2022 taxonomic update of phylum Negarnaviricota (Riboviria: Orthornavirae), including the large orders Bunyavirales and Mononegavirales

Abstract

In March 2022, following the annual International Committee on Taxonomy of Viruses (ICTV) ratification vote on newly proposed taxa, the phylum Negarnaviricota was amended and emended. The phylum was expanded by two new families (bunyaviral Discoviridae and Tulasviridae), 41 new genera, and 98 new species. Three hundred forty-nine species were renamed and/or moved. The accidentally misspelled names of seven species were corrected. This article presents the updated taxonomy of Negarnaviricota as now accepted by the ICTV.

Full text

J. H. Kuhn etal.
1 3
MassimoTurina158 · IoannisE.Tzanetakis159 · AnnaMariaVaira158 · BernadettevandenHoogen160 ·
BertVanmechelen161 · NikosVasilakis(Νίκος Βασιλάκης)91 · MartinVerbeek162 · SusannevonBargen10 ·
JiroWada(和田治郎)163 · VictoriaWahl164 · PeterJ.Walker165 · AnnaE.Whiteld166 · JohnV.Williams167 ·
YuriI.Wolf168 · JunkiYamasaki(山﨑淳紀)169· HironobuYanagisawa(栁澤広宣)170 · GongyinYe(叶恭银)171 ·
Yong‑ZhenZhang(张永振)172· ArnnnLoddenØkland173
This is a US Government work and not under copyright protection in the US; foreign copyright protection may apply 2022
Abstract
In March 2022, following the annual International Committee on Taxonomy of Viruses (ICTV) ratification vote on newly
proposed taxa, the phylum Negarnaviricota was amended and emended. The phylum was expanded by two new families
(bunyaviral Discoviridae and Tulasviridae), 41 new genera, and 98 new species. Three hundred forty-nine species were
renamed and/or moved. The accidentally misspelled names of seven species were corrected. This article presents the updated
taxonomy of Negarnaviricota as now accepted by the ICTV.
Introduction
The phylum Negarnaviricota was established in 2019 by the
International Committee on Taxonomy of Viruses (ICTV)
for negative-sense RNA viruses that can be connected evo-
lutionarily through the possession of virally encoded RNA-
directed RNA polymerases (RdRps). The phylum includes
two subphyla, Haploviricotina and Polyploviricotina, for
negative-sense RNA viruses that encode RdRps with or
without mRNA capping activity, respectively. The two sub-
phyla include four classes (Chunqiuviricetes, Milnevirice-
tes, Monjiviricetes, and Yunchangviricetes) and two classes
(Ellioviricetes and Insthoviricetes), respectively [33, 59, 74].
The phylum was last amended/emended in 2021 [34]. Here,
we present the changes that were proposed to the phylum
via official taxonomic proposals (TaxoProps) in 2021 and
accepted by the ICTV in March and May 2022 [23, 70].
These changes are now part of the official ICTV taxonomy
[24].
Taxonomic changes insubphylum
Haploviricotina
Chunqiuviricetes
The names of all species in the class were changed to binomi-
als as required by the recently amended International Code
of Virus Classification and Nomenclature (ICVCN) [25, 69,
81] (TaxoProp 2021.018M.R.Negarnaviricota_sprename).
Milneviricetes
The names of all species in the class were changed to bino-
mials as required by the recently amended ICVCN [25, 69,
81] (TaxoProp 2021.014P.R.Aspiviridae_rename).
Monjiviricetes
Family Aliusviridae
The species name Olluvirus shayangense was
corrected to Ollusvirus shayangense (TaxoProp
2021.042M.R.Corrections_Riboviria_Duplodnaviria).
Family Artoviridae
The names of all species in the family were changed to
binomials as required by the recently amended ICVCN [25,
69, 81] (TaxoProp 2021.009M.R.Artoviridae_sprename).
Genus Peropuvirus was expanded by one new species,
Peropuvirus melongenae, for Solanum melongena rhabdo-
like virus (SmRLV) identified in a metagenomic study of
eggplant (Solanum melongena L.) leaf tissue sampled in
Zhènjiāng (镇江市), Jiāngsū Province (江苏省), China [78]
(TaxoProp 2021.029M.R.Peropuvirus_1nsp).
Family Bornaviridae
The names of all species in the family were changed to
binomials as required by the recently amended ICVCN [25,
69, 81] (2021.010M.R.Bornaviridae_sprename).
Genus Orthobornavirus was expanded by one new spe-
cies, Orthobornavirus caenophidiae, for Caribbean water-
snake bornavirus (CWBV) and Mexican black-tailed rat-
tlesnake bornavirus (MRBV) identified in archived raw
transcriptomic read data of a Caribbean watersnake (colubrid
Tretanorhinus variabilis A.M.C. Duméril, Bibron & A.H.A.

Taxonomic update of phylum Negarnaviricota
1 3
Duméril, 1854) and a Mexican black-tailed rattlesnake (vip-
erid Crotalus molossus nigrescens Gloyd, 1936), respectively
[49] (TaxoProp 2021.021M.R.Orthobornavirus_1nsp).
in Hángzhōu (杭州市), Zhèjiāng Province (浙江省),
China [72];
• genus Copasivirus was created to include two new spe-
cies:
o Copasivirus manlyvaleense for Jimsystermes virus
(JIMV) discovered by HTS in termites (termitid
Occasitermes sp.) sampled in Manly Vale, New
South Wales, Australia [35]; and
o Copasivirus ivindoense for isopteran arli-related
virus OKIAV103 (IARV-103) discovered by HTS
in termites (rhinotermitid Coptotermes sp.) sampled
in Ivindo National Park (Parc national d'Ivindo),
Ogooué-Ivindo Province, Gabon [27];
• genus Cybitervirus was created to include new species
Cybitervirus niederense for coleopteran arli-related virus
OKIAV107 (CARV-107) discovered by HTS in beetles
(dytiscid Cybister lateralimarginalis (De Geer, 1774))
sampled in Lüchow-Dannenberg, Lower Saxony (Nied-
ersachsen), Germany [27];
• genus Phelinovirus was created to include new species
Phelinovirus aphidis for hymenopteran arli-related virus
OKIAV99 (HARV-99) discovered by HTS in parasi-
toid wasps (aphelinid Aphelinus abdominalis (Dalman,
1820)) in laboratory culture samples of unknown geo-
graphical origin [27];
• genus Rivapovirus was created to include new species
Rivapovirus aleyrodidae for hemipteran arli-related virus
OKIAV94 (HARV-94) discovered by HTS in greenhouse
whiteflies (aleyrodid Trialeurodes vaporariorum West-
wood, 1856) of commercial laboratory cultures sampled
in Brandenburg, Germany [27];
• genus Stylovirus was created to include new species
Stylovirus niederense for strepsipteran arli-related virus
OKIAV104 (SARV-104) discovered by HTS in endo-
parasitic insects (stylopid Stylops melittae Kirby, 1802)
sampled in Osnabrück, Lower Saxony (Niedersachsen),
Germany [27];
• genus Supelovirus was created to include the new species
Supelovirus thailandense for blattodean arli-related virus
OKIAV102 (BARV-102) discovered by HTS in brown-
banded cockroaches (ectobiid Supella longipalpa Fab-
ricius, 1798) of laboratory culture samples originating
from Thailand [27];
• genus Synelinevirus was created to include two new spe-
cies:
o Synelinevirus paranaense for Linepithema humile
rhabdo-like virus 1 (LhuRLV-1) 1 discovered by
HTS in Argentine ants (dolichoderine Linepithema
Family Filoviridae
Family Filoviridae was expanded by two genera:
• genus Oblavirus was established to include one new
species, Oblavirus percae, for Oberland virus (OBLV)
discovered by high-throughput sequencing (HTS) in
European perch (percid Perca fluviatilis Linnaeus, 1758)
sampled in Switzerland [20] (TaxoProp 2021.011M.R.Fi
loviridae_1ngen_3nsp); and
• genus Tapjovirus was established to include one new
species, Tapjovirus bothropis, for Tapajós virus (TAPV)
discovered in sequence read archive (SRA) data obtained
from a common lancehead (viperid Bothrops atrox
(Linnaeus, 1758)) sampled in Brazil [21] (TaxoProp
2021.005M.R.Filoviridae_1ngen_1nsp).
Genus Thamnovirus was expanded by two new species,
Thamnovirus percae for Fiwi virus (FIWIV) and Thamnovi-
rus kanderense for Kander virus (KNDV), both discovered
by HTS in European perch (percid Perca fluviatilis Linnaeus,
1758) sampled in Switzerland [20] (TaxoProp 2021.011M.R
.Filoviridae_1ngen_3nsp).
Family Lispiviridae
Family Lispiviridae was thoroughly revised (TaxoProp
2021.016M.R.Lispiviridae_16ngen_13nsp):
• in genus Arlivirus, species Lishi arlivirus was renamed
Arlivirus arachnae. Species Gerrid arlivirus, Hubei
arlivirus, Odonate arlivirus, Tacheng arlivirus, and
Wuchang arlivirus were renamed Sanstrivirus gerridis,
Leocovirus coleopteris, Damravirus dentatis, Gani-
avirus tachengense, and Nematovirus wuchangense,
respectively, and moved into the newly created genera
Sanstrivirus, Leocovirus, Damravirus, Ganiavirus, and
Nematovirus, respectively;
• genus Anicalvirus was created to include the new species
Anicalvirus hangzhouense for Anisopteromalus caland-
rae negative-strand RNA virus 2 (AcNSRV-2) discovered
by HTS in a parasitoid wasp (pteromalid Anisoptero-
malus calandrae (Howard, 1881)) sampled in Hángzhōu
(杭州市), Zhèjiāng Province (浙江省), China [72];
• genus Anidravirus was created to include the new spe-
cies Anidravirus hangzhouense for Anisopteromalus
calandrae negative-strand RNA virus 1 (AcNSRV-1)
discovered by HTS in parasitoid wasps (pteromalid
Anisopteromalus calandrae (Howard, 1881)) sampled

J. H. Kuhn etal.
1 3
humile (Mayr, 1868)) sampled in Catalonia (Cata-
luña), Spain [68]; and
o Synelinevirus bonnense for hymenopteran arli-
related virus OKIAV98 (HARV-98) discovered by
HTS in gall wasps (cynipid Synergus umbraculus
Walker, 1874) sampled in Bonn, North Rhine-West-
phalia (Nordrhein-Westfalen), Germany [27];
• genus Usmuvirus was created to include the new species
Usmuvirus newyorkense for Amsterdam virus (AMSV)
discovered by HTS in house mice (murid Mus musculus
Linnaeus, 1758) sampled in New York, New York, USA
[73]; and
• genus Xenophyvirus was created to include the new spe-
cies Xenophyvirus mathesonense for hemipteran arli-
related virus OKIAV95 (HARV-95) discovered by HTS
in moss bugs (peloridiid Xenophyes metoponcus Burck-
hardt, 2011) sampled in Lake Matheson, South Island
Westland District, New Zealand [27].
Family Mymonaviridae
Genus Auricularimonavirus was expanded by one new
species, Auricularimonavirus bondarzewiae, for Bondar-
zewia berkeleyi negative-strand RNA virus 1 (BbNSRV-1)
discovered by HTS in stump blossoms (bondarzewiaceaen
Bondarzewia berkeleyi (Fr.) Bondartsev & Singer (1941))
[65] (TaxoProp 2021.041M.R.Mymonaviridae_5nsp).
Genus Botrytimonavirus was expanded by one new spe-
cies, Botrytimonavirus alphabotrytidis, for Botrytis cinerea
negative-stranded RNA virus 5 (BcNSRV-5) and Sclerotinia
sclerotiorum negative-stranded RNA virus 10 (SsNSRV-10)
discovered in metatranscriptomes of sclerotiniaceaen fungi
(Botrytis cinerea Pers. (1794) and Sclerotinia sclerotiorum
(Lib.) de Bary (1884) sampled in Italy/Spain and China,
respectively) [26, 53] (TaxoProp 2021.041M.R.Mymonaviri
dae_5nsp). Sclerotinia sclerotiorum negative-stranded RNA
virus 11 (SsNSRV-11) discovered in a Sclerotinia sclerotio-
rum (Lib.) de Bary (1884) metatranscriptome dataset [26]
was assigned to the already established species Botrytimo-
navirus botrytidis (TaxoProp 2021.041M.R.Mymonavirid
ae_5nsp).
Genus Sclerotimonavirus was expanded by three new spe-
cies (TaxoProp 2021.041M.R.Mymonaviridae_5nsp):
• Sclerotimonavirus alphabotrytidis for Botrytis cinerea
negative-stranded RNA virus 3 (BcNSRV-3) and Scle-
rotinia sclerotiorum negative-stranded RNA virus 9
(SsNSRV-9) discovered in metatranscriptomes of scle-
rotiniaceaen fungi (Botrytis cinerea Pers. (1794) and
Sclerotinia sclerotiorum (Lib.) de Bary (1884) sampled
in Italy/Spain and China, respectively) [26, 53];
• Sclerotimonavirus betabotrytidis for Botrytis cinerea
negative-stranded RNA virus 4 (BcNSRV-4), discovered
in the metatranscriptome of a sclerotiniaceaen fungus
(Botrytis cinerea Pers. (1794)) sampled in Italy/Spain
[53]; and
• Sclerotimonavirus cryphonectriae for Cryphonectria
parasitica sclerotimonavirus 1 (CpSV-1), discovered by
HTS in a cryphonectriaceaen fungus (Cryphonectria
parasitica (Murrill) M. E. Barr (1978)) sampled in Azer-
baijan [18].
Family Nyamiviridae
The names of all species in the family were changed to
binomials as required by the recently amended ICVCN [25,
69, 81] (TaxoProp 2021.020M.R.Nyamiviridae_sprename).
Genus Formivirus was expanded by one new species,
Formivirus solenopsi, for Solenopsis invicta virus 15
(SoINV-15) identified in the transcriptome of red fire ants
(myrmicine Solenopsis invicta Buren, 1972) sampled in the
US and Taiwan [77] (TaxoProp 2021.019M.R.Nyamivirid
ae_3nsp).
Genus Nyavirus was expanded by two new species (Taxo-
Prop 2021.019M.R.Nyamiviridae_3nsp):
• Nyavirus argatis, for Sekira virus (SEKRV) identified in
the RNA virome of a pool of ticks (argasid Argas japoni-
cus Yamaguti, Clifford and Tipton, 1968) sampled in
Mishima (三島市), Shizuoka Prefecture (静岡県), Japan
[28]; and
• Nyavirus somateriae for Jeremy Point nyavirus (JPNV)
identified in the spleen of a common eider (anatid Soma-
teria mollissima (Linnaeus, 1758)) collected at Jeremy
Point, Massachusetts, USA, in 2013 (unpublished; Gen-
Bank: MN045233).
Family Paramyxoviridae
Genus Jeilongvirus was expanded by eight new species
(TaxoProp: 2021.014M.R.Jeilongvirus_8nsp):
• Jeilongvirus anhuiense for Miniopterus schreibersii para-
myxovirus (MisPV) discovered by HTS in Schreibers’s
long-fingered bats (miniopterid Miniopterus schreibersii
(Kuhl, 1817)) sampled in China [75];
• Jeilongvirus apodemi for rodent paramyxovirus (RoPV)
discovered by HTS in a Korean field mouse (murid
Apodemus peninsulae (Thomas, 1907)) sampled in China
[76];
• Jeilongvirus comorosense for bat paramyxovirus 16797
(BatPV-1) identified in a Comoran long-fingered bat
(miniopterid Miniopterus griveaudi Harrison, 1959) sam-
pled in Comoros (unpublished; GenBank: MG203877);

Taxonomic update of phylum Negarnaviricota
1 3
• Jeilongvirus erinacei for belerina virus (BeV) discovered
by HTS in West European hedgehogs (erinaceid Erina-
ceus europaeus Linnaeus, 1758) sampled in Belgium
[66];
• Jeilongvirus felis for feline paramyxovirus (FPaV) identi-
fied in domestic cats (feline Felis catus Linnaeus, 1758)
sampled in Germany [55, 61];
• Jeilongvirus madagascarense for bat paramyxovirus
17770 (BatPV-2) identified in a Comoran long-fin-
gered bat (miniopterid Miniopterus griveaudi Harrison,
1959) sampled in Madagascar (unpublished; GenBank:
MG203878);
• Jeilongvirus murinae for bat paramyxovirus (BatPV-
3) identified in a greater tube-nosed bat (vespertilionid
Murina leucogaster (Milne-Edwards, 1872) sampled in
China [75]; and
• Jeilongvirus rungweense for ruloma virus (RulV)
detected by HTS in a Machangu’s brush-furred rat (murid
Lophuromys machangui Verheyen, Hulselmans, Dierckx,
Mulungu, Leirs, Corti & Verheyen, 2007) sampled in
Rungwe District, Mbeya Region, Tanzania [67].
Family Rhabdoviridae
The names of all species in the family were changed to
binomials as required by the recently amended ICVCN [25,
69, 81] (TaxoProps 2021.036M.R.Rhabdoviridae_sprename,
2021.042M.R.Corrections_Riboviria_Duplodnaviria, and
2021.043M.R.Corrections_Mononegavirales_Bunyavirales).
Five new genera were created and included in the subfam-
ily Alpharhabdovirinae:
• genus Amplylivirus, including one novel species,
Amplylivirus cinereus, for frog lyssa-like virus 1 (FLLV-
1) detected in an American green tree frog (hylid Dryo-
phytes cinereus (Schneider, 1799)) (unpublished; Gen-
Bank: MK473367) (2021.035M.R.Rhabdoviridae_2nge
n_2nsp);
• genus Cetarhavirus for two new species (2021.003M.R.
Alpharhabdovirinae_3ngen_7nsp):
o Cetarhavirus laganorhynchus for dolphin rhabdovi-
rus (DRV) isolated from a white-beaked dolphin
(delphinid Lagenorhynchus albirostris (Gray, 1846))
sampled in Schiermonnikoog, Friesland, Holland
[45]; and
o Cetarhavirus phocoena for harbour porpoise rhab-
dovirus (HPRV) isolated from a harbour porpoise
(phocoenid Phocoena phocoena (Linnaeus, 1758))
stranded off the coast of Alaska [15];
• genus Replylivirus, including one novel species, Replyli-
virus allogus, for anole lyssa-like virus 1 (ALLV-1) iden-
tified in RNAseq data from a Spanish flag anole (dac-
tyloid Anolis allogus Barbour & Ramsden, 1919) [22]
(2021.035M.R.Rhabdoviridae_2ngen_2nsp);
• genus Siniperhavirus for two new species (2021.003M.R
.Alpharhabdovirinae_3ngen_7nsp):
o Siniperhavirus chuatsi for Siniperca chuatsi rhab-
dovirus (SRCV) isolated from mandarin fish (sin-
ipercid Siniperca chuatsi (Basilewsky, 1855))
collected in Guǎngdōng Province (广东省),
China [86, 87]; Chinese rice-field eel rhabdovirus
(CrERV) isolated from Chinese ricefield eels (syn-
branchid Monopterus albus (Zuiew, 1793)) sam-
pled in Qiánjiāng (潜江市), Húběi Province (湖
北省), China [38]; hybrid snakehead rhabdovirus
(HSHRV) isolated from hybrid snakehead (chan-
nid Channa maculata × Channa argus) sampled in
Fóshān (佛山市), Guǎngdōng Province (广东省),
China [80]; and Micropterus salmoides rhabdovirus
(MSRV) isolated from largemouth bass (centrarchid
Micropterus salmoides (Lacépède, 1802)) sampled
in Yúháng (余杭区), Zhèjiāng Province (浙江省),
China [40]; and
o Siniperhavirus zoarces for harbour eelpout rhab-
dovirus (EPRV) detected by HTS in eelpout (zoarcid
Zoarces viviparous (Linnaeus, 1758)) near Stock-
holm, Sweden [2];
• genus Scophrhavirus for two new species
(2021.003M.R.Alpharhabdovirinae_3ngen_7nsp):
o Scophrhavirus chanodychthys for Wuhan redfin cul-
ter dimarhabdovirus (WhRFCRV) detected by high-
throughput sequencing in redfin culter (cyprinid
Chanodichthys erythropterus (Basilewsky, 1855))
collected in “Húběi Province (湖北省)”, China [58];
and
o Scophrhavirus maximus for Scophthalmus maxi-
mus rhabdovirus (SMRV) isolated from turbot fish
(scophthalmid Scophthalmus maximus (Linnaeus,
1758)) sampled in Shāndōng Province (山东省),
China [82].
Genus Alphanucleorhabdovirus was expanded by one
new species, Alphanucleorhabdovirus joa, for joa yellow
blotch associated virus (JYBaV) identified in joá (solanace-
aen Solanum aculeatissimum L.) plants sampled in Manaus,
State of Amazonas (Estado de Amazonas), Brazil [11] (Tax-
oProp 2021.001M.A.v1.Betarhabdovirinae_7nsp).
Genus Betanucleorhabdovirus was expanded by one new
species, Betanucleorhabdovirus bacopae, for Bacopa mon-
nieri virus 2 (BmV-2) identified in transcriptomic data of
water hyssop (plantaginaceaen Bacopa monnieri L.) tissues

J. H. Kuhn etal.
1 3
sampled in Lucknow, Uttar Pradesh, India [60] (TaxoProp
2021.001M.A.v1.Betarhabdovirinae_7nsp).
Genus Cytorhabdovirus was expanded by five new spe-
cies (TaxoProp 2021.001M.A.v1.Betarhabdovirinae_7nsp):
• Cytorhabdovirus bacopae for Bacopa monnieri virus 1
(BmV-1) identified in transcriptomic data of water hys-
sop (plantaginaceaen Bacopa monnieri L.) tissues sam-
pled in Lucknow, Uttar Pradesh, India [60];
• Cytorhabdovirus chrysanthemi for chrysanthemum yel-
low dwarf associated virus (CYDaV) identified in grey-
leaved euryops (asteraceaen Euryops pectinatus (L.)
Cass.) plant sampled in Chóngqìng (重庆市), China [37];
• Cytorhabdovirus cucurbitae for cucurbit cytorhabdovi-
rus 1 (CuCV-1) identified in zucchini (cucurbitaceaen
Cucurbita pepo L.) plants sampled in Greece [44];
• Cytorhabdovirus broussonetiae for paper mulberry
mosaic associated virus (PMuMaV) identified in paper
mulberry (moraceaen Broussonetia papyrifera L.) plants
sampled in Chóngqìng (重庆市), China [51]; and
• Cytorhabdovirus rosae for rose virus R (RVR), identified
in a rose ‘Hugh Dickson’ plant from Maryland, USA [5].
Genus Hapavirus was expanded by two new species
(TaxoProp 2021.004M.R.Alpharhabdovirinae_13nsp):
• Hapavirus bangoran for Bangoran virus (BGNV) found
in mosquitoes (culicid Culex perfuscus Edwards, 1914)
sampled in Bangoran, Bamingui-Bangoran Prefecture,
Central African Republic [6, 14]; and
• Hapavirus porton for Porton virus (PORV) found in mos-
quitoes (culicid Mansonia uniformis (Theobald, 1901))
sampled in Sarawak state, Malaysia [6].
Genus Ledantevirus was expanded by one new species,
Ledantevirus taiyi, for Taiyi bat virus (TYBV) found in Chi-
nese rufous horseshoe bats (rhinolophid Rhinolophus sinicus
K. Andersen, 1905) sampled in Xiánníng (咸宁市), Húběi
Province (湖北省), China [39] (TaxoProp 2021.004M.R.A
lpharhabdovirinae_13nsp).
Genus Perhabdovirus was expanded by one new species,
Perhabdovirus leman, for Leman virus (LeRV) isolated from
perch fry sampled from Lake Geneva (Lac Léman), France [47]
(TaxoProp 2021.003M.R.Alpharhabdovirinae_3ngen_7nsp).
Genus Sunrhavirus was expanded by eight new species
(TaxoProp 2021.004M.R.Alpharhabdovirinae_13nsp):
• Sunrhavirus sandjimba for Sandjimba virus (SJAV)
found in sedge warblers (acrocephalid Acrocephalus
schoenobaenus (Linnaeus, 1758)) collected at the Land-
jia river, Ombella-M'Poko Prefecture, Central African
Republic [6, 14];
• Sunrhavirus nasoule for Nasoule virus (NASV) found
in little greenbuls (pycnonotid Eurillas virens (Cassin,
1857)) sampled in Nasoule, Mambéré-Kadéi Prefecture,
Central African Republic [6, 14];
• Sunrhavirus boteke for Boteke virus (BOTV) found
in mosquitoes (culicid Coquillettidia maculipennis
(Theobald, 1911)) sampled in Boteke, Ombella-M'Poko
Prefecture, Central African Republic [6, 14];
• Sunrhavirus bimbo for Bimbo virus (BBOV) found in
yellow-crowned bishop (ploceid Euplectes afer (Gmelin,
1789)) sampled in Kolongo, Ombella-M'Poko Prefecture,
Central African Republic [6, 14];
• Sunrhavirus kolongo for Kolongo virus (KOLV) found in
yellow-crowned bishop (ploceid Euplectes afer (Gmelin,
1789)) sampled in Bangui, Central African Republic [6, 14];
• Sunrhavirus ouango for Ouango virus (OUAV) found in
black-headed weaves (ploceid Ploceus melanocephalus
(Linnaeus, 1758)) collected at the Landjia river, Ombella-
M'Poko Prefecture, Central African Republic [6, 14];
• Sunrhavirus alexandria for Burg el Arab virus (BEAV)
found in lesser whitethroats (sylviid Curruca curruca
(Linnaeus, 1758)) sampled in Borg El Arab/Burj al-'Arab,
Alexandria Governorate, Egypt [6]; and
• Sunrhavirus matariya for Matariya virus (MTYV) found
in lesser whitethroats (sylviid Curruca curruca
(Linnaeus, 1758)) sampled in Port Saïd/Būrsaʿīd, Port
Saïd Governorate, Egypt [6].
Genus Vesiculovirus was expanded by two new species
(TaxoProp 2021.004M.R.Alpharhabdovirinae_13nsp):
• Vesiculovirus mediterranean for Mediterranean bat virus
(MBV) found in bats sampled in Algeria, Morocco, and
Spain (unpublished; GenBank: MW557331); and
• Vesiculovirus yinshui for Yinshui bat virus (YSBV) found
in Chinese rufous horseshoe bats (rhinolophid Rhinolo-
phus sinicus K. Andersen, 1905) sampled in Xiánníng
(咸宁市), Húběi Province (湖北省), China [39].
Family Sunviridae
The name of the only species in the family was changed
to a binomial as required by the recently amended ICVCN
[25, 69, 81] (2021.037M.R.Sunviridae_sprename).
Family Xinmoviridae
Family Xinmoviridae was thoroughly revised
(2021.039M.R.Xinmoviridae_11ngen_8nsp):
• in the genus Anphevirus, the species Shuangao anphevi-
rus was abolished, and the species Xincheng anphevirus
was renamed Anphevirus xinchengense. Species Dipteran
anphevirus, Odonate anphevirus, Drosophilid anphevi-

