32.Núcleo de Apoio à Pesquisa em Microscopia Eletrônica Aplicada a Agricultura, Escola Superior de Agricultura “Luiz de Queiroz”Universidade de São PauloPiracicabaBrazil
  33. 33.Special Pathogens Program, National Microbiology LaboratoryPublic Health Agency of CanadaWinnipegCanada
  34. 34.Institute of Plant Science and ResourcesOkayama UniversityKurashikiJapan
  35. 35.US Geological Survey Western Fisheries Research CenterSeattleUSA
  36. 36.Department of Molecular BiosciencesNorthwestern UniversityEvanstonUSA
  37. 37.Howard Hughes Medical InstituteNorthwestern UniversityEvanstonUSA
  38. 38.Centre International de Recherches Médicales de Franceville, Institut de Recherche pour le DéveloppementFrancevilleGabon
  39. 39.State Key Laboratory for Infectious Disease Prevention and Control, National Institute for Communicable Disease Control and Prevention, Chinese Center for Disease Control and PreventionBeijingChina
  40. 40.Collaborative Innovation Center for Diagnosis and Treatment of Infectious DiseasesHangzhouChina
  41. 41.Wēnzhōu Center for Disease Control and PreventionWenzhouChina
  42. 42.Department of GeneticsUniversity of CambridgeCambridgeUK
  43. 43.Department of Microbiology and National Emerging Infectious Diseases LaboratoryBoston University School of MedicineBostonUSA
  44. 44.Novosibirsk State UniversityNovosibirskRussia
  45. 45.Institute of VirologyUniversity of Veterinary MedicineViennaAustria
  46. 46.Department of Basic Medical Sciences, College of MedicineMohammed Bin Rashid University of Medicine and Health SciencesDubaiUnited Arab Emirates
  47. 47.Department of Virology and ImmunologyTexas Biomedical Research InstituteSan AntonioUSA
  48. 48.Department of Veterinary Pathobiology, College of Veterinary Medicine and Biomedical SciencesTexas A&M UniversityCollege StationUSA
  49. 49.Center for Emerging and Zoonotic Diseases, National Institute for Communicable Diseases of the National Health Laboratory ServiceSandringham-JohannesburgGautengSouth Africa
  50. 50.Biomedical Sciences Research ComplexUniversity of St. AndrewsSt. AndrewsUK
  51. 51.Centre for Experimental Medicine, School of Medicine, Dentistry and Biomedical SciencesThe Queen’s University of BelfastBelfastUK
  52. 52.National Center for Immunization and Respiratory Diseases, Centers for Disease Control and PreventionAtlantaUSA
  53. 53.Institute for Virology, Faculty of MedicineMedical Center—University of Freiburg, University of FreiburgFreiburgGermany
  54. 54.CBR Division, Dstl, Porton DownSalisburyUK
  55. 55.Department of Microbiology, Immunology, and Pathology, College of Veterinary Medicine and Biomedical SciencesColorado State UniversityFort CollinsUSA
  56. 56.Centre for Environment, Fisheries and Aquaculture Science WeymouthDorsetUK
  57. 57.Division of Global EpidemiologyHokkaido University Research Center for Zoonosis ControlSapporoJapan
  58. 58.Istituto Zooprofilattico Sperimentale delle Venezie, Department of Comparative Biomedical Sciences, National/OIE Reference Laboratory for Newcastle Disease and Avian InfluenzaFAO Reference Centre for Animal Influenza and Newcastle Disease, OIE Collaborating Centre for Diseases at the Human-Animal InterfaceLegnaroItaly
  59. 59.Wǔhàn Center for Disease Control and PreventionWuhanChina
  60. 60.Institute for Virus ResearchKyoto UniversityKyotoJapan
  61. 61.Institut Pasteur, Unité des Stratégies AntiviralesParisFrance
  62. 62.Institut Pasteur de GuinéeConakryGuinea
  63. 63.Viral Special Pathogens Branch, Division of High-Consequence Pathogens PathologyNational Center for Emerging and Zoonotic Infectious Diseases, Centers for Disease Control and PreventionAtlantaUSA
  64. 64.Center for Biodefense and Emerging Infectious Diseases, Department of PathologyThe University of Texas Medical BranchGalvestonUSA
  65. 65.Center for Tropical Diseases, Institute for Human Infections and ImmunityThe University of Texas Medical BranchGalvestonUSA
  66. 66.Wageningen University and ResearchWageningenThe Netherlands
  67. 67.Molecular Basis of Viral Pathogenicity, CIRI, INSERM U1111, CNRS UMR5308, Université de Lyon, Université Claude Bernard Lyon 1, Ecole Normale Supérieure de LyonLyonFrance
  68. 68.National Biodefense Analysis and Countermeasures Center, Fort DetrickFrederickUSA
  69. 69.Departments of Molecular Microbiology and Pathology and ImmunologyWashington University School of MedicineSt. LouisUSA
  70. 70.Department of Agriculture and FisheriesBiosecurity QueenslandBrisbaneAustralia
  71. 71.Program in Emerging Infectious Diseases, Duke-NUS Graduate Medical SchoolSingaporeSingapore
  72. 72.DLR Rheinpfalz, Institute of Plant ProtectionNeustadt an der WeinstrasseGermany
  73. 73.Plant PathologyKansas State UniversityManhattanUSA
  74. 74.State Key Laboratory of Emerging Infectious Diseases, Department of MicrobiologyUniversity of Hong Kong, Hong Kong Special Administrative RegionHong KongChina

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