Taxonomic update of phylum Negarnaviricota
1 3
rus, Bolahun anphevirus, and Orthopteran anphevirus
were renamed Alasvirus muscae, Draselvirus dentati,
Drunivirus chambonense, Gambievirus bolahunense, and
Hoptevirus orthopteris, respectively, and moved into the
newly created genera Alasvirus, Draselvirus, Drunivirus,
Gambievirus, and Hoptevirus, respectively;
• genus Doupovirus was created to include the new species
Doupovirus australiaense for Culex mononega-like virus
2 (CMLV-2) discovered by HTS in mosquitoes (culicid
Culex australicus Dobrotworsky & Drummond, 1953)
sampled in Point Douro, Western Australia, Australia [57];
• genus Gambievirus was created to include the new spe-
cies Gambievirus senegalense for Gambie virus (GAMV)
discovered by HTS in mosquitoes (culicid Anopheles
gambiae Giles, 1902) sampled in Senegal [17];
• genus Gylbovirus was created to include new species
Gylbovirus aagae for Aedes anphevirus (AeAV) discov-
ered by HTS in yellow fever mosquitoes (culicid Aedes
aegypti (Linnaeus in Hasselquist, 1762)) and the Aag2
Aedes albopictus cell line [10, 48];
• genus Madalivirus was created to include two new spe-
cies:
o Madalivirus amapaense for Anopheles marajoara virus
(AnMV) discovered by HTS in mosquitoes (culicid
Anopheles marajoara Galvao & Damesceno 1942)
sampled in Macapá, Amapá State, Brazil [56]; and
o Madalivirus amazonaense for Anopheles darlingi
virus (AnDV) discovered by HTS in American
malaria mosquitoes (culicid Anopheles darlingi
Root, 1926) sampled in Macapá, Amapá State, and
Manaus, Amazonas State, Brazil [56];
• genus Pelmivirus was created to include new species
Pelmivirus eymattense for hymenopteran anphe-related
virus OKIAV71 (HARV-71) discovered by HTS in para-
sitoid wasps (ichneumonid Heteropelma amictum (Fab-
ricius, 1775)) sampled in Eymatt, Bern, Switzerland [27];
• genus Triniovirus was created to include the new species Trin-
iovirus yonagoense for Culex tritaeniorhynchus anphevirus
(CtAV) discovered by HTS in mosquitoes (culicid Culex tri-
taeniorhynchus Giles, 1901) sampled in Yonago (米子市),
Tottori Prefecture (鳥取県), Japan [16]; and
• genus Ulegvirus was created for new species Ulegvi-
rus freckenfeldense for odonatan anphe-related virus
OKIAV59 (OARV-59) discovered by HTS in golden-
ringed dragonflies (cordulegastrid Cordulegaster boltonii
(Donovan, 1807)) sampled in Freckenfeld, Rhineland-
Palatinate (Rheinland-Pfalz), Germany [27].
Yunchangviricetes
The names of all species in the class were changed to bino-
mials as required by the recently amended International
Code of Virus Classification and Nomenclature (ICVCN)
[25, 69, 81] (TaxoProp 2021.018M.R.Negarnaviricota
_sprename).
Taxonomic changes insubphylum
Polyploviricotina
Taxonomic changes withinorder Articulavirales
(Polyploviricotina: Insthoviricetes)
Family Amnoonviridae
The name of the only species in the family was
changed to a binomial as required by the recently
amended International Code of Virus Classification
and Nomenclature (ICVCN) [25, 69, 81] (TaxoProp
2021.018M.R.Negarnaviricota_sprename).
Family Orthomyxoviridae
The names of all species in the family were
changed to binomials as required by the recently
amended ICVCN [25, 69, 81] (TaxoProp
2021.024M.R.Orthomyxoviridae_sprename).
One new genus, Mykissvirus, was established to
include one new species, Mykissvirus tructae, for rain-
bow trout orthomyxovirus (RbtOV) and steelhead trout
orthomyxovirus (SttOV-1) first isolated from commer-
cially reared rainbow trout (salmonid Oncorhynchus
mykiss (Walbaum, 1792)) in Hagerman Valley, Idaho,
USA [4] (TaxoProp 2021.022M.R.Orthomyxoviridae_1
ngen_1nsp_Mykiss).
One new genus, Sardinovirus, was established to include
one new species, Sardinovirus pilchardi, for pilchard
orthomyxovirus (POMV) first isolated from blue pilchards
(clupeid Sardinops sagax (Jenyns, 1842)) sampled off the
coast of South Australia and subsequently Tasmania [19,
41] (TaxoProp 2021.023M.R.Orthomyxoviridae_1ngen_1
nsp_Sardino).
Genus Quaranjavirus was expanded by four new species
(TaxoProp 2021.034M.R.Quaranjavirus_4nsp):
• Quaranjavirus araguariense for Araguari virus (ARAV)
isolated from a gray four-eyed opossum (didelphid Phi-
lander opossum (Linnaeus, 1758)) sampled in Serra do
Navio, Amapá State, Brazil [9, 79];
• Quaranjavirus chadense for Lake Chad virus (LKCV)
isolated from a vitelline masked weaver (ploceid Ploceus

J. H. Kuhn etal.
1 3
vitellinus (Lichtenstein, 1823)), sampled at Lake Chad,
Nigeria [50];
• Quaranjavirus tyulekense for Tyulek virus (TLKV)
isolated from ticks (argasid Argas vulgaris Filippova,
1961) sampled near Tûlëk/Tölök (Tюлёк/Tөлөк), Naryn
Region (Hapынcкaя oблacть), Kirghiz Soviet Socialist
Republic, USSR [85]; and
• Quaranjavirus wellfleetense for Wellfleet Bay virus
(WFBV) isolated from common eiders (anatid Somate-
ria mollissima (Linnaeus, 1758)) sampled at Cape Cod,
Massachusetts, USA [1].
Genus Thogotovirus was expanded by six new species
(TaxoProp 2021.038M.R.Thogotovirus_6nsp):
• Thogotovirus bourbonense for Bourbon virus (BRBV)
isolated from a human sampled in Bourbon County, Kan-
sas, USA [30];
• Thogotovirus josense for Jos virus (JOSV) isolated from
a zebu (bovid Bos indicus Linnaeus, 1758) and ticks
(ixodid Amblyomma variegatum (Fabricius, 1794) and
Rhipicephalus decoloratus Koch, 1844)) sampled in Jos,
Plateau State, Nigeria [36];
• Thogotovirus ozense for Oz virus (OZV) isolated from
ticks (ixodid Amblyomma testudinarium Koch, 1844)
sampled in Ehime Prefecture (愛媛県), Japan [13];
• Thogotovirus sinuense for Sinu virus (SINUV) isolated
from a pool of mosquitoes collected in San Bernardo del
Viento, Córdoba Department, Colombia [8];
• Thogotovirus thailandense for Thailand tick thogotovirus
(TT-THOV) detected by HTS in ticks (ixodid Boophilus
sp.) sampled in Tha Wang Pha District (ท่าวังผา), Nan
Province (น่าน), Thailand [63]; and
• Thogotovirus upoluense for Upolu virus (UPOV) isolated
from ticks (ixodid Ornithodoros capensis Neumann,
1901) sampled on Upolu Cay, Great Barrier Reef, Aus-
tralia [12].
Taxonomic changes withinorder Bunyavirales
(Polyploviricotina: Ellioviricetes)
The order Bunyavirales was expanded by two new families
(2021.040M.R.Bunyavirales_2nfam_2ngen_6nsp):
• Discoviridae: genus Orthodiscovirus was created for five
species:
o Orthodiscovirus coniellae for Coniothyrium diplo-
diella negative-stranded RNA virus 1 (CdNSRV-1)
and Plasmopara viticola lesion associated mycobun-
yavirales-like virus 9, discovered by HTS in an isolate
of a fungus (melanconidaceaen Coniella diplodiella
(Speg.) Petr. & Syd., (1927)) and in plant lesions
caused by grapevine downy mildew (peronosporace-
aen Plasmopara viticola (Berk. & M.A. Curtis) Berl.
& De Toni, (1888)), respectively [42];
o Orthodiscovirus hispaniae for Plasmopara viticola
lesion associated mycobunyavirales-like virus 8
(PvLAM-LV8) discovered by HTS in plant lesions
caused by grapevine downy mildew (peronosporace-
aen Plasmopara viticola (Berk. & M.A. Curtis)
Berl. & De Toni, (1888)) [7];
o Orthodiscovirus iberiae for Plasmopara viticola
lesion associated mycobunyavirales-like virus 4
(PvLAM-LV4) discovered by HTS in plant lesions
caused by grapevine downy mildew (peronosporace-
aen Plasmopara viticola (Berk. & M.A. Curtis)
Berl. & De Toni, (1888)) [7];
o Orthodiscovirus missouriense for Penicillium disco-
virus (PDV) discovered by HTS in a fungus (tricho-
comaceaen Penicillium atramentosum Thom, C.
1910) [31]; and
o Orthodiscovirus penicillii for Penicillium roseopur-
pureum negative ssRNA virus 1 (PrNssV-1) discov-
ered by HTS in a fungus (trichocomaceaen Penicil-
lium roseopurpureum Dierckx, R.P. 1901) [42].
• Tulasviridae: genus Orthotulasvirus was created to
include one species, Orthotulasvirus tulasnellae, for
Tulasnella bunyavirales-like virus 1 (TB-LV) discovered
by HTS in an orchid mycorrhizal virome and confirmed
to be present in an isolate of an effused fungus (tulasnel-
laceaen Tulasnella sp.) by full length Sanger sequencing
of its cDNA [62].
Family Arenaviridae
Genus Mammarenavirus was expanded by two new
species:
• Mammarenavirus bituense for Bitu virus (BITV) detected
by HTS in Namaqua micaelamys (Micaelamys namaq-
uensis (A. Smith, 1834)) sampled in Angola (TaxoProp
2021.006M.R.Mammarenavirus_1nsp_Bitu) [64]; and
• Mammarenavirus kwanzaense, for Kwanza virus
(KWAV) detected by HTS in gray-bellied mice (Mus
triton (Thomas, 1909)) sampled in Angola (TaxoProp
2021.007M.R.Mammarenavirus_1nsp_Kwanza) [64].
Family Fimoviridae
The names of all species in the family were changed to
binomials as required by the recently amended ICVCN [25,
69, 81] (TaxoProp 2021.013P.R.Fimoviridae_rename).
Genus Emaravirus was expanded by three new species:

Taxonomic update of phylum Negarnaviricota
1 3
• Emaravirus aceris for maple mottle-associated emara-
virus (MaMaV) discovered by HTS in maple (sap-
indaceaen Acer sp.) sampled in Germany (TaxoProp
2021.010P.R.Emaravirus_1ns) [54];
• Emaravirus chrysanthemi for chrysanthemum
mosaic-associated emaravirus (ChMaV) discov-
ered by HTS in chrysanthemum (asteraceaen Chry-
santhemum morifolium L.) sampled in Japan
(TaxoProps 2021.011P.R.Emaravirus_1ns and
2021.043M.R.Corrections_Mononegavirales_Bunyavi-
rales) [32]; and
• Emaravirus quercus for common oak ringspot-associated
emaravirus (CORaV) discovered by HTS in oak (fagace-
aen Quercus robur L.) sampled in Germany and Scandi-
navia (TaxoProp 2021.012P.R.Emaravirus_1ns) [52].
Family Hantaviridae
The species name Seewis orhtohantavirus was
corrected to Seewis orthohantavirus (TaxoProp
2021.042M.R.Corrections_Riboviria_Duplodnaviria).
The species name Thottopalayam thottimvirus was
corrected to Thottapalayam thottimvirus (TaxoProp
2021.042M.R.Corrections_Riboviria_Duplodnaviria).
Family Leishbuviridae
The name of the only species in the fam-
ily was changed to a binomial as required by the
recently amended ICVCN [25, 69, 81] (TaxoProp
2021.018M.R.Negarnaviricota_sprename).
Table 1 ICTV-accepted
taxonomy of the order
Muvirales (Negarnaviricota:
Haploviricotina:
Chunqiuviricetes) as of March
2022
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, fami-
lies, and orders are taxa
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper
noun. This column lists the virus names with their correct (lack of) capitalization
Genus Species¶Virus (Abbreviation)&
Family Qinviridae
Yingvirus Yingvirus beihaiense Běihǎi sesarmid crab virus 4 (BhSCV-4)
Yingvirus charybdis Wēnzhōu qínvirus-like virus 2 (WzQLV-2)
Yingvirus hubeiense Húběi qínvirus-like virus 1 (HbQLV-1)
Yingvirus sanxiaense Sānxiá qínvirus-like virus 1 (SxQLV-1)
Yingvirus shaheense Shāhé qínvirus-like virus 1 (ShQLV-1)
Yingvirus wenzhouense Wēnzhōu qínvirus-like virus 1 (WzQLV-1)
Yingvirus wuhanense Wǔhàn insect virus 15 (WhIV-15)
Yingvirus xinzhouense Xīnzhōu nematode virus 3 (XzNV-3)
Table 2 ICTV-accepted
taxonomy of the
order Serpentovirales
(Negarnaviricota:
Haploviricotina: Milneviricetes)
as of March 2022
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, fami-
lies, and orders are taxa
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper
noun. This column lists the virus names with their correct (lack of) capitalization
Genus Species¶Virus (Abbreviation)&
Family Aspiviridae
Ophiovirus Ophiovirus citri citrus psorosis virus (CPsV)
Ophiovirus freesiae freesia sneak virus (FreSV)
Ophiovirus lactucae lettuce ring necrosis virus (LRNV)
Ophiovirus mirafioriense Mirafiori lettuce big-vein virus (MLBVV)
Ophiovirus ranunculi ranunculus white mottle virus (RWMV)
Ophiovirus tulipae tulip mild mottle mosaic virus (TMMMV)
Ophiovirus vaccinii blueberry mosaic associated virus (BlMaV)

J. H. Kuhn etal.
1 3
Table 3 ICTV-accepted taxonomy of the order Jingchuvirales (Negarnaviricota: Haploviricotina: Monjiviricetes) as of March 2022
Genus Species¶Virus (Abbreviation)&
Family Aliusviridae
Obscuruvirus Obscuruvirus quintum Atrato chu-like virus 5 (AClV-5)
Ollusvirus Ollusvirus coleopteri Húběi coleoptera virus 3 (HbCV-3)
Ollusvirus culvertonense Culverton virus (CvV)
Ollusvirus hanchengense Hánchéng leafhopper mivirus (HLMV)
Ollusvirus hymenopteri hymenopteran chu-related virus 123 (HCrV-123)
Ollusvirus insectii hymenopteran chu-related virus 126 (HCrV-126)
Ollusvirus scaldisense Scaldis River bee virus (SRBV)
Ollusvirus shayangense Shāyáng fly virus 1 (SyFV-1)
Ollusvirus taiyuanense Tàiyuán leafhopper virus (TYLeV)
Family Chuviridae
Boscovirus Boscovirus hippoboscidae Wǔhàn louse fly virus 7 (WhLFV-7)
Boscovirus hypoboscidae Wǔhàn louse fly virus 6 (WhLFV-6)
Chuvivirus Chuvivirus brunnichi Wēnlǐng crustacean virus 14 (WlCV-14)
Chuvivirus canceris Wēnzhōu crab virus 2 (WzCV-2)
Culicidavirus Culicidavirus culicidae Wǔhàn mosquito virus 8 (WhMV-8)
Culicidavirus culicis Culex mosquito virus 5 (ClMV-5)
Culicidavirus imjinense Imjin River virus 1 (IjRV-1)
Culicidavirus quitotaense Culex mosquito virus 4 (ClMV-4)
Demapteravirus Demapteravirus dermapteri dermapteran chu-related virus 142 (DCrV-142)
Doliuvirus Doliuvirus culisetae Mos8Chu0 chuvirus (MoCV)
Mivirus Mivirus amblyommae lone star tick chuvirus 1 (LSTCV-1)
Mivirus boleense Bólè tick virus 3 (BTV-3)
Mivirus changpingense Chāngpíng tick virus 2 (CpTV-2)
Mivirus dermacentoris Chāngpíng tick virus 3 (CpTV-3)
Tǎchéng tick virus 5 (TcTV-5)
Mivirus genovaense Genoa virus (GeV)
Mivirus karukeraense Karukera tick virus (KtV)
Mivirus rhipicephali brown dog tick mivirus 1 (BDTMV-1)
Mivirus suffolkense Suffolk virus (SFKV)
Mivirus wuhanense Wǔhàn tick virus 2 (WhTV-2)
Morsusvirus Morsusvirus argatis Tǎchéng tick virus 4 (TcTV-4)
Nigecruvirus Nigecruvirus ixodes blacklegged tick chuvirus-2 (BlTC-2)
Odonatavirus Odonatavirus draconis odonatan chu-related virus 137 (OCrV-137)
Odonatavirus fabricii Húběi odonate virus 11 (HbOV-11)
Odonatavirus odontis odonatan chu-related virus 136 (OCrV-136)
Pediavirus Pediavirus cirripedis Běihǎi barnacle virus 9 (BhBV-9)
Piscichuvirus Piscichuvirus franki Herr Frank virus 1 (HFrV-1)
Piscichuvirus lycodontis Guǎngdōng red-banded snake chuvirus-like virus (GRSCV)
Piscichuvirus sanxiaense Sānxiá atyid shrimp virus 4 (SxASC-4)
Piscichuvirus wenlingense Wēnlǐng fish chu-like virus (WFClV)
Pterovirus Pterovirus chulinense hymenopteran chu-related virus OKIAV147 (HCrV-147)
Scarabeuvirus Scarabeuvirus blattae Wǔchāng cockroach virus 3 (WcLFV-3)
Scarabeuvirus dentati Húběi chuvirus-like virus 3 (HbCLV-3)
Scarabeuvirus hubeiense Húběi chuvirus-like virus 1 (HbCLV-1)
Scarabeuvirus lampyris Lampyris noctiluca chuvirus-like virus 1 (LNClV-1)
Scarabeuvirus lishiense Lĭshì spider virus 1 (LsSV-1)
Taceavirus Taceavirus wenlingense Wēnlǐng crustacean virus 15 (WlCV-15)

Taxonomic update of phylum Negarnaviricota
1 3
Family Mypoviridae
The name of the only species in the fam-
ily was changed to a binomial as required by the
recently amended ICVCN [25, 69, 81] (TaxoProp
2021.018M.R.Negarnaviricota_sprename).
Family Peribunyaviridae
Family Peribunyaviridae was expanded by three new
genera (TaxoProp 2021.027M.R.Peribunyaviridae_3ngen_
3nsp):
• Khurdivirus, including one new species, Khurdivirus
volgaense, for Khurdun virus (KHURV) isolated from a
Eurasian coot (rallid Fulica atra Linnaeus, 1758) sam-
pled in Astrakhan Region (Acтpaxaнcкaя oблacть), Rus-
sia [83, 84];
• Lakivirus, including one new species, Lakivirus lakam-
haense, for Lakamha virus (LAKV) discovered by HTS
in a mosquito (culicid Wyeomyia complosa (Dyar, 1928))
sampled in Palenque, Chiapas, Mexico [29]; and
• Lambavirus, including one new species, Lambavirus wis-
consinense, for largemouth bass bunyavirus (LBBV) dis-
covered by HTS in largemouth bass (centrarchid Microp-
terus salmoides (Lacépède, 1802)) sampled in the upper
Mississippi River, Wisconsin, USA [71].
The species name Lumbo orthbunyavirus was
corrected to Lumbo orthobunyavirus (TaxoProp
2021.042M.R.Corrections_Riboviria_Duplodnaviria).
Family Phasmaviridae
The family was expanded by one new genus, Cicadellivi-
rus, to include one new species, Cicadellivirus scaphoidei,
for Scaphoideus titanus bunya-like virus 1 (StHV) (Taxo-
Prop 2021.030M.R.Phasmaviridae_1ngen_5nsp) [46].
Genus Orthophasmavirus was expand ed by four new species
(TaxoProps 2021.030M.R.Phasmaviridae_1ngen_5nsp and
2021.043M.R.Corrections_Mononegavirales_Bunyavirales):
• Orthophasmavirus coredoense for Coredo virus (CORV)
discovered by HTS in mosquitoes (culicid Mansonia titil-
lans (Walker, 1848)) sampled in Colombia (unpublished;
GenBank: MN661021–3);
• Orthophasmavirus barstukasense for Barstukas virus
(BARV) discovered by HTS in mosquitoes (culicid Aedes
sp.) sampled in California, USA [3];
• Orthophasmavirus miglotasense for Miglotas virus
(MIGV) discovered by HTS in mosquitoes (culicid Aedes
sp. and Culex sp.) sampled in California, USA [3]; and
• Orthophasmavirus flenense for Flen virus (FLNV) dis-
covered by HTS in mosquitoes (culicid Aedes cantans
(Meigen, 1818)) sampled in Flen, Södermanland County
(Södermanlands län), Sweden [43].
Family Wupedeviridae
The name of the only species in the fam-
ily was changed to a binomial as required by the
recently amended ICVCN [25, 69, 81] (TaxoProp
2021.018M.R.Negarnaviricota_sprename).
Summary
A summary of the current, ICTV-accepted taxonomy of the
phylum Negarnaviricota is presented in Tables1, 2, 3, 4,
5, 6 and 7 and in the form of two posters (Supplementary
Figure1).
Table 3 (continued)
Genus Species¶Virus (Abbreviation)&
Family Crepuscuviridae
Aqualaruvirus Aqualaruvirus sialis megalopteran chu-related virus 119 (MCrV-119)
Family Myriaviridae
Myriavirus Myriavirus myriapedis Húběi myriapoda virus 8 (HbMV-8)
Family Natareviridae
Charybdivirus Charybdivirus charybdis Wēnzhōu crab virus 3 (WzCV-3)
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, families, and orders are taxa.
¶ Taxon names are always italicized and always begin with a capital letter.
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper noun. This column lists the virus
names with their correct (lack of) capitalization. Lists of viruses within a given species are provisional at this point and will likely be amended in
the near future.

J. H. Kuhn etal.
1 3
Table 4 ICTV-accepted taxonomy of the order Mononegavirales (Negarnaviricota: Haploviricotina: Monjiviricetes) as of March 2022
Genus Species¶Virus (Abbreviation)&
Family Artoviridae
Hexartovirus Hexartovirus cirripedis Běihai barnacle virus 8 (BhBV-8)
Hexartovirus lepeophtheiri Lepeophtheirus salmonis negative-stranded RNA virus 1 (LsNSRV-1)
Peropuvirus Peropuvirus beihaiense Běihai rhabdo-like virus 1 (BhRLV-1)
Peropuvirus dentati Húběi rhabdo-like virus 8 (HbRLV-8)
Peropuvirus hubeiense Húběi rhabdo-like virus 6 (HbRLV-6)
Peropuvirus juli Húběi rhabdo-like virus 5 (HbRLV-5)
Peropuvirus lignarii Běihai rhabdo-like virus 2 (BhRLV-2)
Peropuvirus melongenae Solanum melongena rhabdo-like virus (SmRLV)
Peropuvirus pteromali Pteromalus puparum negative-strand RNA virus 1 (PpNSRV-1)
Family Bornaviridae
Carbovirus Carbovirus queenslandense jungle carpet python virus (JCPV)
Carbovirus tapeti southwest carpet python virus (SWCPV)
Cultervirus Cultervirus hemicultri Wǔhàn sharpbelly bornavirus (WhSBV)
Orthobornavirus Orthobornavirus alphapsittaciforme 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)
Orthobornavirus avisaquaticae aquatic bird bornavirus 1 (ABBV-1)
aquatic bird bornavirus 2 (ABBV-2)
Orthobornavirus betapsittaciforme parrot bornavirus 5 (PaBV-5)
Orthobornavirus bornaense Borna disease virus 1 (BoDV-1)
Borna disease virus 2 (BoDV-2)
Orthobornavirus caenophidiae Caribbean watersnake bornavirus (CWBV)
Mexican black-tailed rattlesnake bornavirus (MRBV)
Orthobornavirus elapsoideae Loveridge’s garter snake virus 1 (LGSV-1)
Orthobornavirus estrildidae estrildid finch bornavirus 1 (EsBV-1)
Orthobornavirus sciuri variegated squirrel bornavirus 1 (VSBV-1)
Orthobornavirus serini canary bornavirus 1 (CnBV-1)
canary bornavirus 2 (CnBV-2)
canary bornavirus 3 (CnBV-3)
Family Filoviridae
Cuevavirus Lloviu cuevavirus Lloviu virus (LLOV)
Dianlovirus Mengla dianlovirus Měnglà virus (MLAV)
Ebolavirus Bombali ebolavirus Bombali virus (BOMV)
Bundibugyo ebolavirus Bundibugyo virus (BDBV)
Reston ebolavirus Reston virus (RESTV)
Sudan ebolavirus Sudan virus (SUDV)
Tai Forest ebolavirus Taï Forest virus (TAFV)
Zaire ebolavirus Ebola virus (EBOV)
Marburgvirus Marburg marburgvirus Marburg virus (MARV)
Ravn virus (RAVV)
Oblavirus Oblavirus percae Oberland virus (OBLV)
Striavirus Xilang striavirus Xīlǎng virus (XILV)
Tapjovirus Tapjovirus bothropis Tapajós virus (TAPV)
Thamnovirus Huangjiao thamnovirus Huángjiāo virus (HUJV)
Thamnovirus kanderense Kander virus (KNDV)
Thamnovirus percae Fiwi virus (FIWIV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Family Lispiviridae
Anicalvirus Anicalvirus hangzhouense Anisopteromalus calandrae negative-strand RNA virus 2 (AcNSRV-2)
Anidravirus Anidravirus hangzhouense Anisopteromalus calandrae negative-strand RNA virus 1 (AcNSRV-1)
Arlivirus Arlivirus arachnae Lĭshì spider virus 2 (LsSV-2)
Copasivirus Copasivirus ivindoense isopteran arli-related virus OKIAV103 (IARV-103)
Copasivirus manlyvaleense Jimsystermes virus (JIMV)
Cybitervirus Cybitervirus niederense coleopteran arli-related virus OKIAV107 (CARV-107)
Damravirus Damravirus dentatis Húběi odonate virus 10 (HbOV-10)
Ganiavirus Ganiavirus tachengense Tǎchéng tick virus 6 (TcTV-6)
Leocovirus Leocovirus coleopteris Húběi rhabdo-like virus 3 (HbRLV-3)
Nematovirus Nematovirus wuchangense Wuchāng romanomermis nematode virus 2 (WcRNV-2)
Phelinovirus Phelinovirus aphidis hymenopteran arli-related virus OKIAV99 (HARV-99)
Rivapovirus Rivapovirus aleyrodidae hemipteran arli-related virus OKIAV94 (HARV-94)
Sanstrivirus Sanstrivirus gerridis Sānxiá water strider virus 4 (SxWSV-4)
Stylovirus Stylovirus niederense strepsipteran arli-related virus OKIAV104 (SARV-104)
Supelovirus Supelovirus thailandense blattodean arli-related virus OKIAV102 (BARV-102)
Synelinevirus Synelinevirus bonnense hymenopteran arli-related virus OKIAV98 (HARV-98)
Synelinevirus paranaense Linepithema humile rhabdo-like virus 1 (LhuRLV-1)
Usmuvirus Usmuvirus newyorkense Amsterdam virus (AMSV)
Xenophyvirus Xenophyvirus mathesonense hemipteran arli-related virus OKIAV95 (HARV-95)
Family Mymonaviridae
Auricularimonavirus Auricularimonavirus auriculariae Auricularia heimuer negative-stranded RNA virus 1 (AhNRSV-1)
Auricularimonavirus bondarzewiae Bondarzewia berkeleyi negative-strand RNA virus 1 (BbNSRV-1)
Botrytimonavirus Botrytimonavirus alphabotrytidis Botrytis cinerea negative-stranded RNA virus 5 (BcNSRV-5)
Sclerotinia sclerotiorum negative-stranded RNA virus 10 (SsNSRV-10)
Botrytimonavirus botrytidis Botrytis cinerea negative-stranded RNA virus 7 (BcNSRV-7)
Sclerotinia sclerotiorum negative-stranded RNA virus 11 (SsNSRV-11)
Botrytimonavirus glycinis soybean leaf-associated negative-stranded RNA virus 3 (SLaNSRV-3)
Botrytimonavirus sclerotiniae Sclerotinia sclerotiorum negative-stranded RNA virus 2 (SsNSRV-2)
Sclerotinia sclerotiorum negative-stranded RNA virus 2-A (SsNSRV-2-A)
Sclerotinia sclerotiorum negative-stranded RNA virus 4 (SsNSRV-4)
Sclerotinia sclerotiorum negative-stranded RNA virus 4-A (SsNSRV-4-A)
Hubramonavirus Hubramonavirus hubeiense Húběi rhabdo-like virus 4 (HbRLV-4)
Hubramonavirus terrae H2BulkLitter1223 virus
Lentimonavirus Lentimonavirus lentinulae Lentinula edodes negative-strand RNA virus 1 (LeNSRV-1)
Penicillimonavirus Penicillimonavirus alphapenicillii Penicillium adametzioides negative-stranded RNA virus 1 (PaNsRV-1)
Penicillimonavirus alphaplasmoparae Plasmopara viticola lesion associated mononegaambi virus 1 (PvaMV-1)
Penicillimonavirus betapenicillii Penicillium glabrum negative-stranded RNA virus 1 (PgRlV-1)
Penicillimonavirus betaplasmoparae Plasmopara viticola lesion associated mononegaambi virus 2 (PvaMV-2)
Plasmopara viticola lesion associated mononegaambi virus 4 (PvaMV-4)
Penicillimonavirus deltaplasmoparae Plasmopara viticola lesion associated mononegaambi virus 5 (PvaMV-5)
Penicillimonavirus epsilonplasmoparae Plasmopara viticola lesion associated mononegaambi virus 6 (PvaMV-6)
Penicillimonavirus etaplasmoparae Plasmopara viticola lesion associated mononegaambi virus 9 (PvaMV-9)
Penicillimonavirus gammaplasmopara Plasmopara viticola lesion associated mononegaambi virus 3 (PvaMV-3)
Penicillimonavirus kilnbarnense Kiln Barn virus (KBV)
Penicillimonavirus zetaplasmoparae Plasmopara viticola lesion associated mononegaambi virus 7 (PvaMV-7)
Plasmopamonavirus Plasmopamonavirus plasmoparae Plasmopara viticola lesion associated mononegaambi virus 8 (PvaMV-8)
Phyllomonavirus Phyllomonavirus gysingense Gysinge virus (GYSV)
Phyllomonavirus phyllospherae soybean leaf-associated negative-stranded RNA virus 4 (SLaNSRV-4)

J. H. Kuhn etal.
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Rhizomonavirus Rhizomonavirus mali apple virus B (APPv-B)
Sclerotimonavirus Sclerotimonavirus alphabotrytidis Botrytis cinerea negative-stranded RNA virus 3 (BcNSRV-3)
Sclerotinia sclerotiorum negative-stranded RNA virus 9 (SsNSRV-9)
Sclerotimonavirus alphaclarireediae Sclerotinia homoeocarpa TSA contig 1 (ShTSA-1)
Sclerotinia homoeocarpa TSA contig 2 (ShTSA-2)
Sclerotimonavirus alphaplasmoparae Plasmopara viticola lesion associated mymonavirus 1 (PvaMV-1)
Sclerotimonavirus alternariae Alternaria tenuissima negative-stranded RNA virus 1 (AtNsRV-1)
Sclerotimonavirus betabotrytidis Botrytis cinerea negative-stranded RNA virus 4 (BcNSRV-4)
Sclerotimonavirus betaclarireediae Sclerotinia homoeocarpa TSA contig 3 (ShTSA-3)
Sclerotimonavirus betaplasmoparae Plasmopara viticola lesion associated mononega virus 2 (PvaMV-2)
Sclerotimonavirus botrytidis Botrytis cinerea mymonavirus 1 (BcMyV-1)
Sclerotinia sclerotiorum negative-stranded RNA virus 7 (SsNSRV-7)
Sclerotimonavirus cryphonectriae Cryphonectria parasitica sclerotimonavirus 1 (CpSV-1)
Sclerotimonavirus fusarii Fusarium graminearum negative-stranded RNA virus 1 (FgNSRV-1)
soybean leaf-associated negative-stranded RNA virus 1 (SLaNSRV-1)
Sclerotimonavirus illinoisense soybean leaf-associated negative-stranded RNA virus 2 (SLaNSRV-2)
Sclerotimonavirus penicillii Penicillium cairnsense negative-stranded RNA virus 1 (PcNsRV-1)
Sclerotimonavirus sclerotiniae Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1)
Sclerotinia sclerotiorum negative-stranded RNA virus 1-A (SsNSRV-1-A)
Sclerotinia sclerotiorum negative-stranded RNA virus 3 (SsNSRV-3)
Sclerotinia sclerotiorum negative-stranded RNA virus 3-A (SsNSRV-3-A)
Sclerotimonavirus terrae H4BulkLitter234 virus
Family Nyamiviridae
Berhavirus Berhavirus beihaiense Běihai rhabdo-like virus 4 (BhRLV-4)
Berhavirus radialis Běihai rhabdo-like virus 5 (BhRLV-5)
Berhavirus sipunculi Běihai rhabdo-like virus 3 (BhRLV-3)
Crustavirus Crustavirus beihaiense Běihai rhabdo-like virus 6 (BhRLV-6)
Crustavirus wenlingense Wēnling crustacean virus 12 (WlCV-12)
Crustavirus wenzhouense Wēnzhōu crab virus 1 (WzCV-1)
Formivirus Formivirus angliae Formica fusca virus 1 (FfusV-1)
Formivirus chalybii hymenopteran orino-related virus OKIAV87 (HORV-87)
Formivirus finnoniae Formica exsecta virus 4 (FeV-4)
Formivirus gorytis hymenopteran orino-related virus OKIAV85 (HORV-85)
Formivirus solenopsi Solenopsis invicta virus 15 (SoINV-15)
Nyavirus Nyavirus argatis Sekira virus (SEKRV)
Nyavirus midwayense Midway virus (MIDWV)
Nyavirus nyamaniniense Nyamanini virus (NYMV)
Nyavirus sanjacintoense San Jacinto virus (SJCV)
Nyavirus sierranevadaense Sierra Nevada virus (SNVV)
Nyavirus somateriae Jeremy Point nyavirus (JPNV)
Orinovirus Orinovirus pasiphilae Orinoco virus (ONCV)
Socyvirus Socyvirus heteroderae soybean cyst nematode virus 1 (SbCNV-1)
Tapwovirus Tapwovirus cesti Wēnzhōu tapeworm virus 1 (WzTWV-1)
Family Paramyxoviridae
Subfamily Avulavirinae
Metaavulavirus Avian metaavulavirus 2 avian paramyxovirus 2 (APMV-2)
Avian metaavulavirus 5 avian paramyxovirus 5 (APMV-5)
Avian metaavulavirus 6 avian paramyxovirus 6 (APMV-6)
Avian metaavulavirus 7 avian paramyxovirus 7 (APMV-7)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Avian metaavulavirus 8 avian paramyxovirus 8 (APMV-8)
Avian metaavulavirus 10 avian paramyxovirus 10 (APMV-10)
Avian metaavulavirus 11 avian paramyxovirus 11 (APMV-11)
Avian metaavulavirus 14 avian paramyxovirus 14 (APMV-14)
Avian metaavulavirus 15 avian paramyxovirus 15 (APMV-15)
Avian metaavulavirus 20 avian paramyxovirus 20 (APMV-20)
Avian metaavulavirus 22 avian paramyxovirus 22 (APMV-22)
Orthoavulavirus Avian orthoavulavirus 1 avian paramyxovirus 1 (APMV-1)1
Avian orthoavulavirus 9 avian paramyxovirus 9 (APMV-9)
Avian orthoavulavirus 12 avian paramyxovirus 12 (APMV-12)
Avian orthoavulavirus 13 avian paramyxovirus 13 (APMV-13)
Avian orthoavulavirus 16 avian paramyxovirus 16 (APMV-16)
Avian orthoavulavirus 17 Antarctic penguin virus A (APV-A)
Avian orthoavulavirus 18 Antarctic penguin virus B (APV-B)
Avian orthoavulavirus 19 Antarctic penguin virus C (APV-C)
Avian orthoavulavirus 21 avian paramyxovirus 21 (APMV-21)
Paraavulavirus Avian paraavulavirus 3 avian paramyxovirus 3 (APMV-3)
Avian paraavulavirus 4 avian paramyxovirus 4 (APMV-4)
Subfamily Metaparamyxovirinae
Synodonvirus Synodus synodonvirus Wēnlǐng triplecross lizardfish paramyxovirus (WTLPV)
Subfamily Orthoparamyxovirinae
Aquaparamyxovirus Oncorhynchus aquaparamyxovirus Pacific salmon paramyxovirus (PSPV)
Salmo aquaparamyxovirus Atlantic salmon paramyxovirus (AsaPV)
Ferlavirus Reptilian ferlavirus fer-de-lance virus (FDLV)
Henipavirus Cedar henipavirus Cedar virus (CedV)
Ghanaian bat henipavirus Ghana virus (GhV)
Hendra henipavirus Hendra virus (HeV)
Mojiang henipavirus Mòjiāng virus (MojV)
Nipah henipavirus Nipah virus (NiV)
Jeilongvirus Beilong jeilongvirus Beilong virus (BeiV)
Jeilongvirus anhuiense Miniopterus schreibersii paramyxovirus (MisPV)
Jeilongvirus apodemi rodent paramyxovirus (RoPV)
Jeilongvirus comorosense bat paramyxovirus 16797 (BatPV-1)
Jeilongvirus erinacei belerina virus (BeV)
Jeilongvirus felis feline paramyxovirus (FPaV)
Jeilongvirus madagascarense bat paramyxovirus 17770 (BatPV-2)
Jeilongvirus murinae bat paramyxovirus (BatPV-3)
Jeilongvirus rungweense ruloma virus (RulV)
Jun jeilongvirus J virus (JV)
Lophuromys jeilongvirus 1 Mount Mabu Lophuromys virus 1 (MMLV-1)
Lophuromys jeilongvirus 2 Mount Mabu Lophuromys virus 2 (MMLV-2)
Miniopteran jeilongvirus Shaan virus (ShaV)
Myodes jeilongvirus Pohorje Myodes paramyxovirus 1 (PMPV-1)
Tailam jeilongvirus Tailam virus (TaiV)
Morbillivirus Canine morbillivirus canine distemper virus (CDV)
Cetacean morbillivirus cetacean morbillivirus (CeMV)
Feline morbillivirus feline morbillivirus (FeMV)
Measles morbillivirus measles virus (MeV)
Phocine morbillivirus phocine distemper virus (PDV)

J. H. Kuhn etal.
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Rinderpest morbillivirus rinderpest virus (RPV)
Small ruminant morbillivirus peste-des-petits-ruminants virus (PPRV)
Narmovirus Mossman narmovirus Mossman virus (MossV)
Myodes narmovirus bank vole virus 1 (BaV-1)
Nariva narmovirus Nariva virus (NarV)
Tupaia narmovirus Tupaia paramyxovirus (TupV)
Respirovirus Bovine respirovirus 3 bovine parainfluenza virus 3 (BPIV-3)
Caprine respirovirus 3 caprine parainfluenzavirus 3 (CPIV-3)
Human respirovirus 1 human parainfluenza virus 1 (HPIV-1)
Human respirovirus 3 human parainfluenza virus 3 (HPIV-3)
Murine respirovirus Sendai virus (SeV)
Porcine respirovirus 1 porcine parainfluenza virus 1 (PPIV-1)
Squirrel respirovirus giant squirrel virus (GSqV)
Salemvirus Salem salemvirus Salem virus (SalV)
Subfamily Rubulavirinae
Orthorubulavirus Human orthorubulavirus 2 human parainfluenza virus 2 (HPIV-2)
Human orthorubulavirus 4 human parainfluenza virus 4a (HPIV-4a)
human parainfluenza virus 4b (HPIV-4b)
Mammalian orthorubulavirus 5 parainfluenza virus 5 (PIV-5)
Mammalian orthorubulavirus 6 Alston virus (AlsV)
Mapuera orthorubulavirus Mapuera virus (MapV)
Mumps orthorubulavirus mumps virus (MuV)
Porcine orthorubulavirus La Piedad Michoacán Mexico virus (LPMV)
Simian orthorubulavirus simian virus 41 (SV-41)
Pararubulavirus Achimota pararubulavirus 1 Achimota virus 1 (AchPV-1)
Achimota pararubulavirus 2 Achimota virus 2 (AchPV-2)
Hervey pararubulavirus Hervey virus (HerV)
Menangle pararubulavirus Menangle virus (MenPV)
Sosuga pararubulavirus Sosuga virus (SOSV)
Teviot pararubulavirus Teviot virus (TevPV)
Tioman pararubulavirus Tioman virus (TioPV)
Tuhoko pararubulavirus 1 Tuhoko virus 1 (ThkPV-1)
Tuhoko pararubulavirus 2 Tuhoko virus 2 (ThkPV-2)
Tuhoko pararubulavirus 3 Tuhoko virus 3 (ThkPV-3)
Unassigned (to subfamilies)
Cynoglossusvirus Cynoglossus cynoglossusvirus Wēnlǐng tonguesole paramyxovirus (WTSPV)
Hoplichthysvirus Hoplichthys hoplichthysvirus Wēnlǐng hoplichthys paramyxovirus (WHPV)
Scoliodonvirus Scoliodon scoliodonvirus Wēnzhōu pacific spadenose shark paramyxovirus (WPSSPV)
Family Pneumoviridae
Metapneumovirus Avian metapneumovirus avian metapneumovirus (AMPV)
Human metapneumovirus human metapneumovirus (HMPV)
Orthopneumovirus Bovine orthopneumovirus bovine respiratory syncytial virus (BRSV)
Human orthopneumovirus human respiratory syncytial virus (HRSV)
Murine orthopneumovirus murine pneumonia virus (MPV)
Family Rhabdoviridae
Subfamily Alpharhabdovirinae
Almendravirus Almendravirus almendras Puerto Almendras virus (PTAMV)
Almendravirus arboretum Arboretum virus (ABTV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Almendravirus balsa Balsa virus (BALV)
Almendravirus chico Rio Chico virus (RCHV)
Almendravirus cootbay Coot Bay virus (CBV)
Almendravirus menghai Menghai virus (MRV)
Alphanemrhavirus Alphanemrhavirus xingshan Xingshan nematode virus 4 (XsNV-4)
Alphanemrhavirus xinzhou Xinzhou nematode virus 4 (XzNV-4)
Alphapaprhavirus Alphapaprhavirus hubei Hubei lepidoptera virus 2 (HbLV-2)
Alphapaprhavirus pararge Pararge aegeria rhabdovirus (PAeRV)
Alpharicinrhavirus Alpharicinrhavirus blanchseco Blanchseco virus (BCOV)
Alpharicinrhavirus bole Bole tick virus 2 (BlTV-2)
Alpharicinrhavirus wuhan Wuhan tick virus 1 (WhTV-1)
Amplylivirus Amplylivirus cinereus frog lyssa-like virus 1 (FLLV-1)
Arurhavirus Arurhavirus aruac Aruac virus (ARUV)
Arurhavirus inhangapi Inhangapi virus (INHV)
Arurhavirus santabarbara Santa Barbara virus (SBAV)
Arurhavirus xiburema Xiburema virus (XIBV)
Barhavirus Barhavirus bahia Bahia Grande virus (BGV)
Harlingen virus (HARV)
Barhavirus muir Muir Springs virus (MSV)
Caligrhavirus Caligrhavirus caligus Caligus rogercresseyi rhabdovirus (CRogRV)
Caligrhavirus lepeophtheirus Lepeophtheirus salmonis rhabdovirus 127 (LSalRV-127)
Caligrhavirus salmonlouse Lepeophtheirus salmonis rhabdovirus 9 (LSalRV-9)
Cetarhavirus Cetarhavirus laganorhynchus dolphin rhabdovirus (DRV)
Cetarhavirus phocoena harbour porpoise rhabdovirus (HPRV)
Curiovirus Curiovirus curionopolis Curionopolis virus (CURV)
Curiovirus iriri Iriri virus (IRIRV)
Curiovirus itacaiunas Itacaiunas virus (ITAV)
Curiovirus rochambeau Rochambeau virus (RBUV)
Ephemerovirus Ephemerovirus adelaide Adelaide River virus (ARV)
Ephemerovirus berrimah Berrimah virus (BRMV)
Ephemerovirus febris bovine ephemeral fever virus (BEFV)
Ephemerovirus hayes Hayes Yard virus (HYV)
Ephemerovirus kent New Kent County virus (NKCV)
Ephemerovirus kimberley Kimberley virus (KIMV)
Malakal virus (MALV)
Ephemerovirus koolpinyah Koolpinyah virus (KOOLV)
Ephemerovirus kotonkan kotonkan virus (KOTV)
Ephemerovirus obodhiang Obodhiang virus (OBOV)
Ephemerovirus puchong Puchong virus (PUCV)
Ephemerovirus yata Yata virus (YATV)
Hapavirus Hapavirus bangoran Bangoran virus (BGNV)
Hapavirus flanders Flanders virus (FLAV)
Hapavirus graylodge Gray Lodge virus (GLOV)
Hapavirus hartpark Hart Park virus (HPV)
Hapavirus holmes Holmes Jungle virus (HOJV)
Hapavirus joinjakaka Joinjakaka virus (JOIV)
Hapavirus kamese Kamese virus (KAMV)
Hapavirus lajoya La Joya virus (LJV)
Hapavirus landjia Landjia virus (LANV = LJAV)
Hapavirus manitoba Manitoba virus (MANV = MNTBV)

J. H. Kuhn etal.
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Hapavirus marco Marco virus (MCOV)
Hapavirus mosqueiro Mosqueiro virus (MQOV)
Hapavirus mossuril Mossuril virus (MOSV)
Hapavirus ngaingan Ngaingan virus (NGAV)
Hapavirus ord Ord River virus (ORV)
Hapavirus parry Parry Creek virus (PCV)
Hapavirus porton Porton virus (PORV)
Hapavirus wongabel Wongabel virus (WONV)
Ledantevirus Ledantevirus barur Barur virus (BARV)
Ledantevirus bughendera Bughendera virus (BUGV)
Ledantevirus elgon Mount Elgon bat virus (MEBV)
Ledantevirus fikirini Fikirini virus (FKRV)
Ledantevirus fukuoka Fukuoka virus (FUKV)
Ledantevirus kanyawara Kanyawara virus (KYAV)
Ledantevirus kern Kern Canyon virus (KCV)
Ledantevirus keuraliba Keuraliba virus (KEUV)
Ledantevirus kolente Kolente virus (KOLEV)
Ledantevirus kumasi Kumasi rhabdovirus (KRV)
Ledantevirus ledantec Le Dantec virus (LDV)
Ledantevirus nishimuro Nishimuro virus (NISV)
Ledantevirus nkolbisson Nkolbisson virus (NKOV)
Ledantevirus oita Oita virus (OITAV)
Ledantevirus taiyi Taiyi bat virus (TYBV)
Ledantevirus vaprio Vaprio virus (VAPV)
Ledantevirus wuhan Wuhan louse fly virus 5 (WLFV-5)
Ledantevirus yongjia Yongjia tick virus 2 (YTV-2)
Lostrhavirus Lostrhavirus hyalomma Xinjiang tick rhabdovirus (XjTRV)
Lostrhavirus lonestar lone star tick rhabdovirus (LITRV)
Lyssavirus Lyssavirus aravan Aravan virus (ARAV)
Lyssavirus australis Australian bat lyssavirus (ABLV)
Lyssavirus bokeloh Bokeloh bat lyssavirus (BBLV)
Lyssavirus caucasicus West Caucasian bat virus (WCBV)
Lyssavirus duvenhage Duvenhage virus (DUVV)
Lyssavirus formosa Taiwan bat lyssavirus (TWBLV)
Lyssavirus gannoruwa Gannoruwa bat lyssavirus (GBLV)
Lyssavirus hamburg European bat lyssavirus 1 (EBLV-1)
Lyssavirus helsinki European bat lyssavirus 2 (EBLV-2)
Lyssavirus ikoma Ikoma lyssavirus (IKOV)
Lyssavirus irkut Irkut virus (IRKV)
Lyssavirus khujand Khujand virus (KHUV)
Lyssavirus lagos Lagos bat virus (LBV)
Lyssavirus lleida Lleida bat lyssavirus (LLEBV)
Lyssavirus mokola Mokola virus (MOKV)
Lyssavirus rabies rabies virus (RABV)
Lyssavirus shimoni Shimoni bat virus (SHIBV)
Merhavirus Merhavirus merida Merida virus (MERDV)
Merhavirus tritaeniorhynchus Culex tritaeniorhynchus rhabdovirus (CTRV)
Mousrhavirus Mousrhavirus moussa Moussa virus (MOUV)
Ohlsrhavirus Ohlsrhavirus angeles Culex rhabdo-like virus Los Angeles (CRLVLA)
Ohlsrhavirus culex Culex rhabdo-like virus (CRLV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Ohlsrhavirus lobeira Lobeira virus (LOBV)
Ohlsrhavirus northcreek North Creek virus (NORCV)
Ohlsrhavirus ohlsdorf Ohlsdorf virus (OHLDV)
Ohlsrhavirus pseudovishnui Culex pseudovishnui rhabdo-like virus (CpRLV)
Ohlsrhavirus riverside Riverside virus (RISV)
Ohlsrhavirus tongilchon Tongilchon virus 1 (TCHV-1)
Perhabdovirus Perhabdovirus anguilla eel virus American (EVA)
eel virus European X (EVEX)
Perhabdovirus leman Leman virus (LeRV)
Perhabdovirus perca perch rhabdovirus (PRV)
Perhabdovirus trutta lake trout rhabdovirus (LTRV)
Replylivirus Replylivirus allogus anole lyssa-like virus 1 (ALLV-1)
Sawgrhavirus Sawgrhavirus connecticut Connecticut virus (CNTV)
Sawgrhavirus longisland Long Island tick rhabdovirus (LITRV)
Sawgrhavirus minto New Minto virus (NMV)
Sawgrhavirus sawgrass Sawgrass virus (SAWV)
Scophrhavirus Scophrhavirus chanodychthys Wuhan redfin culter dimarhabdovirus (WhRFCRV)
Scophrhavirus maximus Scophthalmus maximus rhabdovirus (SMRV)
Sigmavirus Sigmavirus affinis Drosophila affinis sigmavirus (DAffSV)
Sigmavirus ananassae Drosophila ananassae sigmavirus (DAnaSV)
Sigmavirus capitata Ceratitis capitata sigmavirus (CCapSV)
Sigmavirus domestica Wuhan fly virus 2 (WhFV-2)
Sigmavirus hippoboscid Wuhan louse fly virus 9 (WhLFV-9)
Sigmavirus hubei Hubei diptera virus 9 (HbDV-9)
Sigmavirus immigrans Drosophila immigrans sigmavirus (DImmSV)
Sigmavirus lousefly Wuhan louse fly virus 10 (WhLFV-10)
Sigmavirus melanogaster Drosophila melanogaster sigmavirus (DMelSV)
Sigmavirus muscina Muscina stabulans sigmavirus (MStaSV)
Sigmavirus myga Hubei diptera virus 10 (HbDV-10)
Sigmavirus obscura Drosophila obscura sigmavirus (DObsSV)
Sigmavirus shayang Shayang fly virus 2 (SyFV-1)
Sigmavirus sturtevanti Drosophila sturtevanti sigmavirus (DStuSV)
Sigmavirus tristis Drosophila tristis sigmavirus (DTriSV)
Sigmavirus wuhan Wuhan house fly virus 1 (WhHFV-1)
Sigmavirus ying Hubei dimarhabdovirus 1 (HbDRV-1)
Siniperhavirus Siniperhavirus chuatsi Chinese rice-field eel rhabdovirus (CrERV)
hybrid snakehead rhabdovirus (HSHRV)
Siniperca chuatsi rhabdovirus (SCRV)
Micropterus salmoides rhabdovirus (MSRV)
Siniperhavirus zoarces eelpout rhabdovirus (EPRV)
Sprivivirus Sprivivirus cyprinus spring viremia of carp virus (SVCV)
Sprivivirus esox grass carp rhabdovirus (GrCRV)
pike fry rhabdovirus (PFRV)
tench rhabdovirus (TenRV)
Sripuvirus Sripuvirus almpiwar Almpiwar virus (ALMV)
Sripuvirus chaco Chaco virus (CHOV)
Sripuvirus charleville Charleville virus (CHVV)
Sripuvirus cuiaba Cuiaba virus (CUIV)
Sripuvirus hainan Hainan black-spectacled toad rhabdovirus (HnBSTRV)
Sripuvirus madureira Sena Madureira virus (SMV)

J. H. Kuhn etal.
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Sripuvirus niakha Niakha virus (NIAV)
Sripuvirus sripur Sripur virus (SRIV)
Sunrhavirus Sunrhavirus alexandria Burg el Arab virus (BEAV)
Sunrhavirus bimbo Bimbo virus (BBOV)
Sunrhavirus boteke Boteke virus (BOTV)
Sunrhavirus dillard Dillard’s Draw virus (DDRV)
Sunrhavirus garba Garba virus (GARV)
Sunrhavirus harrison Harrison Dam virus (HARDV)
Sunrhavirus kolongo Kolongo virus (KOLV)
Sunrhavirus kwatta Kwatta virus (KWAV)
Sunrhavirus matariya Matariya virus (MTYV)
Sunrhavirus nasoule Nasoule virus (NASV)
Sunrhavirus oakvale Oak Vale virus (OVV)
Sunrhavirus ouango Ouango virus (OUAV)
Sunrhavirus sandjimba Sandjimba virus (SJAV)
Sunrhavirus sunguru Sunguru virus (SUNV)
Sunrhavirus walkabout Walkabout Creek virus (WACV)
Tibrovirus Tibrovirus alphaekpoma Ekpoma virus 1 (EKV-1)
Tibrovirus beatrice Beatrice Hill virus (BHV)
Tibrovirus betaekpoma Ekpoma virus 2 (EKV-2)
Tibrovirus coastal Coastal Plains virus (CPV)
Tibrovirus congo Bas-Congo virus (BASV)
Tibrovirus sweetwater Sweetwater Branch virus (SWBV)
Tibrovirus tibrogargan Bivens Arm virus (BAV)
Tibrogargan virus (TIBV)
Tupavirus Tupavirus durham Durham virus (DURV)
Tupavirus klamath Klamath virus (KLAV)
Tupavirus tupaia tupaia rhabdovirus (TUPV)
Vesiculovirus Vesiculovirus alagoas vesicular stomatitis Alagoas virus (VSAV)
Vesiculovirus bogdanovac Yug Bogdanovac virus (YBV)
Vesiculovirus carajas Carajás virus (CJSV)
Vesiculovirus chandipura Chandipura virus (CHPV)
Vesiculovirus cocal Cocal virus (COCV)
Vesiculovirus eptesicus American bat vesiculovirus (ABVV)
Vesiculovirus indiana vesicular stomatitis Indiana virus (VSIV)
Vesiculovirus isfahan Isfahan virus (ISFV)
Vesiculovirus jurona Jurona virus (JURV)
Vesiculovirus malpais Malpais Spring virus (MSPV)
Vesiculovirus maraba Maraba virus (MARAV)
Vesiculovirus mediterranean Mediterranean bat virus (MBV)
Vesiculovirus morreton Morreton virus (MORV)
Vesiculovirus newjersey vesicular stomatitis New Jersey virus (VSNJV)
Vesiculovirus perinet Perinet virus (PERV)
Vesiculovirus piry Piry virus (PIRYV)
Vesiculovirus radi Radi virus (RADV)
Vesiculovirus rhinolophus Jinghong bat virus (JhBV)
Qiongzhong bat virus (QZBV)
Vesiculovirus yinshui Yinshui bat virus (YSBV)
Zarhavirus Zarhavirus zahedan Zahedan rhabdovirus (ZARV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Subfamily Betarhabdovirinae
Alphanucleorhabdovirus Alphanucleorhabdovirus colocasiae taro vein chlorosis virus (TaVCV)
Alphanucleorhabdovirus constrictae constricta yellow dwarf virus (CYDV)
Alphanucleorhabdovirus joa joa yellow blotch associated virus (JYBaV)
Alphanucleorhabdovirus maydis maize mosaic virus (MMV)
Alphanucleorhabdovirus melongenae eggplant mottled dwarf virus (EMDV)
Alphanucleorhabdovirus morogoromaydis Morogoro maize-associated virus (MMaV)
Alphanucleorhabdovirus oryzae rice transitory yellowing virus (RTYV)
rice yellow stunt virus (RYSV)
Alphanucleorhabdovirus physostegiae Physostegia chlorotic mottle virus (PhCMoV)
Alphanucleorhabdovirus pruni peach virus 1 (PeV1)
Alphanucleorhabdovirus tritici wheat yellow striate virus (WYSV)
Alphanucleorhabdovirus tuberosum potato yellow dwarf virus (PYDV)
Alphanucleorhabdovirus zeairanense maize Iranian mosaic virus (MIMV)
Betanucleorhabdovirus Betanucleorhabdovirus bacopae Bacopa monnieri virus 2 (BmV-2)
Betanucleorhabdovirus cardamomi cardamom vein clearing virus (CdVCV)
Betanucleorhabdovirus daturae datura yellow vein virus (DYVV)
Betanucleorhabdovirus loti birds-foot trefoil-associated virus (BFTV)
Betanucleorhabdovirus mali apple rootstock virus A (ApRVA)
Betanucleorhabdovirus medicagonis alfalfa-associated nucleorhabdovirus (AaNV)
Betanucleorhabdovirus retesonchi Sonchus yellow net virus (SYNV)
Betanucleorhabdovirus ribes blackcurrant-associated rhabdovirus (BCaRV)
Betanucleorhabdovirus venasonchi sowthistle yellow vein virus (SYVV)
Betanucleorhabdovirus zanthoxyli Zhuye pepper nucleorhabdovirus (ZPNRV)
Cytorhabdovirus Cytorhabdovirus alphatrifolii Trifolium pratense virus A (TpVA)
Cytorhabdovirus alphawuhaninsectum Wuhan insect virus 4 (WuIV-4)
Cytorhabdovirus bacopae Bacopa monnieri virus 1 (BmV-1)
Cytorhabdovirus betatrifolii Trifolium pratense virus B (TpVB)
Cytorhabdovirus betawuhaninsectum Wuhan insect virus 5 (WuIV-5)
Cytorhabdovirus brassicae broccoli necrotic yellows virus (BNYV)
Cytorhabdovirus brassicicolae cabbage cytorhabdovirus 1 (CCyV-1)
Cytorhabdovirus broussonetiae paper mulberry mosaic associated virus (PMuMaV)
Cytorhabdovirus caricae papaya virus E (PpVE)
Cytorhabdovirus chrysanthemi chrysanthemum yellow dwarf associated virus (CYDaV)
Cytorhabdovirus colocasiae Colocasia bobone disease-associated virus (CBDaV)
Cytorhabdovirus cucurbitae cucurbit cytorhabdovirus 1 (CuCV-1)
Cytorhabdovirus festucae Festuca leaf streak virus (FLSV)
Cytorhabdovirus flaviyerbamate yerba mate chlorosis-associated virus (YmCaV)
Cytorhabdovirus fragariae strawberry-associated virus 1 (SaV-1)
strawberry virus 1 (StrV-1)
Cytorhabdovirus fragariarugosus strawberry crinkle virus (SCV)
Cytorhabdovirus gammawuhaninsectum Wuhan insect virus 6 (WuIV-6)
Cytorhabdovirus gramineae northern cereal mosaic virus (NCMV)
Cytorhabdovirus hordei barley yellow striate mosaic virus (BYSMV)
Cytorhabdovirus lactucamaculante lettuce yellow mottle virus (LYMoV)
Cytorhabdovirus lactucanecante lettuce necrotic yellows virus (LNYV)
Cytorhabdovirus lycopersici tomato yellow mottle-associated virus (TYMaV)
Cytorhabdovirus maydis maize-associated cytorhabdovirus (MaCV)
Cytorhabdovirus maysflavostriatis maize yellow striate virus (MYSV)
Cytorhabdovirus medicagonis alfalfa dwarf virus (ADV)

J. H. Kuhn etal.
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Cytorhabdovirus oryzae rice stripe mosaic virus (RSMV)
Cytorhabdovirus persimmon persimmon virus A (PeVA)
Cytorhabdovirus rosae rose virus R (RVR)
Cytorhabdovirus rubus raspberry vein chlorosis virus (RVCV)
Cytorhabdovirus sonchi Sonchus virus (SonV)
Cytorhabdovirus trichosanthei Trichosanthes associated rhabdovirus 1 (TrARV1)
Cytorhabdovirus tritici wheat American striate mosaic virus (WASMV)
Cytorhabdovirus yerbamate yerba mate virus A (YmVA)
Dichorhavirus Dichorhavirus citri citrus chlorotic spot virus (CiCSV)
Dichorhavirus clerodendri clerodendrum chlorotic spot virus (ClCSV)
Dichorhavirus coffeae coffee ringspot virus (CoRSV)
Dichorhavirus leprosis citrus leprosis virus N (CiLV-N)
Dichorhavirus orchidaceae orchid fleck virus (OFV)
Gammanucleorhabdovirus Gammanucleorhabdovirus maydis maize fine streak virus (MFSV)
Varicosavirus Varicosavirus alopecuri Alopecurus myosuroides varicosavirus 1 (AMVV-1)
Varicosavirus lactucae lettuce big-vein associated virus (LBVaV)
Varicosavirus trifolii red clover associated varicosavirus (RCaVV)
Subfamily Gammarhabdovirinae
Novirhabdovirus Novirhabdovirus hirame hirame rhabdovirus (HIRRV = HIRV)
Novirhabdovirus piscine viral hemorrhagic septicemia virus (VHSV)
Novirhabdovirus salmonid infectious hematopoietic necrosis virus (IHNV)
Novirhabdovirus snakehead snakehead rhabdovirus (SHRV)
Unassigned (to subfamilies)
Alphacrustrhavirus Alphacrustrhavirus wenling Wenling crustacean virus 10 (WlCV-10)
Alphacrustrhavirus zhejiang Wenling crustacean virus 11 (WlCV-11)
Alphadrosrhavirus Alphadrosrhavirus hubei Wuhan house fly virus 2 (WhHFV-2)
Alphadrosrhavirus shayang Shayang fly virus 3 (SyFV-3)
Alphahymrhavirus Alphahymrhavirus cinereus hymenopteran rhabdo-related virus 38 (HyRRV-38)
Alphahymrhavirus hirtum hymenopteran rhabdo-related virus 109 (HyRRV-109)
Alphahymrhavirus neglectus Lasius neglectus virus 2 (LnegV-2)
Alphahymrhavirus radians hymenopteran rhabdo-related virus 46 (HyRRV-46)
Betahymrhavirus Betahymrhavirus austriaca hymenopteran rhabdo-related virus 23 (HyRRV-23)
Betahymrhavirus heterodontonyx hymenopteran rhabdo-related virus 24 (HyRRV-24)
Betanemrhavirus Betanemrhavirus hubei Hubei rhabdo-like virus 9 (HbRLV-9)
Betanemrhavirus shayang Shayang ascaridia galli virus 2 (SyAGV-2)
Betapaprhavirus Betapaprhavirus frugiperda Spodoptera frugiperda rhabdovirus (SfruRV)
Betapaprhavirus sylvina lepidopteran rhabdo-related virus 34 (LeRRV-34)
Betaricinrhavirus Betaricinrhavirus chimay Chimay rhabdovirus (CRV)
Betaricinrhavirus scapularis blacklegged tick rhabdovirus 1 (BLTRV-1)
Family Sunviridae
Sunshinevirus Sunshinevirus reptilis Sunshine Coast virus (SunCV)
Family Xinmoviridae
Alasvirus Alasvirus muscae Húběi diptera virus 11 (HbDV-11)
Anphevirus Anphevirus xinchengense Xīnchéng mosquito virus (XcMV)
Doupovirus Doupovirus australiaense Culex mononega-like virus 2 (CMLV-2)
Draselvirus Draselvirus dentati Húběi rhabdo-like virus 7 (HbRLV-7)
Drunivirus Drunivirus chambonense Drosophila unispina virus 1 (DuniV-1)
Gambievirus Gambievirus bolahunense Bolahun virus (BLHV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 4 (continued)
Genus Species¶Virus (Abbreviation)&
Gambievirus senegalense Gambie virus (GAMV)
Gylbovirus Gylbovirus aagae Aedes anphevirus (AeAV)
Hoptevirus Hoptevirus orthopteris Húběi orthoptera virus 5 (HbOV-5)
Madalivirus Madalivirus amapaense Anopheles marajoara virus (AnMV)
Madalivirus amazonaense Anopheles darlingi virus (AnDV)
Pelmivirus Pelmivirus eymattense hymenopteran anphe-related virus OKIAV71 (HARV-71)
Triniovirus Triniovirus yonagoense Culex tritaeniorhynchus anphevirus (CtAV)
Ulegvirus Ulegvirus freckenfeldense odonatan anphe-related virus OKIAV59 (OARV-59)
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, subfamilies, families, and orders are taxa
1 Includes: Newcastle disease virus (NDV)
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper noun. This column lists the virus
names with their correct (lack of) capitalization. Lists of viruses within a given species are provisional at this point and will likely be amended in
the near future
Table 5 ICTV-accepted
taxonomy of the
order Goujianvirales
(Negarnaviricota:
Haploviricotina:
Yunchangviricetes) as of March
2022 Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, fami-
lies, and orders are taxa
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper
noun. This column lists the virus names with their correct (lack of) capitalization
Genus Species¶Virus (Abbreviation)&
Family Yueviridae
Yuyuevirus Yuyuevirus beihaiense Běihǎi sesarmid crab virus 3 (BhSCV-3)
Yuyuevirus shaheense Shāhé yuèvirus-like virus 1 (ShYLV-1)

J. H. Kuhn etal.
1 3
Table 6 ICTV-accepted taxonomy of the order Bunyavirales (Negarnaviricota: Polyploviricotina: Ellioviricetes) as of March 2022
Genus Species¶Virus (Abbreviation)&
Family Arenaviridae
Antennavirus Hairy antennavirus Wēnlǐng frogfish arenavirus 2 (WlFAV-2)
Salmon antennavirus salmon pescarenavirus 1 (SPAV-1)
salmon pescarenavirus 2 (SPAV-2)
Striated antennavirus Wēnlǐng frogfish arenavirus 1 (WlFAV-1)
Hartmanivirus Haartman hartmanivirus Haartman Institute snake virus 1 (HISV-1)
Haartman Institute snake virus 2 (HISV-2)
Heimat hartmanivirus andere Heimat virus 1 (aHeV-1)
Muikkunen hartmanivirus Dante Muikkunen virus 1 (DaMV-1)
Schoolhouse hartmanivirus old schoolhouse virus 1 (OScV-1)
old schoolhouse virus 2 (OScV-2)
Setpatvet hartmanivirus SetPatVet virus 1 (SPVV-1)
Zurich hartmanivirus veterinary pathology Zurich virus 1 (VPZV-1)
veterinary pathology Zurich virus 2 (VPZV-2)
Mammarenavirus Allpahuayo mammarenavirus Allpahuayo virus (ALLV)
Alxa mammarenavirus Alxa virus (ALXV)
Argentinian mammarenavirus Junín virus (JUNV)
Bear Canyon mammarenavirus Bear Canyon virus (BCNV)
Brazilian mammarenavirus Sabiá virus (SBAV)
Cali mammarenavirus Pichindé virus (PICHV)
Chapare mammarenavirus Chapare virus (CHAPV)
Chevrier mammarenavirus Lìjiāng virus (LIJV)
Cupixi mammarenavirus Cupixi virus (CUPXV)
Flexal mammarenavirus Flexal virus (FLEV)
Gairo mammarenavirus Gairo virus (GAIV)
Guanarito mammarenavirus Guanarito virus (GTOV)
Ippy mammarenavirus Ippy virus (IPPYV)
Kitale mammarenavirus Kitale virus (KTLV)
Lassa mammarenavirus Lassa virus (LASV)
Latino mammarenavirus Latino virus (LATV)
Loei River mammarenavirus Loei River virus (LORV)
Lujo mammarenavirus Lujo virus (LUJV)
Luna mammarenavirus Luli virus (LULV)
Luna virus (LUAV)
Lunk mammarenavirus Lunk virus (LNKV)
Lymphocytic choriomeningitis mammare-
navirus Dandenong virus (DANV)
lymphocytic choriomeningitis virus (LCMV)
Machupo mammarenavirus Machupo virus (MACV)
Mammarenavirus bituense Bitu virus (BITV)
Mammarenavirus kwanzaense Kwanza virus (KWAV)
Mariental mammarenavirus Mariental virus (MRLV)
Merino Walk mammarenavirus Merino Walk virus (MRWV)
Mobala mammarenavirus mobala virus (MOBV)
Mopeia mammarenavirus Mopeia virus (MPOV)
Morogoro virus (MORV)
Okahandja mammarenavirus Okahandja virus (OKAV)
Oliveros mammarenavirus Oliveros virus (OLVV)
Paraguayan mammarenavirus Paraná virus (PRAV)
Planalto mammarenavirus Aporé virus (APOV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Pirital mammarenavirus Pirital virus (PIRV)
Ryukyu mammarenavirus Ryukyu virus (RYKV)
Serra do Navio mammarenavirus Amaparí virus (AMAV)
Solwezi mammarenavirus Solwezi virus (SOLV)
Souris mammarenavirus souris virus (SOUV)
Tacaribe mammarenavirus Tacaribe virus (TCRV)
Tamiami mammarenavirus Tamiami virus (TMMV)
Wenzhou mammarenavirus Wēnzhōu virus (WENV)
Whitewater Arroyo mammarenavirus Big Brushy Tank virus (BBRTV)
Catarina virus (CTNV)
Skinner Tank virus (SKTV)
Tonto Creek virus (TTCV)
Whitewater Arroyo virus (WWAV)
Xapuri mammarenavirus Xapuri virus (XAPV)
Reptarenavirus California reptarenavirus CAS virus (CASV)
Giessen reptarenavirus University of Giessen virus 1 (UGV-1)
University of Giessen virus 2 (UGV-2)
University of Giessen virus 3 (UGV-3)
Golden reptarenavirus Golden Gate virus (GOGV)
Ordinary reptarenavirus tavallinen suomalainen mies virus 2 (TSMV-2)
Rotterdam reptarenavirus ROUT virus (ROUTV)
University of Helsinki virus 1 (UHV-1)
Family Cruliviridae
Lincruvirus Lincruvirus europense European shore crab virus 1 (EscV-1)
Lincruvirus sinense Chinese mitten crab virus 1 (CmcV-1)
Lincruvirus wenlingense Wēnlǐng crustacean virus 9 (WlCV-9)
Family Discoviridae
Orthodiscovirus Orthodiscovirus coniellae Coniothyrium diplodiella negative-stranded RNA virus 1
(CdNSRV-1)
Plasmopara viticola lesion associated mycobunyavirales-like virus 9
(PvLAM-LV-9)
Orthodiscovirus hispaniae Plasmopara viticola lesion associated mycobunyavirales-like virus 8
(PvLAM-LV8)
Orthodiscovirus iberiae Plasmopara viticola lesion associated mycobunyavirales-like virus 4
(PvLAM-LV4)
Orthodiscovirus missouriense Penicillium discovirus (PDV)
Orthodiscovirus penicillii Penicillium roseopurpureum negative ssRNA virus 1 (PrNssV-1)
Family Fimoviridae
Emaravirus Emaravirus aceris maple mottle-associated virus (MaMaV)
Emaravirus actinidiae Actinidia chlorotic ringspot-associated virus (AcCRaV)
Emaravirus cajani pigeonpea sterility mosaic virus 1 (PPSMV-1)
Emaravirus camelliae Camellia japonica-associated virus 1 (CjaV-1)
Emaravirus cercidis redbud yellow ringspot-associated virus (RYRaV)
Emaravirus chrysanthemi chrysanthemum mosaic-associated virus (ChMaV)
Emaravirus cordylinae ti ringspot-associated virus (TiRSaV)
Emaravirus fici fig mosaic virus (FMV)
Emaravirus idaeobati raspberry leaf blotch virus (RLBV)
Emaravirus kiwii Actinidia virus 2 (AcV-2)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Emaravirus parkinsoniae palo verde broom virus (PVBV)
Emaravirus perillae perilla mosaic virus (PerMV)
Emaravirus pistaciae Pistacia virus B (PiVB)
Emaravirus populi aspen mosaic-associated virus (AsMaV)
Emaravirus pyri pear chlorotic leaf spot-associated virus (PCLSaV)
Emaravirus quercus common oak ringspot-associated virus (CORaV)
Emaravirus rosae rose rosette virus (RRV)
Emaravirus rubi blackberry leaf mottle-associated virus (BLMaV)
Emaravirus sorbi European mountain ash ringspot-associated virus (EMARaV)
Emaravirus syringae lilac chlorotic ringspot-associated virus (LiCRaV)
Emaravirus toordali pigeonpea sterility mosaic virus 2 (PPSMV-2)
Emaravirus tritici High Plains wheat mosaic virus (HPWMoV)
Emaravirus verbanni Camellia japonica-associated virus 2 (CjaV-2)
Emaravirus ziziphi jujube yellow mottle-associated virus (JYMaV)
Family Hantaviridae
Subfamily Actantavirinae
Actinovirus Batfish actinovirus Wēnlǐng minipizza batfish virus (WEMBV)
Goosefish actinovirus Wēnlǐng yellow goosefish virus (WEYGV)
Perch actinovirus Bern perch virus (BRPV)
Spikefish actinovirus Wēnlǐng red spikefish virus (WERSV)
Subfamily Agantavirinae
Agnathovirus Hagfish agnathovirus Wēnlǐng hagfish virus (WEHV)
Subfamily Mammantavirinae
Loanvirus Brno loanvirus Brno virus (BRNV)
Longquan loanvirus Lóngquán virus (LQUV)
Mobatvirus Laibin mobatvirus Láibīn virus (LAIV)
Lena mobatvirus Lena virus (LENV)
Nova mobatvirus Nova virus (NVAV)
Quezon mobatvirus Quezon virus (QZNV)
Xuan Son mobatvirus Xuân Sơn virus (XSV)
Orthohantavirus Andes orthohantavirus Andes virus (ANDV)
Castelo dos Sonhos virus (CASV)
Lechiguanas virus (LECV = LECHV)
Orán virus (ORNV)
Asama orthohantavirus Asama virus (ASAV)
Asikkala orthohantavirus Asikkala virus (ASIV)
Bayou orthohantavirus bayou virus (BAYV)
Catacamas virus (CATV)
Black Creek Canal orthohantavirus Black Creek Canal virus (BCCV)
Bowe orthohantavirus Bowé virus (BOWV)
Bruges orthohantavirus Bruges virus (BRGV)
Cano Delgadito orthohantavirus Caño Delgadito virus (CADV)
Cao Bang orthohantavirus Cao Bằng virus (CBNV)
Liánghé virus (LHEV)
Choclo orthohantavirus Choclo virus (CHOV)
Dabieshan orthohantavirus Dàbiéshān virus (DBSV)
Dobrava-Belgrade orthohantavirus Dobrava virus (DOBV)
Kurkino virus (KURV)
Saaremaa virus (SAAV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Sochi virus (SOCV)
El Moro Canyon orthohantavirus Carrizal virus (CARV)
El Moro Canyon virus (ELMCV)
Huitzilac virus (HUIV)
Fugong orthohantavirus Fúgòng virus (FUGV)
Fusong orthohantavirus Fǔsōng virus (FUSV)
Hantaan orthohantavirus Amur virus (AMRV)
Hantaan virus (HTNV)
Soochong virus (SOOV)
Jeju orthohantavirus Jeju virus (JJUV)
Kenkeme orthohantavirus Kenkeme virus (KKMV)
Khabarovsk orthohantavirus Khabarovsk virus (KHAV)
Topografov virus (TOPV)
Laguna Negra orthohantavirus Laguna Negra virus (LANV)
Maripa virus (MARV)
Rio Mamoré virus (RIOMV)
Luxi orthohantavirus Lúxī virus (LUXV)
Maporal orthohantavirus Maporal virus (MAPV)
Montano orthohantavirus Montaño virus (MTNV)
Necocli orthohantavirus Necoclí virus (NECV)
Oxbow orthohantavirus Oxbow virus (OXBV)
Prospect Hill orthohantavirus Prospect Hill virus (PHV)
Puumala orthohantavirus Hokkaido virus (HOKV)
Muju virus (MUJV)
Puumala virus (PUUV)
Robina orthohantavirus Robina virus (ROBV)
Rockport orthohantavirus Rockport virus (RKPV)
Sangassou orthohantavirus Sangassou virus (SANGV)
Seewis orthohantavirus Seewis virus (SWSV)
Seoul orthohantavirus gōu virus (GOUV)
Seoul virus (SEOV)
Sin Nombre orthohantavirus New York virus (NYV)
Sin Nombre virus (SNV)
Tatenale orthohantavirus Tatenale virus (TATV)
Thailand orthohantavirus Anjozorobe virus (ANJZV)
Serang virus (SERV)
Thailand virus (THAIV)
Tigray orthohantavirus Tigray virus (TIGV)
Tula orthohantavirus Adler virus (ADLV)
Tula virus (TULV)
Yakeshi orthohantavirus Yákèshí virus (YKSV)
Thottimvirus Imjin thottimvirus Imjin virus (MJNV)
Thottapalayam thottimvirus Thottapalayam virus (TPMV)
Subfamily Repantavirinae
Reptillovirus Gecko reptillovirus Hǎinán oriental leaf-toed gecko virus (HOLGV)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Family Leishbuviridae
Shilevirus Shilevirus leptomonadis Leptomonas moramango virus (LEPMV)
Family Mypoviridae
Hubavirus Hubavirus myriapedis Húběi myriapoda virus 5 (HbMV-5)
Family Nairoviridae
Norwavirus Grotenhout norwavirus Grotenhout virus (GRHV)
Ocetevirus Blattodean ocetevirus red goblin roach virus 1 (RGRV-1)
Orthonairovirus Abu Hammad orthonairovirus Abū Ḥammād virus (AHV)
Abu Mina orthonairovirus Abū Mīnā virus (AMV)
Artashat orthonairovirus Artashat virus (ARTSV)
Avalon orthonairovirus Avalon virus (AVAV)
Chim orthonairovirus Chim virus (CHIMV)
Bandia orthonairovirus Bandia virus (BDAV)
Burana orthonairovirus Burana virus (BURV)
Congoid orthonairovirus Aigai virus (AIGV)
Crimean-Congo hemorrhagic fever ortho-
nairovirus Crimean-Congo hemorrhagic fever virus (CCHFV)
Dera Ghazi Khan orthonairovirus Dera Ghazi Khan virus (DGKV)
Dugbe orthonairovirus Dugbe virus (DUGV)
Erve orthonairovirus Erve virus (ERVEV)
Estero Real orthonairovirus Estero Real virus (ERV)
Gossas orthonairovirus Gossas virus (GOSV)
Hazara orthonairovirus Hazara virus (HAZV)
Huangpi orthonairovirus Huángpí tick virus 1 (HpTV-1)
Hughes orthonairovirus Caspiy virus (CASV)
Farallon virus (FARV)
Great Saltee virus (GRSV)
Hughes virus (HUGV)
Raza virus (RAZAV)
Issyk-kul orthonairovirus Issyk-kul virus (ISKV)
Kasokero orthonairovirus Kasokero virus (KASV = KASOV)
Keterah orthonairovirus Keterah virus (KTRV)
Uzun-Agach virus (UZAV)
Kupe orthonairovirus kupe virus (KUPEV)
Leopards Hill orthonairovirus Leopards Hill virus (LPHV)
Meram orthonairovirus Meram virus (MEMV)
Nairobi sheep disease orthonairovirus Nairobi sheep disease virus (NSDV)
Pacific Coast orthonairovirus Pacific Coast tick nairovirus (PCTNV)
Punta orthonairovirus Punta Salinas virus (PSV)
Qalyub orthonairovirus Geran virus (GERV)
Qalyub virus (QYBV)
Sakhalin orthonairovirus Sakhalin virus (SAKV)
Tillamook virus (TILLV)
Sapphire orthonairovirus Sapphire II virus (SAPV)
Scot orthonairovirus Clo Mor virus (CMV = CLMV)
Soldado orthonairovirus Soldado virus (SOLV)
Tacheng orthonairovirus Tǎchéng tick virus 1 (TcTV-1)
Taggert orthonairovirus Taggert virus (TAGV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Tamdy orthonairovirus Tamdy virus (TAMV)
Thiafora orthonairovirus Thiafora virus (TFAV)
Tofla orthonairovirus tofla virus (TFLV)
Tunis orthonairovirus Tunis virus (TUNV)
Vinegar Hill orthonairovirus Vinegar Hill virus (VINHV)
Wenzhou orthonairovirus Wēnzhōu tick virus (WzTV)
Yogue orthonairovirus Yogue virus (YOGV)
Zirqa orthonairovirus Zirqa virus (ZIRV)
Sabavirus South Bay sabavirus South Bay virus (SBV)
Shaspivirus Spider shaspivirus Shāyáng spider virus 1 (SySV-1)
Striwavirus Strider striwavirus Sānxiá water strider virus 1 (SxWSV-1)
Xinspivirus Xinzhou xinspivirus Xīnzhōu spider virus (XSV)
Family Peribunyaviridae
Herbevirus Herbert herbevirus Herbert virus (HEBV)
Kibale herbevirus Kibale virus (KIBV)
Tai herbevirus Taï virus (TAIV)
Khurdivirus Khurdivirus volgaense Khurdun virus (KURV)
Lakivirus Lakivirus lakamhaense Lakamha virus (LAKV)
Lambavirus Lambavirus wisconsinense largemouth bass bunyavirus (LBBV)
Orthobunyavirus Abras orthobunyavirus Abras virus (ABRV)
Acara orthobunyavirus Acará virus (ACAV)
Moriche virus (MORV)
Aino orthobunyavirus Aino virus (AINOV)
Akabane orthobunyavirus Akabane virus (AKAV)
Tinaroo virus (TINV)
Yaba-7 virus (Y7V)
Alajuela orthobunyavirus Alajuela virus (ALJV)
San Juan virus (SJV)
Anadyr orthobunyavirus Anadyr virus (ANADV)
Ananindeua orthobunyavirus Ananindeua virus (ANUV)
Anhembi orthobunyavirus Anhembi virus (AMBV)
Anopheles A orthobunyavirus Anopheles A virus (ANAV)
Arumateua virus (ARTV = ARMTV)
Caraipé virus (CPEV = CRPV)
Las Maloyas virus (LMV)
Lukuni virus (LUKV)
Trombetas virus (TRMV)
Tucuruí virus (TUCV = TUCRV)
Anopheles B orthobunyavirus Anopheles B virus (ANBV)
Boracéia virus (BORV)
Apeu orthobunyavirus Apeú virus (APEUV)
Bakau orthobunyavirus Bakau virus (BAKV)
Ketapang virus (KETV)
Nola virus (NOLAV)
Tanjong Rabok virus (TRV)
Telok Forest virus (TFV)
Batai orthobunyavirus Batai virus (BATV)
Batama orthobunyavirus Batama virus (BMAV)
Bellavista orthobunyavirus Bellavista virus (BELLV)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Benevides orthobunyavirus Benevides virus (BVSV = BENV)
Bertioga orthobunyavirus Bertioga virus (BERV)
Cananéia virus (CNAV)
Guaratuba virus (GTBV)
Itimirim virus (ITIV)
Mirim virus (MIRV)
Bimiti orthobunyavirus bimiti virus (BIMV)
Birao orthobunyavirus Birao virus (BIRV)
Botambi orthobunyavirus Botambi virus (BOTV)
Bozo orthobunyavirus Bozo virus (BOZOV)
Brazoran orthobunyavirus brazoran virus (BRAZV)
Bruconha orthobunyavirus Bruconha virus (BRUV)
Buffalo Creek orthobunyavirus Buffalo Creek virus (BUCV)
Bunyamwera orthobunyavirus Bunyamwera virus (BUNV)
Germiston virus (GERV)
Lokern virus (LOKV)
Mboké virus (MBOV)
Ngari virus (NRIV)
Northway virus (NORV)
Santa Rosa virus (SARV)
Shokwe virus (SHOV)
Stanfield virus (STAV)
Xingu virus (XINV)
Bushbush orthobunyavirus Benfica virus (BENV = BNFV)
Bushbush virus (BSBV)
Juan Díaz virus (JDV)
Buttonwillow orthobunyavirus Buttonwillow virus (BUTV)
Bwamba orthobunyavirus Bwamba virus (BWAV)
Pongola virus (PGAV)
Cache Valley orthobunyavirus Cache Valley virus (CVV)
Cholul virus (CHLV)
Tlacotalpan virus (TLAV)
Cachoeira Porteira orthobunyavirus Cachoeira Porteira virus (CPOV)
California encephalitis orthobunyavirus California encephalitis virus (CEV)
Morro Bay virus (MBV)
Capim orthobunyavirus Capim virus (CAPV)
Caraparu orthobunyavirus Caraparú virus (CARV)
El Huayo virus (EHUV)
Itaya virus (ITYV)
Ossa virus (OSSAV)
Vinces virus (VINV)
Cat Que orthobunyavirus Cát Quế virus (CQV)
Oya virus (OYAV)
Catu orthobunyavirus Catú virus (CATUV)
Enseada orthobunyavirus Enseada virus (ENSV)
Faceys paddock orthobunyavirus Facey’s paddock virus (FPV)
Fort Sherman orthobunyavirus Fort Sherman virus (FSV)
Gamboa orthobunyavirus Brus Laguna virus (BLAV)
Calchaquí virus (CQIV)
Gamboa virus (GAMV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Pueblo Viejo virus (PVV)
Soberanía virus (SOBV)
Gan Gan orthobunyavirus Gan Gan virus (GGV)
Guajara orthobunyavirus Guajará virus (GJAV)
Guama orthobunyavirus Guamá virus (GMAV)
Guaroa orthobunyavirus Guaroa virus (GROV)
Iaco orthobunyavirus Iaco virus (IACOV)
Ilesha orthobunyavirus Ilesha virus (ILEV)
Ingwavuma orthobunyavirus Ingwavuma virus (INGV)
Jamestown Canyon orthobunyavirus Inkoo virus (INKV)
Jamestown Canyon virus (JCV)
Jerry Slough virus (JSV)
South River virus (SORV)
Jatobal orthobunyavirus Jatobal virus (JATV)
Kaeng Khoi orthobunyavirus Kaeng Khoi virus (KKV)
Kairi orthobunyavirus kairi virus (KRIV)
Keystone orthobunyavirus Keystone virus (KEYV)
Koongol orthobunyavirus koongol virus (KOOV)
wongal virus (WONV)
La Crosse orthobunyavirus La Crosse virus (LACV)
Leanyer orthobunyavirus Leanyer virus (LEAV)
Lumbo orthobunyavirus Lumbo virus (LUMV)
Macaua orthobunyavirus Macauã virus (MCAV)
Madrid orthobunyavirus Madrid virus (MADV)
Maguari orthobunyavirus Maguari virus (MAGV)
Playas virus (PLAV)
Mahogany Hammock orthobunyavirus Mahogany Hammock virus (MHV)
Main Drain orthobunyavirus Main Drain virus (MDV)
Manzanilla orthobunyavirus Manzanilla virus (MANV)
Inini virus (INIV)
Mapputta orthobunyavirus Mapputta virus (MAPV)
Maprik orthobunyavirus Maprik virus (MPKV)
Marituba orthobunyavirus Gumbo Limbo virus (GLV)
Marituba virus (MTBV)
Murutucú virus (MURV)
Nepuyo virus (NEPV)
Restan virus (RESV)
Zungarococha virus (ZUNV)
Matruh orthobunyavirus Matruh virus (MTRV)
Melao orthobunyavirus Melao virus (MELV)
Moju orthobunyavirus Moju virus (MOJUV)
Mermet orthobunyavirus Mermet virus (MERV)
Minatitlan orthobunyavirus Minatitlán virus (MNTV)
Palestina virus (PLSV)
MPoko orthobunyavirus M'Poko virus (MPOV)
Yaba-1 virus (Y1V)
Nyando orthobunyavirus Eretmapodites virus (ERETV)
Mojuí dos Campos virus (MDCV)
Nyando virus (NDV)
Olifantsvlei orthobunyavirus Bobia virus (BIAV)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Dabakala virus (DABV)
Olifantsvlei virus (OLIV)
Oubi virus (OUBIV)
Oriboca orthobunyavirus Itaquí virus (ITQV)
Oriboca virus (ORIV)
Oropouche orthobunyavirus Iquitos virus (IQTV)
Madre de Dios virus (MDDV)
Oropouche virus (OROV)
Perdões virus (PDEV)
Pintupo virus (PINTV)
Oyo orthobunyavirus Oyo virus (OYOV)
Patois orthobunyavirus Babahoya virus (BABV)
Pahayokee virus (PAHV)
Patois virus (PATV)
Peaton orthobunyavirus peaton virus (PEAV)
Potosi orthobunyavirus Potosi virus (POTV)
Sabo orthobunyavirus Sabo virus (SABOV)
San Angelo orthobunyavirus San Angelo virus (SAV)
Sango orthobunyavirus Sango virus (SANV)
Schmallenberg orthobunyavirus Douglas virus (DOUV)
Sathuperi virus (SATV)
Schmallenberg virus (SBV)
Shamonda virus (SHAV)
Sedlec orthobunyavirus Sedlec virus (SEDV)
Serra do Navio orthobunyavirus Serra do Navio virus (SDNV)
Shark River orthobunyavirus Shark River virus (SRV)
Shuni orthobunyavirus Kaikalur virus (KAIV)
Shuni virus (SHUV)
Simbu orthobunyavirus Para virus (PARAV)
Simbu virus (SIMV)
Snowshoe hare orthobunyavirus Khatanga virus (KHATV)
snowshoe hare virus (SSHV)
Sororoca orthobunyavirus Sororoca virus (SORV)
Tacaiuma orthobunyavirus CoAr 1071 virus (CA1071V)
CoAr 3627 virus (CA3626V)
Tacaiuma virus (TCMV)
Virgin River virus (VRV)
Tahyna orthobunyavirus Ťahyňa virus (TAHV)
Tataguine orthobunyavirus Tataguine virus (TATV)
Tensaw orthobunyavirus Tensaw virus (TENV)
Tete orthobunyavirus Bahig virus (BAHV)
Tete virus (TETEV)
Tsuruse virus (TSUV)
Weldona virus (WELV)
Thimiri orthobunyavirus Thimiri virus (THIV)
Timboteua orthobunyavirus Timboteua virus (TBTV)
Triniti orthobunyavirus Triniti virus (TNTV)
Trivittatus orthobunyavirus Achiote virus (ACHOV)
Trivittatus virus (TVTV)
Turlock orthobunyavirus Lednice virus (LEDV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Turlock virus (TURV)
Umbre virus (UMBV)
Utinga orthobunyavirus Utinga virus (UTIV)
Witwatersrand orthobunyavirus Witwatersrand virus (WITV)
Wolkberg orthobunyavirus Wolkberg virus (WBV)
Wyeomyia orthobunyavirus Rio Pracupi virus
Taiassui virus (TAIAV)
Tucunduba virus (TUCV)
Wyeomyia virus (WYOV)
Pacuvirus Caimito pacuvirus Caimito virus (CAIV)
Chilibre pacuvirus Chilibre virus (CHIV)
Pacui pacuvirus Pacui virus (PACV)
Rio Preto da Eva pacuvirus Rio Preto da Eva virus (RPEV)
Tapirape pacuvirus Tapirapé virus (TAPV)
Shangavirus Insect shangavirus Shuāngào insect virus 1 (SgIV-1)
Family Phasmaviridae
Feravirus Ferak feravirus Ferak virus (FRKV)
Hemipteran feravirus hemipteran phasma-related virus OKIAV247 (HeFV)
Neuropteran feravirus neuropteran phasma-related virus OKIAV248 (NeFV)
Hymovirus Hymenopteran hymovirus 1 hymenopteran phasma-related virus OKIAV252 (HyHV-1)
Hymenopteran hymovirus 2 hymenopteran phasma-related virus OKIAV250 (HyHV-2)
Jonvirus Jonchet jonvirus jonchet virus (JONV)
Orthophasmavirus Anopheles orthophasmavirus Anopheles triannulatus orthophasmavirus (AtOPV)
Coleopteran orthophasmavirus coleopteran phasma-related virus OKIAV235 (CPRV)
Culex orthophasmavirus Culex phasma-like virus (CPLV)
Ganda orthophasmavirus Ganda bee virus (GBEEV)
Hymenopteran orthophasmavirus 1 hymenopteran phasma-related virus OKIAV228 (HyOV-1)
Hymenopteran orthophasmavirus 2 hymenopteran phasma-related virus OKIAV227 (HyOV-2)
Kigluaik phantom orthophasmavirus Kigluaik phantom virus (KIGV)
Niukluk phantom orthophasmavirus Niukluk phantom virus (NUKV)
Odonate orthophasmavirus Húběi odonate virus 8 (HbOV-8)
Orthophasmavirus barstukasense Barstukas virus (BARV)
Orthophasmavirus coredoense Coredo virus (CORV)
Orthophasmavirus flenense Flen virus (FLNV)
Orthophasmavirus miglotasense Miglotas virus (MIGV)
Qingling orthophasmavirus Húběi odonate virus 9 (HbOV-9)
Wuchang cockroach orthophasmavirus 1 Wǔchāng cockroach virus 1 (WcCV-1)
Wuhan mosquito orthophasmavirus 1 Wǔhàn mosquito virus 1 (WhMV-1)
Wuhan mosquito orthophasmavirus 2 Wǔhàn mosquito virus 2 (WhMV-2)
Sawastrivirus Sanxia sawastrivirus Sānxiá water strider virus 2 (SxWSV-2)
Wuhivirus Insect wuhivirus Wǔhàn insect virus 2 (WhIV-2)
Family Phenuiviridae
Bandavirus Bhanja bandavirus Bhanja virus (BHAV)
Dabie bandavirus severe fever with thrombocytopenia syndrome virus (SFTSV)
Guertu bandavirus Gùěrtú virus (GTV)
Heartland bandavirus Heartland virus (HRTV)
Hunter Island bandavirus Albatross Island virus (ABIV)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Kismaayo bandavirus Kismaayo virus (KISV)
Lone star bandavirus lone star virus (LSV)
Razdan bandavirus Razdan virus (RAZV)
Beidivirus Dipteran beidivirus Húběi diptera virus 3 (HbDV-3)
Coguvirus Citrus coguvirus citrus concave gum-associated virus (CCGaV)
Coguvirus eburi citrus virus A (CiV-A)
Grapevine coguvirus grapevine associated cogu-like virus 1 (GaCLV-1)
Entovirus Entoleuca entovirus Entoleuca phenui-like virus 1 (EnPLV-1)
Goukovirus Cumuto goukovirus Cumuto virus (CUMV)
Gouleako goukovirus Gouléako virus (GOLV)
Yichang insect goukovirus Yíchāng insect virus (YcIV)
Horwuvirus Horsefly horwuvirus Wǔhàn horsefly virus (WhHV)
Kimberley horwuvirus Fitzroy Crossing tenui-like virus 1 (FCTenV1)
Hudivirus Dipteran hudivirus Húběi diptera virus 4 (HbDV-4)
Hudovirus Lepidopteran hudovirus Húběi lepidoptera virus 1 (HbLV-1)
Ixovirus Blackleg ixovirus blacklegged tick virus 1 (BLTV-1)
Norway ixovirus Fairhair virus (FHAV)
Scapularis ixovirus blacklegged tick virus 3 (BLTV-3)
Laulavirus Grapevine laulavirus 2 grapevine associated cogu-like virus 2 (GaCLV-2)
Grapevine laulavirus 3 grapevine associated cogu-like virus 3 (GaCLV-3)
Grapevine laulavirus 4 grapevine associated cogu-like virus 4 (GaCLV-4)
Laurel Lake laulavirus Laurel Lake virus (LLV)
Lentinuvirus Lentinula lentinuvirus Lentinula edodes negative-strand RNA virus 2 (LeNSRV-2)
Mobuvirus Mothra mobuvirus Mothra virus (MTHV)
Narangue mobuvirus Ñarangue virus (NRGV)
Phasivirus Badu phasivirus Badu virus (BADUV)
Dipteran phasivirus Húběi diptera virus 5 (HbDV-5)
Fly phasivirus Wǔhàn fly virus 1 (WhFV-1)
Guadeloupe phasivirus Guadeloupe mosquito phasivirus (GMPV)
Kimberley phasivirus Parry’s Creek phasivirus 1 (PCPhasV1)
Phasi Charoen-like phasivirus Phasi Charoen-like virus (PCLV)
Wutai mosquito phasivirus Wǔtái mosquito virus (WtMV)
Phlebovirus Adana phlebovirus Adana virus (ADAV)
Aguacate phlebovirus Aguacate virus (AGUV)
Alcube phlebovirus Alcube virus (ACBV)
Alenquer phlebovirus Alenquer virus (ALEV)
Ambe phlebovirus Ambé virus (ABEV)
Anhanga phlebovirus Anhangá virus (ANHV)
Arumowot phlebovirus Arumowot virus (AMTV)
Buenaventura phlebovirus Buenaventura virus (BUEV)
Bogoria phlebovirus Bogoria virus (BGRV)
Bujaru phlebovirus Bujaru virus (BUJV)
Cacao phlebovirus Cacao virus (CACV)
Campana phlebovirus Campana virus (CMAV)
Candiru phlebovirus Ariquemes virus (ARQV)
Candirú virus (CDUV)
Jacundá virus (JCNV)
Morumbi virus (MR(M)BV)
Mucura virus (MCRV/MRAV)
Serra Norte virus (SRNV)

Taxonomic update of phylum Negarnaviricota
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Chagres phlebovirus Chagres virus (CHGV)
Cocle phlebovirus Coclé virus (CCLV)
Corfou phlebovirus Corfou virus (CFUV)
Dashli phlebovirus Dāshlī virus (DASV)
Durania phlebovirus Durania virus (DRNV)
Echarate phlebovirus Echarate virus (ECHV)
Embossos phlebovirus Embossos virus (EMBV)
Gabek phlebovirus Gabek forest virus (GFV)
Gordil phlebovirus Gordil virus (GORV)
Icoaraci phlebovirus Icoaraci virus (ICOV)
Itaituba phlebovirus Itaituba virus (ITAV)
Itaporanga phlebovirus Itaporanga virus (ITPV)
Ixcanal phlebovirus Ixcanal virus (IXCV)
Karimabad phlebovirus Karimabad virus (KARV)
Kiborgoch phlebovirus Kiborgoch virus (KBGV)
La Gloria phlebovirus La Gloria virus (LAGV)
Lara phlebovirus Rio Claro virus (RICV)
Leticia phlebovirus Leticia virus (LTCV)
Maldonado phlebovirus Maldonado virus (MLOV)
Massilia phlebovirus Massilia virus (MASV)
Medjerda phlebovirus Medjerda Valley virus (MVV)
Mona Grita phlebovirus Mona Grita virus (MOGV)
Mukawa phlebovirus Mukawa virus (MKWV)
Munguba phlebovirus Munguba virus (MUNV)
Naples phlebovirus Arrábida virus (ARRV)
Balkan virus (BALKV)
Fermo virus (FERV)
Granada virus (GRV = GRAV)
Saddaguia virus (SADV)
sandfly fever Naples virus (SFNV)
Nique phlebovirus Nique virus (NIQV)
Ntepes phlebovirus Ntepes virus (NTPV)
Odrenisrou phlebovirus Odrénisrou virus (ODRV)
Oriximina phlebovirus Oriximiná virus (ORXV)
Pena Blanca phlebovirus Peña Blanca virus (PEBV)
Penshurt phlebovirus Penshurt virus (PEHV)
Perkerra phlebovirus Perkerra virus (PKEV)
Punique phlebovirus Punique virus (PUNV)
Punta Toro phlebovirus Capira virus (CAPIV)
Punta Toro virus (PTV)
Rift Valley fever phlebovirus Rift Valley fever virus (RVFV)
Rio Grande phlebovirus Rio Grande virus (RGV)
Saint Floris phlebovirus Saint-Floris virus (SAFV)
Salanga phlebovirus Salanga virus (SLGV)
Salehabad phlebovirus Adria virus (ADRV)
Arbia virus (ARBV)
Bregalaka virus (BREV)
Olbia virus (OLBV)
Salehabad virus (SALV)
Zaba virus (ZABAV)

J. H. Kuhn etal.
1 3
Table 6 (continued)
Genus Species¶Virus (Abbreviation)&
Salobo phlebovirus Salobo virus (SLBOV)
Sicilian phlebovirus sandfly fever Sicilian virus (SFSV)
Tapara phlebovirus Tapará virus (TPRV)
Tehran phlebovirus Tehran virus (THEV)
Tico phlebovirus Tico virus (TICV)
Toros phlebovirus Toros virus (TORV)
Toscana phlebovirus Toscana virus (TOSV)
Tres Almendras phlebovirus Tres Almendras virus (TRAV)
Turuna phlebovirus Turuna virus (TUAV)
Uriurana phlebovirus Uriurana virus (URIV)
Urucuri phlebovirus Urucuri virus (URUV)
Viola phlebovirus viola virus (VIOV)
Zerdali phlebovirus Zerdali virus (ZERV)
Pidchovirus Coleopteran pidchovirus coleopteran phenui-related virus 308 (CoPrV-308)
Pidgey pidchovirus Pidgey virus (PGYV)
Rubodvirus Apple rubodvirus 1 apple rubbery wood virus 1 (ARWV-1)
Apple rubodvirus 2 apple rubbery wood virus 2 (ARWV-2)
Grapevine rubodvirus 1 grapevine Garan dmak virus (GGDV)
Grapevine rubodvirus 2 grapevine Muscat rose virus (GMRV)
Tanzavirus Human tanzavirus Dar es Salaam virus (DeSV)
Tenuivirus Echinochloa hoja blanca tenuivirus Echinochloa hoja blanca virus (EHBV)
European wheat striate mosaic tenuivirus European wheat striate mosaic virus (EWSMV)
Iranian wheat stripe tenuivirus Iranian wheat stripe virus (IWSV)
Maize stripe tenuivirus maize stripe virus (MStV = MSpV)
Melon tenuivirus melon chlorotic spot virus (MeCSV)
Rice grassy stunt tenuivirus rice grassy stunt virus (RGSV)
Rice hoja blanca tenuivirus rice hoja blanca virus (RHBV)
Rice stripe tenuivirus rice stripe virus (RSV = RStV)
Urochloa hoja blanca tenuivirus Urochloa hoja blanca virus (UHBV)
Uukuvirus American dog uukuvirus American dog tick virus (ADAV)
Dabieshan uukuvirus Dàbiéshān tick virus (DBSH)
Grand Arbaud uukuvirus Grand Arbaud virus (GAV)
Huangpi uukuvirus Huángpí tick virus 2 (HpTV-2)
Kabuto mountain uukuvirus Kabuto mountain virus (KAMV)
Kaisodi uukuvirus Kaisodi virus (KASDV)
Lihan uukuvirus Lǐhán tick virus (LITV)
Murre uukuvirus murre virus (MURV)
Pacific coast uukuvirus Pacific coast tick virus (PACV)
Precarious Point uukuvirus Precarious Point virus (PPV)
Rukutama uukuvirus Rukutama virus (RUKV)
Schmid uukuvirus Nile Warbler virus (NIWV)
Silverwater uukuvirus Silverwater virus (SILV)
Tacheng uukuvirus Tǎchéng tick virus 2 (TCGV)
Uukuniemi uukuvirus Chizé virus (CHZV)
Fin V 707 virus (FINV)
Oceanside virus (OCV = OCEV)
Pontevès virus (PTVV)
St. Abbs Head virus (SAHV)
Uukuniemi virus (UUKV)

Taxonomic update of phylum Negarnaviricota
1 3
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, subfamilies, families, and orders are taxa
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper noun. This column lists the virus
names with their correct (lack of) capitalization. Lists of viruses within a given species are provisional at this point and will likely be amended in
the near future
Genus Species¶Virus (Abbreviation)&
Yongjia uukuvirus Yǒngjiā tick virus (YONV)
Zaliv Terpeniya uukuvirus Zaliv Terpeniya virus (ZTV)
Wenrivirus Shrimp wenrivirus Mourilyan virus (MoV)
Family Tospoviridae
Orthotospovirus Alstroemeria necrotic streak orthotospovirus Alstroemeria necrotic streak virus (ANSV)
Alstroemeria yellow spot orthotospovirus Alstroemeria yellow spot virus (AYSV)
Bean necrotic mosaic orthotospovirus bean necrotic mosaic virus (BeNMV)
Calla lily chlorotic spot orthotospovirus calla lily chlorotic spot virus (CCSV)
Capsicum chlorosis orthotospovirus Capsicum chlorosis virus (CaCV)
Chrysanthemum stem necrosis
orthotospovirus Chrysanthemum stem necrosis virus (CSNV)
Groundnut bud necrosis orthotospovirus groundnut bud necrosis virus (GBNV)
Groundnut chlorotic fan spot
orthotospovirus groundnut chlorotic fan-spot virus (GCFSV)
Groundnut ringspot orthotospovirus groundnut ringspot virus (GRSV)
Groundnut yellow spot orthotospovirus groundnut yellow spot virus (GYSV)
Hippeastrum chlorotic ringspot
orthotospovirus Hippeastrum chlorotic spot virus (HCRV)
Impatiens necrotic spot orthotospovirus impatiens necrotic spot virus (INSV)
Iris yellow spot orthotospovirus iris yellow spot virus (IYSV)
Melon severe mosaic orthotospovirus melon severe mosaic virus (MSMV)
Melon yellow spot orthotospovirus melon yellow spot virus (MYSV)
Mulberry vein banding associated
orthotospovirus mulberry vein banding-associated virus (MVBaV)
Pepper chlorotic spot orthotospovirus pepper chlorotic spot virus (PCSV)
Polygonum ringspot orthotospovirus Polygonum ringspot virus (PolRSV)
Soybean vein necrosis orthotospovirus soybean vein necrosis virus (SVNV)
Tomato chlorotic spot orthotospovirus tomato chlorotic spot virus (TCSV)
Tomato spotted wilt orthotospovirus tomato spotted wilt virus (TSWV)
Tomato yellow ring orthotospovirus tomato yellow ring virus (TYRV)
Tomato zonate spot orthotospovirus tomato zonate spot virus (TZSV)
Watermelon bud necrosis orthotospovirus watermelon bud necrosis virus (WBNV)
Watermelon silver mottle orthotospovirus watermelon silver mottle virus (WSMoV)
Zucchini lethal chlorosis orthotospovirus zucchini lethal chlorosis virus (ZLCV)
Family Tulasviridae
Orthotulasvirus Orthotulasvirus tulasnellae Tulasnella bunyavirales-like virus 1 (TB-LV)
Family Wupedeviridae
Wumivirus Wumivirus millepedae Wǔhàn millipede virus 2 (WhMV-2)
Table 6 (continued)

J. H. Kuhn etal.
1 3
Supplementary Information The online version contains supplemen-
tary material available at https:// doi. org/ 10. 1007/ s00705- 022- 05546-z.
Acknowledgements Members of the 2020–2023 International Com-
mittee on Taxonomy of Viruses (ICTV):
Piet Maes; Marco Marklewitz; Gustavo Palacios; Takahide Sasaya
(笹谷孝英); Massimo Turina; Yong-Zhen Zhang (张永振));
Filoviridae (Jens H. Kuhn, Nadine Biedenkopf; Alexander
Bukreyev; Kartik Chandran; Nicholas Di Paola; Pierre B. H. Formenty;
Anthony Griffiths; Adam J. Hume; Elke Mühlberger; Sergey V. Netesov
(Heтёcoв Cepгeй Bиктopoвич); Gustavo Palacios; Janusz T. Pawęska;
Sophie Smither; Ayato Takada (高田礼人); Victoria Wahl);
Fimoviridae (Michele Digiaro; Toufic Elbeaino; Kenji Kubota
(久保田健嗣); Francisco M. Ochoa-Corona; Susanne von Bargen);
Hantaviridae (Steven B. Bradfute; Charles H. Calisher; Boris
Klempa; Jonas Klingström; Lies Laenen; Piet Maes; Nicole D.
Tischler);
Jingchuvirales (Jens H. Kuhn; Nolwenn M. Dheilly;
Nicholas Di Paola; Sandra Junglen; Sofia Paraskevopoulou
(Σοφία Παρασκευοπούλου); Mang Shi (施莽));
Lispiviridae (Jun-Min Li (李俊敏); Sofia Paraskevopoulou
(Σοφία Παρασκευοπούλου); Gongyin Ye (叶恭银));
Monjiviricetes (Jens H. Kuhn; Nicholas Di Paola; Ralf G.
Dietzgen; W. Paul Duprex; Juliana Freitas-Astúa; Timothy H.
Hyndman; Dàohóng Jiāng (姜道宏); Piet Maes; Sofia Paraskevopoulou
(Σοφία Παρασκευοπούλου); Janusz T. Pawęska; Dennis Rubbenstroth;
Peter J. Walker; Yong-Zhen Zhang (张永振));
Table 7 ICTV-accepted
taxonomy of the
order Articulavirales
(Negarnaviricota:
Polyploviricotina:
Insthoviricetes) as of March
2022
Note that viruses are real objects that are assigned to concepts that are called taxa. Species, genera, fami-
lies, and orders are taxa
¶ Taxon names are always italicized and always begin with a capital letter
& Virus names are not italicized and are not capitalized, except if the name or a name component is a proper
noun. This column lists the virus names with their correct (lack of) capitalization
Genus Species¶Virus (Abbreviation)&
Family Amnoonviridae
Tilapinevirus Tilapinevirus tilapiae tilapia lake virus (TiLV)
Family Orthomyxoviridae
Alphainfluenzavirus Alphainfluenzavirus influenzae influenza A virus (FLUAV)
Betainfluenzavirus Betainfluenzavirus influenzae influenza B virus (FLUBV)
Deltainfluenzavirus Deltainfluenzavirus influenzae influenza D virus (FLUDV)
Gammainfluenzavirus Gammainfluenzavirus influenzae influenza C virus (FLUCV)
Isavirus Isavirus salaris infectious salmon anemia virus (ISAV)
Mykissvirus Mykissvirus tructae rainbow trout orthomyxovirus (RbtOV)
steelhead trout orthomyxovirus (SttOV-1)
Quaranjavirus Quaranjavirus araguariense Araguari virus (ARAV)
Quaranjavirus chadense Lake Chad virus (LKCV)
Quaranjavirus johnstonense Johnston Atoll virus (JAV)
Quaranjavirus quaranfilense Quaranfil virus (QRFV)
Quaranjavirus tyulekense Tyulek (Tjuloc) virus (TLKV)
Quaranjavirus wellfleetense Wellfleet Bay virus (WFBV)
Sardinovirus Sardinovirus pilchardi pilchard orthomyxovirus (POMV)
Thogotovirus Thogotovirus bourbonense Bourbon virus (BRBV)
Thogotovirus dhoriense Dhori virus (DHOV)
Thogotovirus josense Jos virus (JOSV)
Thogotovirus ozense Oz virus (OZV)
Thogotovirus sinuense Sinu virus (SINUV)
Thogotovirus thailandense Thailand tick thogotovirus (TT-THOV)
Thogotovirus thogotoense Thogoto virus (THOV)
Thogotovirus upoluense Upolu virus (UPOV)
Arenaviridae (Jens H. Kuhn; Michael J. Buchmeier; Rémi N.
Charrel; Juan Carlos de la Torre; Xavier de Lamballerie; Jean-Paul
J. Gonzalez; Stephan Günther; Jussi Hepojoki; Igor S. Lukashevich;
Sheli R. Radoshitzky; Víctor Romanowski; Manuela Sironi; Mark D.
Stenglein);
Artoviridae (Jens H. Kuhn; Ralf G. Dietzgen; Nikos Vasilakis
(Νίκος Βασιλάκης); Gongyin Ye (叶恭银); Arnfinn Lodden Økland);
Aspiviridae (Elena Dal Bó; Selma Gago-Zachert; María Laura
García; Tomohide Natsuaki (夏秋知英); José A. Navarro; Yutaro
Neriya (煉谷裕太朗); John Hammond; Vicente Pallás; Carina A.
Reyes; Takahide Sasaya (笹谷孝英); Ioannis E. Tzanetakis; Anna
Maria Vaira; Martin Verbeek);
Bornaviridae (Jens H. Kuhn; Thomas Briese; Ralf Dürrwald; Mas-
ayuki Horie (堀江真行); Timothy H. Hyndman; Norbert Nowotny;
Florian Pfaff; Dennis Rubbenstroth; Keizō Tomonaga (朝長啓造);
Ellioviricetes (Jens H. Kuhn, Scott Adkins; Juan Carlos de la Torre;
Michele Digiaro; Holly R. Hughes; Sandra Junglen; Amy J. Lambert;

Taxonomic update of phylum Negarnaviricota
1 3
Mymonaviridae (María A. Ayllón; Dàohóng Jiāng (姜道宏);
Hideki Kondō (近藤秀樹); Shin-Yi L. Marzano; Massimo Turina);
Nairoviridae (Jens H. Kuhn, Sergey V. Alkhovsky (Aльxoвcкий
Cepгeй Bлaдимиpoвич), Tatjana Avšič-Županc, Éric Bergeron; Felic-
ity Burt; Koray Ergünay; Aura R. Garrison; Marco Marklewitz; Ali
Mirazimi; Gustavo Palacios; Anna Papa (Άννα Παπά); Janusz T.
Pawęska; Jessica R. Spengler);
Negarnaviricota (Jens H. Kuhn; Eugene V. Koonin; Mart
Krupovic; Yuri I. Wolf);
Nyamiviridae (Jens H. Kuhn; Ralf G. Dietzgen; Andrew E. Firth;
Sofia Paraskevopoulou (Σοφία Παρασκευοπούλου); Nikos Vasilakis
(Νίκος Βασιλάκης));
Orthomyxoviridae (Justin Bahl; Inmaculada Casas; Adolfo García-
Sastre; Seiji Hongō (本郷誠治); Sergio H. Marshall; John W. McCau-
ley; Gabriele Neumann; Colin R. Parrish; Daniel R. Pérez; Jonathan
A. Runstadler; Martin Schwemmle);
Paramyxoviridae (Anne Balkema-Buschmann; J. Felix Drexler;
W. Paul Duprex; Piet Maes; Richard K. Plemper; Bert Vanmechelen);
Peribunyaviridae (Sergey V. Alkhovsky (Aльxoвcкий Cepгeй
Bлaдимиpoвич); Martin Beer; Carol D. Blair; Charles H. Calisher;
William M. de Souza; Holly R. Hughes; Amy J. Lambert; Marco
Marklewitz);
Phasmaviridae (Matthew J. Ballinger; Michael A. Drebot; Sandra
Junglen; Alex Pauvolid-Corrêa);
Phenuiviridae (Thomas Briese; Francesco Di Serio; Martin H. Gro-
schup; Gilda B. Jonson; Yutaro Neriya (煉谷裕太朗; Gustavo Pala-
cios; Takahide Sasaya (笹谷孝英); Jin-Won Song (송진원)); Yasuhiro
Tomitaka (冨髙保弘));
Pneumoviridae (Paul A. Brown; Ursula J. Buchholz; Rik L. de
Swart; J. Felix Drexler; W. Paul Duprex; Andrew J. Easton; Jiànróng
Lǐ (李建荣); Kirsten Spann; Natalie J. Thornburg; Bernadette van den
Hoogen; John V. Williams);
Rhabdoviridae (Nicolas Bejerman; Kim R. Blasdell; Ralf G. Dietz-
gen; Anthony R. Fooks; Juliana Freitas-Astúa; Hideki Kondō (近藤秀
樹); Ivan V. Kuzmin; Pedro L. Ramos-González; Mang Shi (施莽);
Robert B. Tesh; Noël Tordo; Nikos Vasilakis (Νίκος Βασιλάκης); Peter
J. Walker; Anna E. Whitfield);
Sunviridae (Jens H. Kuhn; Timothy H. Hyndman);
Tospoviridae (Scott Adkins; Holly R. Hughes; Rayapati Naidu;
Renato O. Resende; Massimo Turina; Anna E. Whitfield);
Xinmoviridae (Guozhong Feng (冯国忠); Sofia Paraskevopoulou
(Σοφία Παρασκευοπούλου); Stephen Sharpe) Study Groups.
2020–2023 ICTV Chair of the Animal dsRNA and ssRNA- Viruses
Subcommittee (Jens H. Kuhn); 2020–2023 ICTV Chair of the Archaeal
Viruses Subcommittee (Mart Krupovic); 2020–2023 ICTV Chair of the
Plant Viruses Subcommittee (Luisa Rubino); 2020–2023 ICTV Chair
Fungal and Protist Viruses Subcommittee (Sead Sabanadzovic); and
ICTV Proposal Secretary (Peter J. Walker)
Chair of the 2020–2023 ICTV Ellioviricetes, Filoviridae, and
Negarnaviricota Study Groups (Jens H. Kuhn); Chair of the 2020–
2023 ICTV Tospoviridae Study Group (Scott Adkins); Chair of the
2020–2023 ICTV Arenaviridae Study Group (Juan Carlos de la Torre);
Chair of the 2020–2023 ICTV Jingchuvirales Study Group (Nicholas
Di Paola); Chair of the 2020–2023 ICTV Artoviridae and Nyamiviri-
dae Study Groups (Ralf G. Dietzgen); Chair of the 2020–2023 ICTV
Fimoviridae Study Group (Michele Digiaro); Chair of the 2020–2023
ICTV Pneumoviridae Study Group (W. Paul Duprex); Co-Chair of
the 2020–2023 ICTV Rhabdoviridae Study Group (Juliana Freitas-
Astúa; Peter J. Walker); Chair of the 2020–2023 ICTV Orthomyxo-
viridae Study Group (Adolfo García-Sastre); Chair of the 2020–2023
ICTV Peribunyaviridae Study Group (Holly R. Hughes); Chair of the
2020–2023 ICTV Sunviridae Study Group (Timothy H. Hyndman);
Chair of the 2020–2023 ICTV Mymonaviridae Study Group (Dàohóng
Jiāng (姜道宏)); Chair of the 2020–2023 ICTV Phasmaviridae Study
Group (Sandra Junglen); Chair of the 2020–2023 ICTV Hantaviridae,
Monjiviricetes, and Paramyxoviridae Study Groups (Piet Maes); Chair
of the 2020–2023 ICTV Nairoviridae Study Group (Gustavo Palacios);
Co-Chair of the 2020–2023 ICTV Phenuiviridae Study Group (Gus-
tavo Palacios; Takahide Sasaya (笹谷孝英)); Resigned Chair of the
2020–2023 ICTV Paramyxoviridae Study Group (Richard K. Plemper);
Chair of the 2020–2023 ICTV Lispiviridae and Xinmoviridae Study
Groups (Sofia Paraskevopoulou (Σοφία Παρασκευοπούλου)); Chair
of the 2020–2023 ICTV Bornaviridae Study Group (Dennis Rubben-
stroth); and Chair of the 2020–2023 ICTV Aspiviridae Study Group
(Anna Maria Vaira).
Funding This work was supported in part through Laulima Govern-
ment Solutions, LLC prime contract with the US National Institute
of Allergy and Infectious Diseases (NIAID) under Contract No.
HHSN272201800013C. J.H.K. performed this work as an employee of
Tunnell Government Services (TGS), a subcontractor of Laulima Gov-
ernment Solutions, LLC under Contract No. HHSN272201800013C.
This work was also funded in part by Contract No. HSHQDC-
15-C-00064 awarded by DHS S&T for the management and operation
of The National Biodefense Analysis and Countermeasures Center,
a federally funded research and development center operated by
the Battelle National Biodefense Institute (V.W.); and NIH contract
HHSN272201000040I/HHSN27200004/D04 and grant R24AI120942
(N.V., R.B.T.). S.S. acknowledges support from the Mississippi Agri-
cultural and Forestry Experiment Station (MAFES), USDA-ARS pro-
ject 58-6066-9-033 and the National Institute of Food and Agriculture,
US Department of Agriculture, Hatch Project, under Accession Num-
ber 1021494.
Declarations
Conflict of interest The authors declare no conflicts of interest.
Ethical approval This article does not contain any studies with human
participants or animals performed by any of the authors.
Compliance with ethical standards The views and conclusions con-
tained in this document are those of the authors and should not be
interpreted as necessarily representing the official policies, either ex-
pressed or implied, of the US Department of the Army, the US De-
partment of Defense, the US Department of Health and Human Ser-
vices, the US 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. The US Government retains and the publisher, by accept-
ing the article for publication, acknowledges that the US Government
retains a non-exclusive, paid up, irrevocable, world-wide license to
publish or reproduce the published form of this manuscript, or allow
others to do so, for US Government purposes.
References
1. Allison AB, Ballard JR, Tesh RB, Brown JD, Ruder MG, Keel
MK, Munk BA, Mickley RM, Gibbs SEJ, Travassos da Rosa
APA, Ellis JC, Ip HS, Shearn-Bochsler VI, Rogers MB, Ghedin
E, Holmes EC, Parrish CR, Dwyer C (2015) Cyclic avian mass
mortality in the northeastern United States is associated with a
novel orthomyxovirus. J Virol 89:1389–1403
2. Axén C, Hakhverdyan M, Boutrup TS, Blomkvist E, Ljunghager
F, Alorden A, Hagström Å, Olesen NJ, Juremalm M, Leijon M,
Valarcher J-F (2017) Emergence of a new rhabdovirus associated

J. H. Kuhn etal.
1 3
with mass mortalities in eelpout (Zoarces viviparous) in the Baltic
Sea. J Fish Dis 40:219–229
3. Batson J, Dudas G, Haas-Stapleton E, Kistler AL, Li LM, Logan
P, Ratnasiri K, Retallack H (2021) Single mosquito metatranscrip-
tomics identifies vectors, emerging pathogens and reservoirs in
one assay. Elife 10:e68353
4. Batts WN, LaPatra SE, Katona R, Leis E, Ng TFF, Brieuc MSO,
Breyta RB, Purcell MK, Conway CM, Waltzek TB, Delwart E,
Winton JR (2017) Molecular characterization of a novel ortho-
myxovirus from rainbow and steelhead trout (Oncorhynchus
mykiss). Virus Res 230:38–49
5. Bolus S, Al Rwahnih M, Grinstead SC, Mollov D (2021) Rose
virus R, a cytorhabdovirus infecting rose. Arch Virol 166:655–658
6. Calisher CH, Karabatsos N, Zeller H, Digoutte J-P, Tesh RB,
Shope RE, Travassos da Rosa APA, St George TD (1989) Anti-
genic relationships among rhabdoviruses from vertebrates and
hematophagous arthropods. Intervirology 30:241–257
7. Chiapello M, Rodríguez-Romero J, Ayllón MA, Turina M (2020)
Analysis of the virome associated to grapevine downy mildew
lesions reveals new mycovirus lineages. Virus Evol 6:veaa058
8. Contreras-Gutiérrez MA, Nunes MRT, Guzman H, Uribe S, Suaza
Vasco JD, Cardoso JF, Popov VL, Widen SG, Wood TG, Vasilakis
N, Tesh RB (2017) Sinu virus, a novel and divergent orthomyxo-
virus related to members of the genus Thogotovirus isolated from
mosquitoes in Colombia. Virology 501:166–175
9. Da Silva EVP, Travassos Da Rosa APA, Nunes MRT, Diniz JA,
Tesh RB, Cruz ACR, Vieira CMA, Vasconcelos PFC (2005)
Araguari virus, a new member of the family Orthomyxoviridae:
serologic, ultrastructural, and molecular characterization. Am J
Trop Med Hyg 73:1050–1058
10. Di Giallonardo F, Audsley MD, Shi M, Young PR, McGraw EA,
Holmes EC (2018) Complete genome of Aedes aegypti anphevi-
rus in the Aag2 mosquito cell line. J Gen Virol 99:832–836
11. Dietzgen RG, Bejerman NE, Mei Y, Jee CLJ, Chabi-Jesus C, Fre-
itas-Astúa J, Veras SM, Kitajima EW (2021) Joá yellow blotch-
associated virus, a new alphanucleorhabdovirus from a wild sola-
naceous plant in Brazil. Arch Virol 166:1615–1622
12. Doherty RL, Whitehead RH, Wetters EJ (1968) Isolation of
viruses from Ornithodoros capensis Neumann from a tern colony
on the Great Barrier Reef, North Queensland. Aust J Sci 1968:31
13. Ejiri H, Lim C-K, Isawa H, Fujita R, Murota K, Sato T, Kobayashi
D, Kan M, Hattori M, Kimura T, Yamaguchi Y, Takayama-Ito M,
Horiya M, Posadas-Herrera G, Minami S, Kuwata R, Shimoda H,
Maeda K, Katayama Y, Mizutani T, Saijo M, Kaku K, Shinomiya
H, Sawabe K (2018) Characterization of a novel thogotovirus
isolated from Amblyomma testudinarium ticks in Ehime, Japan: a
significant phylogenetic relationship to Bourbon virus. Virus Res
249:57–65
14. El Mekki AA, Nieuwenhuysen P, van der Groen G, Pattyn SR
(1981) Characterization of some ungrouped viruses. Trans R Soc
Trop Med Hyg 75:799–806
15. Emelianchik A, Rodrigues TCS, Subramaniam K, Nielsen O,
Burek-Huntington KA, Rotstein D, Popov VL, Stone D, Waltzek
TB (2019) Characterization of a novel rhabdovirus isolated from
a stranded harbour porpoise (Phocoena phocoena). Virus Res
273:197742
16. Faizah AN, Kobayashi D, Isawa H, Amoa-Bosompem M, Murota
K, Higa Y, Futami K, Shimada S, Kim KS, Itokawa K, Watan-
abe M, Tsuda Y, Minakawa N, Miura K, Hirayama K, Sawabe
K (2020) Deciphering the virome of Culex vishnui subgroup
mosquitoes, the major vectors of Japanese encephalitis, in Japan.
Viruses 12:264
17. Fauver JR, Grubaugh ND, Krajacich BJ, Weger-Lucarelli J, Lakin
SM, Fakoli LS III, Bolay FK, Diclaro JW II, Dabiré KR, Foy
BD, Brackney DE, Ebel GD, Stenglein MD (2016) West African
Anopheles gambiae mosquitoes harbor a taxonomically diverse
virome including new insect-specific flaviviruses, mononegavi-
ruses, and totiviruses. Virology 498:288–299
18. Forgia M, Isgandarli E, Aghayeva DN, Huseynova I, Turina M
(2021) Virome characterization of Cryphonectria parasitica iso-
lates from Azerbaijan unveiled a new mymonavirus and a putative
new RNA virus unrelated to described viral sequences. Virology
553:51–61
19. Godwin SE, Morrison RN, Knowles G, Cornish MC, Hayes D,
Carson J (2020) Pilchard orthomyxovirus (POMV). II. Causative
agent of salmon orthomyxoviral necrosis, a new disease of farmed
Atlantic salmon Salmo salar. Dis Aquat Organ 139:51–68
20. Hierweger MM, Koch MC, Rupp M, Maes P, Di Paola N, Brug-
gmann R, Kuhn JH, Schmidt-Posthaus H, Seuberlich T (2021)
Novel filoviruses, hantavirus, and rhabdovirus in freshwater fish,
Switzerland, 2017. Emerg Infect Dis 27:3082–3091
21. Horie M (2021) Identification of a novel filovirus in a common
lancehead (Bothrops atrox (Linnaeus, 1758)). J Vet Med Sci
83:1485–1488
22. Horie M, Akashi H, Kawata M, Tomonaga K (2021) Identification
of a reptile lyssavirus in Anolis allogus provided novel insights
into lyssavirus evolution. Virus Genes 57:40–49
23. International Committee on Taxonomy of Viruses (ICTV) (2022)
Approved Proposals. https:// ictv. global/ files/ propo sals/ appro ved
24. International Committee on Taxonomy of Viruses (ICTV) (2022)
Virus Taxonomy: 2021 Release (MSL #37). https:// ictv. global/
news/ news_ 082322
25. International Committee on Taxonomy of Viruses (ICTV) (2022)
ICTV Code. The International Code of Virus Classification and
Nomenclature (ICVCN). 2021. https:// ictv. global/ about/ code
26. Jia J, Fu Y, Jiang D, Mu F, Cheng J, Lin Y, Li B, Marzano S-YL,
Xie J (2021) Interannual dynamics, diversity and evolution of the
virome in Sclerotinia sclerotiorum from a single crop field. Virus
Evol 7:veab032
27. Käfer S, Paraskevopoulou S, Zirkel F, Wieseke N, Donath A,
Petersen M, Jones TC, Liu S, Zhou X, Middendorf M, Junglen S,
Misof B, Drosten C (2019) Re-assessing the diversity of negative
strand RNA viruses in insects. PLoS Pathog 15:e1008224
28. Kobayashi D, Komatsu N, Faizah AN, Amoa-Bosompem M,
Sawabe K, Isawa H (2021) A novel nyavirus lacking matrix
and glycoprotein genes from Argas japonicus ticks. Virus Res
292:198254
29. Kopp A, Hübner A, Zirkel F, Hobelsberger D, Estrada A, Jordan
I, Gillespie TR, Drosten C, Junglen S (2019) Detection of two
highly diverse peribunyaviruses in mosquitoes from Palenque.
Mexico. Viruses 11:832
30. Kosoy OI, Lambert AJ, Hawkinson DJ, Pastula DM, Goldsmith
CS, Hunt DC, Staples JE (2015) Novel thogotovirus associated
with febrile illness and death, United States, 2014. Emerg Infect
Dis 21:760–764
31. Krishnamurthy SR (2017) The reduction of viral dark matter by
expanding of viral diversity. In: PhD dissertation, Washington
University, Saint Louis, Missouri, USA
32. Kubota K, Yanagisawa H, Chiaki Y, Yamasaki J, Horikawa H,
Tsunekawa K, Morita Y, Kadono F (2021) Complete nucleotide
sequence of chrysanthemum mosaic-associated virus, a novel
emaravirus infecting chrysanthemum. Arch Virol 166:1241–1245
33. Kuhn JH, Wolf YI, Krupovic M, Zhang Y-Z, Maes P, Dolja VV,
Koonin EV (2019) Classify viruses—the gain is worth the pain.
Nature 566:318–320
34. Kuhn JH, Adkins S, Agwanda BR, Al Kubrusli R, Alkhovsky
SV, Amarasinghe GK, Tatjana AŽ, Ayllón MA, Bahl J, Balkema-
Buschmann A, Ballinger MJ, Basler CF, Bavari S, Beer M, Bejer-
man N, Bennett AJ, Bente DA, Bergeron É, Bird BH, Blair CD,
Blasdell KR, Blystad D-R, Bojko J, Borth WB, Bradfute S, Breyta
R, Briese T, Brown PA, Brown JK, Buchholz UJ, Buchmeier MJ,
Bukreyev A, Burt F, Büttner C, Calisher CH, Cao M, Casas I,

Taxonomic update of phylum Negarnaviricota
1 3
Chandran K, Charrel RN, Cheng Q, Chiaki Y, Chiapello M, Choi
I-R, Ciuffo M, Clegg JCS, Crozier I, Dal Bó E, de la Torre JC,
de Lamballerie X, de Swart RL, Debat H, Dheilly NM, Di Cicco
E, Di Paola N, Di Serio F, Dietzgen RG, Digiaro M, Dolnik O,
Drebot MA, Drexler JF, Dundon WG, Duprex WP, Dürrwald R,
Dye JM, Easton AJ, Ebihara H, Elbeaino T, Ergünay K, Fergu-
son HW, Fooks AR, Forgia M, Formenty PBH, Fránová J, Frei-
tas-Astúa J, Fu J, Fürl S, Gago-Zachert S, Gāo GF, García ML,
García-Sastre A, Garrison AR, Gaskin T, Gonzalez J-PJ, Griffiths
A, Goldberg TL, Groschup MH, Günther S, Hall RA, Hammond J,
Han T, Hepojoki J, Hewson R, Hong J, Hong N, Hongo S, Horie
M, Hu JS, Hu T, Hughes HR, Hüttner F, Hyndman TH, Ilyas M,
Jalkanen R, Jiāng D, Jonson GB, Junglen S, Kadono F, Kaukinen
KH, Kawate M, Klempa B, Klingström J, Kobinger G, Koloniuk
I, Kondō H, Koonin EV, Krupovic M, Kubota K, Kurath G, Lae-
nen L, Lambert AJ, Langevin SL, Lee B, Lefkowitz EJ, Leroy
EM, Li S, Li L, Lǐ J, Liu H, Lukashevich IS, Maes P, de Souza
WM, Marklewitz M, Marshall SH, Marzano S-YL, Massart S,
McCauley JW, Melzer M, Mielke-Ehret N, Miller KM, Ming TJ,
Mirazimi A, Mordecai GJ, Mühlbach H-P, Mühlberger E, Naidu
R, Natsuaki T, Navarro JA, Netesov SV, Neumann G, Nowotny
N, Nunes MRT, Olmedo-Velarde A, Palacios G, Pallás V, Pályi
B, Papa A, Paraskevopoulou S, Park AC, Parrish CR, Patterson
DA, Pauvolid-Corrêa A, Pawęska JT, Payne S, Peracchio C, Pérez
DR, Postler TS, Qi L, Radoshitzky SR, Resende RO, Reyes CA,
Rima BK, Luna GR, Romanowski V, Rota P, Rubbenstroth D,
Rubino L, Runstadler JA, Sabanadzovic S, Sall AA, Salvato MS,
Sang R, Sasaya T, Schulze AD, Schwemmle M, Shi M, Shí X,
Shí Z, Shimomoto Y, Shirako Y, Siddell SG, Simmonds P, Sironi
M, Smagghe G, Smither S, Song J-W, Spann K, Spengler JR,
Stenglein MD, Stone DM, Sugano J, Suttle CA, Tabata A, Takada
A, Takeuchi S, Tchouassi DP, Teffer A, Tesh RB, Thornburg NJ,
Tomitaka Y, Tomonaga K, Tordo N, Torto B, Towner JS, Tsuda
S, Tu C, Turina M, Tzanetakis IE, Uchida J, Usugi T, Vaira AM,
Vallino M, van den Hoogen B, Varsani A, Vasilakis N, Verbeek
M, von Bargen S, Wada J, Wahl V, Walker PJ, Wang L-F, Wang
G, Wang Y, Wang Y, Waqas M, Wèi T, Wen S, Whitfield AE,
Williams JV, Wolf YI, Wu J, Xu L, Yanagisawa H, Yang C, Yang
Z, Zerbini FM, Zhai L, Zhang Y-Z, Zhang S, Zhang J, Zhang
Z, Zhou X (2021) 2021 Taxonomic update of phylum Negarna-
viricota (Riboviria: Orthornavirae), including the large orders
Bunyavirales and Mononegavirales. Arch Virol 166:3513–3566
35. Lay CL, Shi M, Buček A, Bourguignon T, Lo N, Holmes EC
(2020) Unmapped RNA virus diversity in termites and their sym-
bionts. Viruses 12:1145
36. Lee VH, Kemp GE, Madbouly MH, Moore DL, Causey OR,
Casals J (1974) Jos, a new tick-borne virus from Nigeria. Am J
Vet Res 35:1165–1167
37. Liu Q, Jin J, Yang L, Zhang S, Cao M (2021) Molecular charac-
terization of a novel cytorhabdovirus associated with chrysanthe-
mum yellow dwarf disease. Arch Virol 166:1253–1257
38. Liu W, Fan Y, Li Z, Zhao J, Zhou Y, Jiang N, Zeng J, Cain K,
Zeng L (2019) Isolation, identification, and classification of a
novel rhabdovirus from diseased Chinese rice-field eels (Monop-
terus albus). Arch Virol 164:105–116
39. Luo DS, Li B, Shen X-R, Jiang R-D, Zhu Y, Wu J, Fan Y, Bourhy
H, Hu B, Ge X-Y, Shi Z-L, Dacheux L (2021) Characterization of
novel rhabdoviruses in Chinese bats. Viruses 13:64
40. Lyu S-j, Yuan X-m, Zhang H-q, Shi W-d, Hang X-y, Liu L, Wu
Y-l (2019) Isolation and characterization of a novel strain (YH01)
of Micropterus salmoides rhabdovirus and expression of its gly-
coprotein by the baculovirus expression system. J Zhejiang Univ
Sci B 20:728–739
41. Mohr PG, Crane MSJ, Hoad J, Williams LM, Cummins D, Neave
MJ, Shiell B, Beddome G, Michalski WP, Peck GR, Samsing F,
Wynne JW, Crameri SG, Hyatt AD, Moody NJG (2020) Pilchard
orthomyxovirus (POMV). I. Characterisation of an emerging virus
isolated from pilchards Sardinops sagax and Atlantic salmon
Salmo salar. Dis Aquat Organ 139:35–50
42. Nerva L, Forgia M, Ciuffo M, Chitarra W, Chiapello M, Vallino
M, Varese GC, Turina M (2019) The mycovirome of a fungal
collection from the sea cucumber Holothuria polii. Virus Res
273:197737
43. Öhlund P, Hayer J, Lundén H, Hesson JC, Blomström AL (2019)
Viromics reveal a number of novel RNA viruses in Swedish mos-
quitoes. Viruses 11:1027
44. Orfanidou CG, Beta C, Reynard J-S, Tsiolakis G, Katis NI, Mali-
ogka VI (2020) Identification, molecular characterization and
prevalence of a novel cytorhabdovirus infecting zucchini crops in
Greece. Virus Res 287:198095
45. Osterhaus ADME, Broeders HWJ, Teppema JS, Kuiken T, House
JA, Vos HW, Visser IKG (1993) Isolation of a virus with rhab-
dovirus morphology from a white-beaked dolphin (Lagenorhyn-
chus albirostris). Arch Virol 133:189–193
46. Ottati S, Chiapello M, Galetto L, Bosco D, Marzachì C, Abbà S
(2020) New viral sequences identified in the flavescence dorée
phytoplasma vector Scaphoideus titanus. Viruses 12:287
47. Pallandre L, Luo D, Feuvrier C, Lieffrig F, Pozet F, Dacheux L,
Bigarré L (2020) Revisiting the classification of percid perhab-
doviruses using new full-length genomes. Viruses 12:649
48. Parry R, Asgari S (2018) Aedes anphevirus: an insect-specific
virus distributed worldwide in Aedes aegypti mosquitoes that has
complex interplays with Wolbachia and dengue virus infection in
cells. J Virol 92:e00224-18
49. Pfaff F, Rubbenstroth D (2021) Two novel bornaviruses identified
in colubrid and viperid snakes. Arch Virol 166:2611–2614
50. Presti RM, Zhao G, Beatty WL, Mihindukulasuriya KA, Travassos
da Rosa APA, Popov VL, Tesh RB, Virgin HW, Wang D (2009)
Quaranfil, Johnston Atoll, and Lake Chad viruses are novel mem-
bers of the family Orthomyxoviridae. J Virol 83:11599–11606
51. Qiu Y, Zhang S, Jin J, Xie J, Cao Y, Cao M (2020) Molecular
characterization of a novel cytorhabdovirus associated with paper
mulberry mosaic disease. Arch Virol 165:2703–2707
52. Rehanek M, von Bargen S, Bandte M, Karlin DG, Büttner C
(2021) A novel emaravirus comprising five RNA segments is
associated with ringspot disease in oak. Arch Virol 166:987–990
53. Ruiz-Padilla A, Rodríguez-Romero J, Gómez-Cid I, Pacifico D,
Ayllón MA (2021) Novel mycoviruses discovered in the mycovi-
rome of a necrotrophic fungus. MBio 12:e03705-20
54. Rumbou A, Candresse T, von Bargen S, Büttner C (2020)
Next-generation sequencing reveals a novel emaravirus in dis-
eased maple trees from a German urban forest. Front Microbiol
11:621179
55. Sakaguchi S, Nakagawa S, Mitsuhashi S, Ogawa M, Sugiyama K,
Tamukai K, Koide R, Katayama Y, Nakano T, Makino S, Imanishi
T, Miyazawa T, Mizutani T (2020) Molecular characterization
of feline paramyxovirus in Japanese cat populations. Arch Virol
165:413–418
56. Scarpassa VM, Debat HJ, Alencar RB, Saraiva JF, Calvo E, Arcà
B, Ribeiro JMC (2019) An insight into the sialotranscriptome and
virome of Amazonian anophelines. BMC Genomics 20:166
57. Shi M, Neville P, Nicholson J, Eden J-S, Imrie A, Holmes EC
(2017) High-resolution metatranscriptomics reveals the ecologi-
cal dynamics of mosquito-associated RNA viruses in Western
Australia. J Virol 91:e00680-17
58. Shi M, Lin X-D, Chen X, Tian J-H, Chen L-J, Li K, Wang W, Eden
J-S, Shen J-J, Liu L, Holmes EC, Zhang Y-Z (2018) The evolu-
tionary history of vertebrate RNA viruses. Nature 556:197–202
59. Siddell SG, Walker PJ, Lefkowitz EJ, Mushegian AR, Adams
MJ, Dutilh BE, Gorbalenya AE, Bz H, Harrison RL, Junglen S,
Knowles NJ, Kropinski AM, Krupovic M, Kuhn JH, Nibert M,
Rubino L, Sabanadzovic S, Sanfaçon H, Simmonds P, Varsani A,

J. H. Kuhn etal.
1 3
Zerbini FM, Davison AJ (2019) Additional changes to taxonomy
ratified in a special vote by the International Committee on Tax-
onomy of Viruses (October 2018). Arch Virol 164:943–946
60. Sidharthan VK, Baranwal VK (2021) Mining of the water hyssop
(Bacopa monnieri) transcriptome reveals genome sequences of
two putative novel rhabdoviruses and a solendovirus. Arch Virol
166:1985–1990
61. Sieg M, Heenemann K, Rückner A, Burgener I, Oechtering G,
Vahlenkamp TW (2015) Discovery of new feline paramyxovi-
ruses in domestic cats with chronic kidney disease. Virus Genes
51:294–297
62. Sutela S, Forgia M, Vainio EJ, Chiapello M, Daghino S, Vallino
M, Martino E, Girlanda M, Perotto S, Turina M (2020) The virome
from a collection of endomycorrhizal fungi reveals new viral taxa
with unprecedented genome organization. Virus Evol 6:veaa076
63. Temmam S, Chrétien D, Bigot T, Dufour E, Petres S, Desquesnes
M, Devillers E, Dumarest M, Yousfi L, Jittapalapong S, Karn-
chanabanthoeng A, Chaisiri K, Gagnieur L, Cosson JF, Vayssier-
Taussat M, Morand S, Moutailler S, Eloit M (2019) Monitoring
silent spillovers before emergence: a pilot study at the tick/human
interface in Thailand. Front Microbiol 10:2315
64. Těšíková J, Krásová J, Goüy de Bellocq J (2021) Multiple mam-
marenaviruses circulating in Angolan rodents. Viruses 13:982
65. Vainio EJ, Sutela S (2020) Mixed infection by a partitivirus and a
negative-sense RNA virus related to mymonaviruses in the poly-
pore fungus Bondarzewia berkeleyi. Virus Res 286:198079
66. Vanmechelen B, Vergote V, Merino M, Verbeken E, Maes P
(2020) Common occurrence of Belerina virus, a novel paramyxo-
virus found in Belgian hedgehogs. Sci Rep 10:19341
67. Vanmechelen B, Meurs S, Zisi Z, Goüy de Bellocq J, Bletsa M,
Lemey P, Maes P (2021) Genome sequence of ruloma Virus, a
novel paramyxovirus clustering basally to members of the genus
Jeilongvirus. Microbiol Resour Announc 10:e00325-21
68. Viljakainen L, Holmberg I, Abril S, Jurvansuu J (2018) Viruses
of invasive Argentine ants from the European Main supercol-
ony: characterization, interactions and evolution. J Gen Virol
99:1129–1140
69. Walker PJ, Siddell SG, Lefkowitz EJ, Mushegian AR, Adriaens-
sens EM, Alfenas-Zerbini P, Davison AJ, Dempsey DM, Dutilh
BE, García ML, Harrach B, Harrison RL, Hendrickson RC, Jun-
glen S, Knowles NJ, Krupovic M, Kuhn JH, Lambert AJ, Łobocka
M, Nibert ML, Oksanen HM, Orton RJ, Robertson DL, Rubino L,
Sabanadzovic S, Simmonds P, Smith DB, Suzuki N, Van Dooer-
slaer K, Vandamme AM, Varsani A, Zerbini FM (2021) Changes
to virus taxonomy and to the International Code of Virus Classi-
fication and Nomenclature ratified by the International Committee
on Taxonomy of Viruses (2021). Arch Virol 166:2633–2648
70. Walker PJ, Siddell SG, Lefkowitz EJ, Mushegian AR, Adriaens-
sens EM, Alfenas-Zerbini P, Dempsey DM, Dutilh BE, García
ML, Hendrickson RC, Junglen S, Krupovic M, Kuhn JH, Lambert
AJ, Łobocka M, Oksanen HM, Orton RJ, Robertson DL, Rubino
L, Sabanadzovic S, Simmonds P, Smith DB, Suzuki N, Van Doo-
erslaer K, Vandamme A-M, Varsani A, Zerbini FM (2022) Recent
changes to virus taxonomy ratified by the International Commit-
tee on Taxonomy of Viruses (2022). Arch Virol 167:2429–2440.
https:// doi. org/ 10. 1007/ s00705- 022- 05516-5.
71. Waltzek TB, Subramaniam K, Leis E, Katona R, Fan Ng TF,
Delwart E, Barbknecht M, Rock K, Hoffman MA (2019) Char-
acterization of a peribunyavirus isolated from largemouth bass
(Micropterus salmoides). Virus Res 273:197761
72. Wang F, Yuan B, Xiao S, Zhang J, Jia W, Fang Q, Wang F,
Song Q, Ye G (2021) Diverse RNA viruses discovered in three
parasitoid wasps of the rice weevil Sitophilus oryzae. mSphere
6:e00331-21
73. Williams SH, Che X, Oleynik A, Garcia JA, Muller D, Zabka
TS, Firth C, Corrigan RM, Briese T, Jain K, Lipkin WI (2019)
Discovery of two highly divergent negative-sense RNA viruses
associated with the parasitic nematode, Capillaria hepatica, in wild
Mus musculus from New York City. J Gen Virol 100:1350–1362
74. Wolf YI, Kazlauskas D, Iranzo J, Lucía-Sanz A, Kuhn JH,
Krupovic M, Dolja VV, Koonin EV (2018) Origins and evolution
of the global RNA virome. MBio 9:e02329-18
75. Wu Z, Yang L, Ren X, He G, Zhang J, Yang J, Qian Z, Dong J,
Sun L, Zhu Y, Du J, Yang F, Zhang S, Jin Q (2016) Deciphering
the bat virome catalog to better understand the ecological diversity
of bat viruses and the bat origin of emerging infectious diseases.
ISME J 10:609–620
76. Wu Z, Lu L, Du J, Yang L, Ren X, Liu B, Jiang J, Yang J, Dong J,
Sun L, Zhu Y, Li Y, Zheng D, Zhang C, Su H, Zheng Y, Zhou H,
Zhu G, Li H, Chmura A, Yang F, Daszak P, Wang J, Liu Q, Jin Q
(2018) Comparative analysis of rodent and small mammal viromes
to better understand the wildlife origin of emerging infectious
diseases. Microbiome 6:178
77. Xavier CAD, Allen ML, Whitfield AE (2021) Ever-increasing
viral diversity associated with the red imported fire ant Solenopsis
invicta (Formicidae: Hymenoptera). Virol J 18:5
78. Yang S, Shan T, Wang Y, Yang J, Chen X, Xiao Y, You Z, He Y,
Zhao M, Lu J, Yang Z, Dai Z, Liu Q, Yao Y, Lu X, Li H, Zhou R,
Li W, Zhou C, Wang X, Shen Q, Xu H, Deng X, Delwart E, Zhang
W (2020) Virome of riverside phytocommunity ecosystem of an
ancient canal. ResearchSquare:https:// www. resea rchsq uare. com/
artic le/ rs- 25620/ v1
79. Zeller HG, Karabatsos N, Calisher CH, Digoutte J-P, Murphy FA,
Shope RE (1989) Electron microscopy and antigenic studies of
uncharacterized viruses. I. Evidence suggesting the placement of
viruses in families Arenaviridae, Paramyxoviridae, or Poxviridae.
Arch Virol 108:191–209
80. Zeng W, Wang Q, Wang Y, Liu C, Liang H, Fang X, Wu S (2014)
Genomic characterization and taxonomic position of a rhabdovi-
rus from a hybrid snakehead. Arch Virol 159:2469–2473
81. Zerbini FM, Siddell SG, Mushegian AR, Walker PJ, Lefkowitz EJ,
Adriaenssens EM, Alfenas-Zerbini P, Dutilh BE, García ML, Jun-
glen S, Krupovic M, Kuhn JH, Lambert AJ, Łobocka M, Oksanen
HM, Robertson DL, Rubino L, Sabanadzovic S, Simmonds P,
Suzuki N, Van Doorslaer K, Vandamme AM, Varsani A (2022)
Differentiating between viruses and virus species by writing their
names correctly. Arch Virol 167:1231–1234
82. Zhang Q-Y, Tao J-J, Gui L, Zhou G-Z, Ruan H-M, Li Z-Q, Gui J-F
(2007) Isolation and characterization of Scophthalmus maximus
rhabdovirus. Dis Aquat Organ 74:95–105
83. Aльxoвcкий CB, Щeтинин AM, Львoв ДК, Щeлкaнoв MЮ,
Дepябин ПГ, Львoв ДH, Caмoxвaлoв EИ, Гитeльмaн AК,
Бoтикoв AГ (2013) Bиpyc Xypдyн (KHURV): нoвый виpyc
poдa Orthobunyavirus (Bunyaviridae). Boпp виpycoл 58:10–13
84. Гaлкинa ИB, Львoв ДH, Гpoмaшeвcкий BЛ, Mocквинa TM
(2005) Bиpyc Xypдyн (Khurdun), пpeдпoлoжитeльнo нoвый
PHК-coдepжaщий виpyc, cвязaнный c лыcyxaм и (Fulica atra),
изoлиpoвaнный в дeльтe Boлги. Boпp виpycoл 50:29–31
85. Кapacь ФP (1979) Экoлoгия apбoвиpycoв гopнoй cиcтeмы
Cpeднeaзиaтcкoгo peгиoнa CCCP. Aвтopeфepaт диccepтaции
нa coиcкaниe yчeнoй cтeпeни дoктopa мeдицинcкиx нayк.
Mocквa, PCФCP, CCCP
86. 张奇亚、李正秋 (1999) 在患病鳜鱼组织中观察到3种病毒。
科学通报 44:192–195
87. 张奇亚、陶建军、黄晓红、李正秋、袁秀平 (2005) 两种鳜病
毒的共感染现象及引起感染细胞的超微变化。水生生物学报
29:473–478
Publisher's Note Springer Nature remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.

Taxonomic update of phylum Negarnaviricota
1 3
Authors and Aliations
JensH.Kuhn1 · ScottAdkins2 · SergeyV.Alkhovsky(Aльxoвcкий Cepгeй Bлaдимиpoвич)3 ·
TatjanaAvšič‑Županc4 · MaríaA.Ayllón5,6 · JustinBahl7 · AnneBalkema‑Buschmann8 ·
MatthewJ.Ballinger9 · MartinaBandte10 · MartinBeer11 · NicolasBejerman12 · ÉricBergeron13 ·
NadineBiedenkopf14 · LaurentBigarré15 · CarolD.Blair16 · KimR.Blasdell17 · StevenB.Bradfute18 ·
ThomasBriese19 · PaulA.Brown20 · RémyBruggmann21 · UrsulaJ.Buchholz22 · MichaelJ.Buchmeier23 ·
AlexanderBukreyev24,91 · FelicityBurt25 · CarmenBüttner10 · CharlesH.Calisher26· ThierryCandresse27 ·
JeremyCarson28 · InmaculadaCasas29 · KartikChandran30 · RémiN.Charrel31 · YuyaChiaki
(千秋祐也)32 · AnyaCrane33 · MarkCrane34· LaurentDacheux35 · ElenaDalBó36· JuanCarlosdelaTorre37 ·
XavierdeLamballerie38· WilliamM.deSouza39 · RikL.deSwart40 · NolwennM.Dheilly41 ·
NicholasDiPaola42 · FrancescoDiSerio43 · RalfG.Dietzgen44 · MicheleDigiaro45· J.FelixDrexler46 ·
W.PaulDuprex47 · RalfDürrwald48 · AndrewJ.Easton49 · ToucElbeaino45 · KorayErgünay50,51,52,53 ·
GuozhongFeng(冯国忠)54 · ClaudetteFeuvrier55· AndrewE.Firth56 · AnthonyR.Fooks57 ·
PierreB.H.Formenty58 · JulianaFreitas‑Astúa59 · SelmaGago‑Zachert60 · MaríaLauraGarcía61 ·
AdolfoGarcía‑Sastre62 · AuraR.Garrison63 · ScottE.Godwin28 · Jean‑PaulJ.Gonzalez64 ·
JoëlleGoüydeBellocq65 · AnthonyGriths66 · MartinH.Groschup67 · StephanGünther68 ·
JohnHammond69 · JussiHepojoki70,71 · MelanieM.Hierweger72· SeijiHongō(本郷誠治)73·
MasayukiHorie(堀江真行)74,75 · HidenoriHorikawa(堀川英則)76· HollyR.Hughes77 · AdamJ.Hume78,79 ·
TimothyH.Hyndman80 · DàohóngJiāng(姜道宏)81 · GildaB.Jonson82 · SandraJunglen46 · FujioKadono
(上遠野冨士夫)83 · DavidG.Karlin10 · BorisKlempa84 · JonasKlingström85 · MichelC.Koch86 ·
HidekiKondō(近藤秀樹)87 · EugeneV.Koonin88· JarmilaKrásová65· MartKrupovic89 · KenjiKubota
(久保田健嗣)90 · IvanV.Kuzmin91 · LiesLaenen92,93 · AmyJ.Lambert94 · JiànróngLǐ(李建荣)95·
Jun‑MinLi(李俊敏)96 · FrançoisLierig97· IgorS.Lukashevich98 · DongshengLuo99,100· PietMaes101 ·
MarcoMarklewitz102 · SergioH.Marshall103 · Shin‑YiL.Marzano104 · JohnW.McCauley105 ·
AliMirazimi106· PeterG.Mohr34 · NickJ.G.Moody34· YasuakiMorita(森田泰彰)107· RichardN.Morrison28 ·
ElkeMühlberger108 · RayapatiNaidu109 · TomohideNatsuaki(夏秋知英)110 · JoséA.Navarro111 ·
YutaroNeriya(煉谷裕太朗)112 · SergeyV.Netesov(Heтёcoв Cepгeй Bиктopoвич)113 · GabrieleNeumann114 ·
NorbertNowotny115,116 · FranciscoM.Ochoa‑Corona117 · GustavoPalacios118 · LauranePallandre119·
VicentePallás120 · AnnaPapa(Άννα Παπά)121 · SoaParaskevopoulou(Σοφία Παρασκευοπούλου)122 ·
ColinR.Parrish123· AlexPauvolid‑Corrêa124 · JanuszT.Pawęska125 · DanielR.Pérez126 · FlorianPfa55·
RichardK.Plemper127 · ThomasS.Postler128 · FrançoisePozet55· SheliR.Radoshitzky129 ·
PedroL.Ramos‑González130 · MariusRehanek10 · RenatoO.Resende131 · CarinaA.Reyes132 ·
VíctorRomanowski133 · DennisRubbenstroth134 · LuisaRubino43 · ArtemisRumbou(Άρτεμις Ρούμπου)10 ·
JonathanA.Runstadler135 · MelanieRupp136· SeadSabanadzovic137 · TakahideSasaya
(笹谷孝英)138 · HeikeSchmidt‑Posthaus139 · MartinSchwemmle140 · TorstenSeuberlich141 ·
StephenR.Sharpe142 · MangShi(施莽)143 · ManuelaSironi144 · SophieSmither145 · Jin‑WonSong
(송진원)146 · KirstenM.Spann147 · JessicaR.Spengler148 · MarkD.Stenglein149 · AyatoTakada
(高田礼人)150 · RobertB.Tesh91· JanaTěšíková65,151· NatalieJ.Thornburg152 · NicoleD.Tischler153 ·
YasuhiroTomitaka(冨髙保弘)154 · KeizōTomonaga(朝長啓造)155 · NoëlTordo156 · KentaTsunekawa157·
MassimoTurina158 · IoannisE.Tzanetakis159 · AnnaMariaVaira158 · BernadettevandenHoogen160 ·
BertVanmechelen161 · NikosVasilakis(Νίκος Βασιλάκης)91 · MartinVerbeek162 · SusannevonBargen10 ·
JiroWada(和田治郎)163 · VictoriaWahl164 · PeterJ.Walker165 · AnnaE.Whiteld166 · JohnV.Williams167 ·
YuriI.Wolf168 · JunkiYamasaki(山﨑淳紀)169· HironobuYanagisawa(栁澤広宣)170 · GongyinYe(叶恭银)171 ·
Yong‑ZhenZhang(张永振)172· ArnnnLoddenØkland173
* Jens H. Kuhn
kuhnjens@mail.nih.gov
Scott Adkins
scott.adkins@ars.usda.gov
Sergey V. Alkhovsky (Aльxoвcкий Cepгeй Bлaдимиpoвич)
salkh@yandex.ru
Tatjana Avšič-Županc
Tatjana.Avsic@mf.uni-lj.si
María A. Ayllón
mariaangeles.ayllon@upm.es
Justin Bahl
justin.bahl@uga.edu

J. H. Kuhn etal.
1 3
Anne Balkema-Buschmann
anne.balkema-buschmann@fli.de
Matthew J. Ballinger
ballinger@biology.msstate.edu
Martina Bandte
martina.bandte@agrar.hu-berlin.de
Martin Beer
martin.beer@fli.de
Nicolas Bejerman
bejerman.nicolas@inta.gob.ar
Éric Bergeron
exj8@cdc.gov
Nadine Biedenkopf
nadine.biedenkopf@staff.uni-marburg.de
Laurent Bigarré
laurent.bigarre@anses.fr
Carol D. Blair
carol.blair@colostate.edu
Kim R. Blasdell
kim.blasdell@csiro.au
Steven B. Bradfute
sbradfute@salud.unm.edu
Thomas Briese
thomas.briese@columbia.edu
Paul A. Brown
paul.brown@anses.fr
Rémy Bruggmann
remy.bruggmann@bioinformatics.unibe.ch
Ursula J. Buchholz
ubuchholz@niaid.nih.gov
Michael J. Buchmeier
m.buchmeier@uci.edu
Alexander Bukreyev
alexander.bukreyev@utmb.edu
Felicity Burt
burt@ufs.ac.za
Carmen Büttner
carmen.buettner@agrar.hu-berlin.de
Charles H. Calisher
calisher6920@gmail.com
Thierry Candresse
thierry.candresse@inrae.fr
Jeremy Carson
jeremy.carson@bioconsulting.com.au
Inmaculada Casas
icasas@isciii.es
Kartik Chandran
kartik.chandran@einstein.yu.edu
Rémi N. Charrel
remi.charrel@univ-amu.fr
Yuya Chiaki (千秋祐也)
chiakiy151@affrc.go.jp
Anya Crane
anya.crane@nih.gov
Mark Crane
Mark.Crane@csiro.au
Laurent Dacheux
laurent.dacheux@pasteur.fr
Elena Dal Bó
elenadalbocastanion@gmail.com
Juan Carlos de la Torre
juanct@scripps.edu
Xavier de Lamballerie
xavier.de-lamballerie@univ-amu.fr
William M. de Souza
wmarciel@hotmail.com
Rik L. de Swart
rik.deswart@wur.nl
Nolwenn M. Dheilly
nolwenn.dheilly@anses.fr
Nicholas Di Paola
nicholas.dipaola.civ@health.mil
Francesco Di Serio
francesco.diserio@ipsp.cnr.it
Ralf G. Dietzgen
r.dietzgen@uq.edu.au
Michele Digiaro
digiaro@iamb.it
J. Felix Drexler
felix.drexler@charite.de
W. Paul Duprex
pduprex@pitt.edu
Ralf Dürrwald
duerrwaldr@rki.de
Andrew J. Easton
A.J.Easton@warwick.ac.uk
Toufic Elbeaino
elbeaino@iamb.it
Koray Ergünay
ekoray@hacettepe.edu.tr
Guozhong Feng (冯国忠)
fengguozhong@caas.cn
Claudette Feuvrier
cfeuvrier@jura.fr
Andrew E. Firth
aef24@cam.ac.uk
Anthony R. Fooks
Tony.Fooks@apha.gov.uk
Pierre B. H. Formenty
formentyp@who.int
Juliana Freitas-Astúa
juliana.astua@embrapa.br
Selma Gago-Zachert
selma.gago-zachert@bct.uni-halle.de

Taxonomic update of phylum Negarnaviricota
1 3
María Laura García
marialauragarcia2010@gmail.com
Adolfo García-Sastre
Adolfo.Garcia-Sastre@mssm.edu
Aura R. Garrison
aura.r.garrison.civ@health.mil
Scott E. Godwin
Scott.Godwin@nre.tas.gov.au
Jean-Paul J. Gonzalez
Jean.Paul.Gonzalez@georgetown.edu
Joëlle Goüy de Bellocq
joellegouy@gmail.com
Anthony Griffiths
ahgriff@bu.edu
Martin H. Groschup
Martin.Groschup@fli.de
Stephan Günther
guenther@bni.uni-hamburg.de
John Hammond
John.Hammond@usda.gov
Jussi Hepojoki
jussi.hepojoki@helsinki.fi; jussi.hepojoki@uzh.ch
Melanie M. Hierweger
melanie_hierweger@web.de
Seiji Hongō (本郷誠治)
shongou@med.id.yamagata-u.ac.jp
Masayuki Horie (堀江真行)
mhorie@omu.ac.jp
Hidenori Horikawa (堀川英則)
hidenori.horikawa@pref.aichi.lg.jp
Holly R. Hughes
ltr8@cdc.gov
Adam J. Hume
hume@bu.edu
Timothy H. Hyndman
T.Hyndman@murdoch.edu.au
Dàohóng Jiāng (姜道宏)
daohongjiang@mail.hzau.edu.cn
Gilda B. Jonson
g.jonson@irri.org
Sandra Junglen
sandra.junglen@charite.de
Fujio Kadono (上遠野冨士夫)
acaritick@yahoo.co.jp
David G. Karlin
davidgkarlin@gmail.com
Boris Klempa
boris.klempa@savba.sk
Jonas Klingström
jonas.klingstrom@ki.se
Michel C. Koch
michel.koch@vetsuisse.unibe.ch
Hideki Kondō (近藤秀樹)
hkondo@okayama-u.ac.jp
Eugene V. Koonin
koonin@ncbi.nlm.nih.gov
Jarmila Krásová
krasova@ivb.cz
Mart Krupovic
mart.krupovic@pasteur.fr
Kenji Kubota (久保田健嗣)
kubotak@affrc.go.jp
Ivan V. Kuzmin
ivankuzmin1965@gmail.com
Lies Laenen
lies.laenen@uzleuven.be
Amy J. Lambert
ahk7@CDC.GOV
Jiànróng Lǐ (李建荣)
li.926@osu.edu
Jun-Min Li (李俊敏)
lijunmin@nbu.edu.cn
François Lieffrig
f.lieffrig@cergroupe.be
Igor S. Lukashevich
isluka01@louisville.edu
Dongsheng Luo
dongshengluo@outlook.com
Piet Maes
piet.maes@kuleuven.be
Marco Marklewitz
marco.marklewitz@finddx.org
Sergio H. Marshall
sergio.marshall@pucv.cl
Shin-Yi L. Marzano
shinyi.marzano@usda.gov
John W. McCauley
john.mccauley@crick.ac.uk
Ali Mirazimi
Ali.Mirazimi@ki.se
Peter G. Mohr
Peter.Mohr@csiro.au
Nick J. G. Moody
Nick.Moody@csiro.au
Yasuaki Morita (森田泰彰)
ojigotakeguro@piano.ocn.ne.jp
Richard N. Morrison
Richard.Morrison@nre.tas.gov.au
Elke Mühlberger
muehlber@bu.edu
Rayapati Naidu
naidu.rayapati@wsu.edu
Tomohide Natsuaki (夏秋知英)
natsuaki@cc.utsunomiya-u.ac.jp

J. H. Kuhn etal.
1 3
José A. Navarro
janavarr@ibmcp.upv.es
Yutaro Neriya (煉谷裕太朗)
neriya@a.utsunomiya-u.ac.jp
Sergey V. Netesov (Heтёcoв Cepгeй Bиктopoвич)
netesov.s@nsu.ru
Gabriele Neumann
gabriele.neumann@wisc.edu
Norbert Nowotny
Norbert.Nowotny@vetmeduni.ac.at
Francisco M. Ochoa-Corona
ochoaco@okstate.edu
Gustavo Palacios
gustavo.palacios@mssm.edu
Laurane Pallandre
laurane.pallandre@anses.fr
Vicente Pallás
vpallas@ibmcp.upv.es
Anna Papa (Άννα Παπά)
annap.med@gmail.com
Sofia Paraskevopoulou (Σοφία Παρασκευοπούλου)
paraskevopoulous@rki.de
Colin R. Parrish
crp3@cornell.edu
Alex Pauvolid-Corrêa
pauvolid-correa@ufv.br
Janusz T. Pawęska
januszp@nicd.ac.za
Daniel R. Pérez
dperez1@uga.edu
Florian Pfaff
Florian.Pfaff@fli.de
Richard K. Plemper
rplemper@gsu.edu
Thomas S. Postler
tp2405@cumc.columbia.edu
Françoise Pozet
fpozet@jura.fr
Sheli R. Radoshitzky
sheliijo@gmail.com
Pedro L. Ramos-González
plrg1970@gmail.com
Marius Rehanek
marius.rehanek@agrar.hu-berlin.de
Renato O. Resende
rresende@unb.br
Carina A. Reyes
carinar@biol.unlp.edu.ar
Víctor Romanowski
vromanowski@gmail.com
Dennis Rubbenstroth
Dennis.Rubbenstroth@fli.de
Luisa Rubino
luisa.rubino@ipsp.cnr.it
Artemis Rumbou (Άρτεμις Ρούμπου)
artemis.rumbou@agrar.hu-berlin.de
Jonathan A. Runstadler
jonathan.runstadler@tufts.edu
Melanie Rupp
rupp@fishdoc.ch
Sead Sabanadzovic
SSabanadzovic@entomology.msstate.edu
Takahide Sasaya (笹谷孝英)
tsasaya@affrc.go.jp
Heike Schmidt-Posthaus
heike.schmidt@unibe.ch
Martin Schwemmle
martin.schwemmle@uniklinik-freiburg.de
Torsten Seuberlich
torsten.seuberlich@vetsuisse.unibe.ch
Stephen R. Sharpe
Stephen.Sharpe@westernsydney.edu.au
Mang Shi (施莽)
shim23@mail.sysu.edu.cn
Manuela Sironi
manuela.sironi@BP.LNF.it
Sophie Smither
sjsmither@mail.dstl.gov.uk
Jin-Won Song (송진원)
jwsong@korea.ac.kr
Kirsten M. Spann
Kirsten.Spann@qut.edu.au
Jessica R. Spengler
JSpengler@cdc.gov
Mark D. Stenglein
Mark.Stenglein@ColoState.edu
Ayato Takada (高田礼人)
atakada@czc.hokudai.ac.jp
Robert B. Tesh
rtesh@utmb.edu
Jana Těšíková
jana.tesikova@gmail.com
Natalie J. Thornburg
nax3@cdc.gov
Nicole D. Tischler
ntischler@cienciavida.org
Yasuhiro Tomitaka (冨髙保弘)
yasut@affrc.go.jp
Keizō Tomonaga (朝長啓造)
tomonaga.keizo.5r@kyoto-u.ac.jp;
tomonaga@infront.kyoto-u.ac.jp
Noël Tordo
ntordo@pasteur.fr
Kenta Tsunekawa
kenta_tsunekawa@pref.aichi.lg.jp

Taxonomic update of phylum Negarnaviricota
1 3
Massimo Turina
massimo.turina@ipsp.cnr.it
Ioannis E. Tzanetakis
itzaneta@uark.edu
Anna Maria Vaira
annamaria.vaira@ipsp.cnr.it
Bernadette van den Hoogen
b.vandenhoogen@erasmusmc.nl
Bert Vanmechelen
bert.vanmechelen@kuleuven.be
Nikos Vasilakis (Νίκος Βασιλάκης)
nivasila@utmb.edu
Martin Verbeek
martin.verbeek@wur.nl
Susanne von Bargen
susanne.von.bargen@agrar.hu-berlin.de
Jiro Wada (和田治郎)
wadaj@niaid.nih.gov
Victoria Wahl
victoria.wahl@st.dhs.gov
Peter J. Walker
peter.walker@uq.edu.au
Anna E. Whitfield
awhitfi@ncsu.edu
John V. Williams
jvw@chp.edu
Yuri I. Wolf
wolf@ncbi.nlm.nih.gov
Junki Yamasaki (山﨑淳紀)
junki_yamasaki@ken3.pref.kochi.lg.jp
Hironobu Yanagisawa (栁澤広宣)
hironobu_yanagisa090@maff.go.jp
Gongyin Ye (叶恭银)
chu@zju.edu.cn
Yong-Zhen Zhang (张永振)
zhangyongzhen@icdc.cn
Arnfinn Lodden Økland
arnfinn.lodden.okland@zoetis.com
1 Integrated Research Facility atFort Detrick (IRF-Frederick),
National Institute ofAllergy andInfectious Diseases,
National Institutes ofHealth (NIH), Fort Detrick, Frederick,
MD, USA
2 United States Department ofAgriculture, Agricultural
Research Service, US Horticultural Research Laboratory,
FortPierce, FL, USA
3 D.I. Ivanovsky Institute ofVirology ofN.F. Gamaleya
National Center onEpidemiology andMicrobiology
ofMinistry ofHealth ofRussian Federation, Moscow, Russia
4 Institute ofMicrobiology andImmunology, Faculty
ofMedicine, University ofLjubljana, Ljubljana, Slovenia
5 Centro de Biotecnología y Genómica de Plantas, Universidad
Politécnica de Madrid—Instituto Nacional de Investigación
y Tecnología Agraria y Alimentaria, Campus de
Montegancedo, Pozuelo de Alarcón, Madrid, Spain
6 Departamento de Biotecnología-Biología Vegetal, Escuela
Técnica Superior de Ingeniería Agronómica, Alimentaria y de
Biosistemas, Universidad Politécnica de Madrid, Madrid, Spain
7 Center forEcology ofInfectious Diseases, Department
ofInfectious Diseases, Department ofEpidemiology
andBiostatistics, Insitute ofBioinformatics, University
ofGeorgia, Athens, GA, USA
8 Friedrich-Loeffler-Institut, Institute ofNovel andEmerging
Infectious Diseases, Greifswald-Insel Riems, Greifswald,
Germany
9 Department ofBiological Sciences, Mississippi State
University, Mississippi State, Starkville, MS, USA
10 Division Phytomedicine, Faculty ofLife Sciences,
Humboldt-Universität zu Berlin, Berlin, Germany
11 Institute ofDiagnostic Virology, Friedrich-Loeffler-Institut,
Greifswald-InselRiems, Germany
12 UFyMA, INTA-CONICET, Córdoba, Argentina
13 Division ofHigh-Consequence Pathogens andPathology,
Viral Special Pathogens Branch, Centers forDisease Control
andPrevention, Atlanta, GA, USA
14 Institute ofVirology, Philipps-University Marburg, Marburg,
Germany
15 French Agency forFood, Environmental
andOccupational Heath Safety ANSES, Laboratory
ofPloufragan-Plouzané-Niort, Ploufragan, France
16 Department ofMicrobiology, Immunology andPathology,
Colorado State University, FortCollins, CO, USA
17 Commonwealth Scientific andIndustrial Research
Organisation (CSIRO), Australian Centre forDisease
Preparedness, Geelong, VIC, Australia
18 University ofNew Mexico Health Sciences Center,
Albuquerque, NM, USA
19 Center forInfection andImmunity, andDepartment
ofEpidemiology, Mailman School ofPublic Health,
Columbia University, NewYork, NewYork, USA
20 French Agency forFood, Environmental
andOccupational Heath Safety ANSES, Laboratory
ofPloufragan-Plouzané-Niort, Ploufragan, France
21 Interfaculty Bioinformatics Unit, University ofBern, Bern,
Switzerland
22 RNA Viruses Section, Laboratory ofInfectious Diseases,
National Institute ofAllergy andInfectious Diseases,
National Institutes ofHealth, Bethesda, MD, USA
23 Department ofMolecular Biology andBiochemistry,
University ofCalifornia, Irvine, CA, USA
24 Galveston National Laboratory, The University ofTexas
Medical Branch atGalveston, Galveston, TX, USA
25 Division ofVirology, National Health Laboratory Service
andDivision ofVirology, University oftheFree State,
Bloemfontein, RepublicofSouthAfrica
26 Colorado State University, FortCollins, CO, USA
27 Univ. Bordeaux, INRAE, UMR BFP, Villenaved’Ornon, France
28 Centre forAquatic Animal Health andVaccines, Department
ofNatural Resources andEnvironment Tasmania,
Launceston, TAS, Australia

J. H. Kuhn etal.
1 3
29 Respiratory Virus andInfluenza Unit, National Microbiology
Center, Instituto de Salud Carlos III, Madrid, Spain
30 Department ofMicrobiology andImmunology, Albert
Einstein College ofMedicine, Bronx, NY, USA
31 Unité des Virus Emergents (Aix-Marseille Univ-IRD
190-Inserm 1207-IHU Méditerranée Infection), Marseille,
France
32 Division ofFruit Tree andTea Pest Control Research,
Institute forPlant Protection, NARO, Tsukuba, Ibaraki, Japan
33 Integrated Research Facility atFort Detrick, National
Institute ofAllergy andInfectious Diseases, National
Institutes ofHealth, Frederick, MD, USA
34 CSIRO Australian Centre forDisease Preparedness,
EastGeelong, VIC, Australia
35 Institut Pasteur, Université Paris Cité, Unit Lyssavirus
Epidemiology andNeuropathology, National Reference
Center forRabies, WHO Collaborating Center forReference
andResearch onRabies, Paris, France
36 CIDEFI. Facultad de Ciencias Agrarias y Forestales,
Universidad de La Plata, LaPlata, Argentina
37 Department ofImmunology andMicrobiology IMM-6, The
Scripps Research Institute, LaJolla, CA, USA
38 Unité des Virus Emergents (Aix-Marseille Univ-IRD
190-Inserm 1207-IHU Méditerranée Infection), Marseille,
France
39 World Reference Center forEmerging Viruses
andArboviruses andDepartment ofMicrobiology
andImmunology, The University ofTexas Medical Branch
atGalveston, Galveston, TX, USA
40 Department ofVirology, Wageningen Bioveterinary
Research, Lelystad, TheNetherlands
41 UMR 1161 Virology ANSES/INRAE/ENVA, ANSES
Animal Health Laboratory, Maisons-Alfort, France
42 United States Army Medical Research Institute ofInfectious
Diseases, Fort Detrick, Frederick, MD, USA
43 Istituto perla Protezione Sostenibile delle Piante, Consiglio
Nazionale delle Ricerche, Bari, Italy
44 Queensland Alliance forAgriculture andFood Innovation,
The University ofQueensland, St.Lucia, QLD, Australia
45 CIHEAM, Istituto Agronomico Mediterraneo di Bari,
Valenzano, Italy
46 Institute ofVirology, Charité-Universitätsmedizin
Berlin, corporate member ofFreie Universität Berlin,
Humboldt-Universität zu Berlin, Berlin, Germany
47 School ofMedicine, University ofPittsburgh, Pittsburgh, PA,
USA
48 Robert Koch Institut, Berlin, Germany
49 School ofLife Sciences, University ofWarwick, Coventry,
UK
50 Department ofMedical Microbiology, Virology Unit,
Hacettepe University Faculty ofMedicine, Ankara, Turkey
51 Walter Reed Biosystematics Unit (WRBU), Smithsonian
Institution, Museum Support Center, Suitland, MD, USA
52 One Health Branch, Walter Reed Army Institute ofResearch
(WRAIR), SilverSpring, MD, USA
53 Department ofEntomology, Smithsonian Institution–
National Museum ofNatural History (NMNH), Washington,
DC, USA
54 China National Rice Research Institute, Hangzhou, China
55 Departemental Analysis Laboratory, Poligny, France
56 Department ofPathology, University ofCambridge,
Cambridge, UK
57 Animal andPlant Health Agency, Weybridge, Surrey, UK
58 World Health Organization, Geneva, Switzerland
59 Embrapa Cassava andFruits, CruzdasAlmas, Bahia, Brazil
60 Institute ofBiochemistry andBiotechnology, Martin Luther
University Halle-Wittenberg, Halle/Saale, Germany
61 Instituto de Biotecnología y Biología Molecular, Facultad de
Ciencias Exactas, CONICET UNLP, LaPlata, Argentina
62 Icahn School ofMedicine atMount Sinai, NewYork, NY,
USA
63 United States Army Medical Research Institute ofInfectious
Diseases, Fort Detrick, Frederick, MD, USA
64 Department ofMicrobiology andImmunology, Division
ofBiomedical Graduate Research Organization, School
ofMedicine, Georgetown University, Washington, DC, USA
65 Institute ofVertebrate Biology oftheCzech Academy
ofSciences, Brno, CzechRepublic
66 Department ofMicrobiology andNational Emerging
Infectious Diseases Laboratories, Boston University School
ofMedicine, Boston, MA, USA
67 Institute ofNovel andEmerging Infectious Diseases,
Friedrich-Loeffler-Institut, Greifswald-InselRiems, Germany
68 Department ofVirology, WHO Collaborating Centre
forArboviruses andHemorrhagic Fever Reference
andResearch, Bernhard-Nocht Institute forTropical
Medicine, Hamburg, Germany
69 United States Department ofAgriculture, Agricultural
Research Service, USNA, Floral andNursery Plants
Research Unit, Beltsville, MD, USA
70 Department ofVirology, University ofHelsinki, Medicum,
Helsinki, Finland
71 Institute ofVeterinary Pathology, Vetsuisse Faculty,
University ofZurich, Zurich, Switzerland
72 Division ofNeurological Sciences, Vetsuisse Faculty,
University ofBern, Bern, Switzerland
73 Department ofInfectious Diseases, Yamagata University
Faculty ofMedicine, Yamagata, Japan
74 Graduate School ofVeterinary Science, Osaka Metropolitan
University, Izumisano, Osaka, Japan
75 Osaka International Research Center forInfectious Diseases,
Osaka Metropolitan University, Izumisano, Osaka, Japan
76 Aichi Agricultural Research Center, Nagakute, Aichi, Japan
77 Centers forDisease Control andPrevention, FortCollins,
CO, USA

Taxonomic update of phylum Negarnaviricota
1 3
78 Department ofMicrobiology andNational Emerging
Infectious Diseases Laboratories, Boston University School
ofMedicine, Boston, MA, USA
79 Center forEmerging Infectious Diseases Policy
andResearch, Boston University School ofMedicine,
Boston, Massachusetts, USA
80 School ofVeterinary Medicine, Murdoch University,
Murdoch, WA, Australia
81 State Key Laboratory ofAgricultural Microbiology,
Huazhong Agricultural University, Wuhan, HubeiProvince,
China
82 International Rice Research Institute, College, Los Baños,
4032Laguna, Philippines
83 Clinical Plant Science Center, Hosei University, Tokyo, Japan
84 Institute ofVirology, Biomedical Research Center, Slovak
Academy ofSciences, Bratislava, Slovakia
85 Department ofMedicine Huddinge, Center forInfectious
Medicine, Karolinska Institutet, Karolinska University
Hospital, Stockholm, Sweden
86 Division ofNeurological Sciences, Vetsuisse Faculty,
University ofBern, Bern, Switzerland
87 Institute ofPlant Science andResources, Okayama
University, Kurashiki, Japan
88 National Center forBiotechnology Information, National
Library ofMedicine, National Institutes ofHealth, Bethesda,
MD, USA
89 Institut Pasteur, Université Paris Cité, CNRS UMR6047,
Archaeal Virology Unit, Paris, France
90 Institute forPlant Protection, NARO, Tsukuba, Ibaraki, Japan
91 The University ofTexas Medical Branch atGalveston,
Galveston, TX, USA
92 KU Leuven, Rega Institute, Zoonotic Infectious Diseases
unit, Leuven, Belgium
93 Department ofLaboratory Medicine, University Hospitals
Leuven, Leuven, Belgium
94 Centers forDisease Control andPrevention, FortCollins,
CO, USA
95 Department ofVeterinary Biosciences, College ofVeterinary
Medicine, The Ohio State University, Columbus, OH, USA
96 Institute ofPlant Virology, Ningbo University, Ningbo,
China
97 CER Groupe, Fish Disease Laboratory, Marche-En-Famenne,
Belgium
98 Department ofPharmacology andToxicology, School
ofMedicine, andtheCenter forPredictive Medicine
forBiodefense andEmerging Infectious Diseases, University
ofLouisville, Louisville, KY, USA
99 Institut Pasteur, Université Paris Cité, Unit Lyssavirus
Epidemiology andNeuropathology, Paris, France
100 CAS Key Laboratory ofSpecial Pathogens andBiosafety,
Wuhan Institute ofVirology, Chinese Academy ofSciences,
Wuhan, Hubei, China
101 KU Leuven, Rega Institute, Zoonotic Infectious Diseases
unit, Leuven, Belgium
102 FIND, theglobal alliance fordiagnostics, Geneva,
Switzerland
103 Instituto de Biología-Laboratorio de Genética
Molecular-Campus Curauma, Valparaíso, Chile
104 United States Department ofAgriculture, Agricultural
Research Service, Toledo, OH, USA
105 Worldwide Influenza Centre, Francis Crick Institute, London, UK
106 Folkhalsomyndigheten, Stockholm, Sweden
107 Kochi Agricultural Research Center, Kochi, Japan
108 Department ofMicrobiology andNational Emerging
Infectious Diseases Laboratories, Boston University School
ofMedicine, Boston, MA, USA
109 Department ofPlant Pathology, Irrigated Agricultural
Research andExtension Center, Washington State University,
Prosser, WA, USA
110 Utsunomiya University, Utsunomiya, Japan
111 Instituto de Biología Molecular y Celular de Plantas,
Universitat Politècnica de València-Consejo Superior de
Investigaciones Científicas, Valencia, Spain
112 School ofAgriculture, Utsunomiya University, Utsunomiya,
Japan
113 Novosibirsk State University, Novosibirsk,
NovosibirskOblast, Russia
114 Department ofPathobiological Sciences, Influenza Research
Institute, University ofWisconsin-Madison, Madison, USA
115 Institute ofVirology, University ofVeterinary Medicine
Vienna, Vienna, Austria
116 College ofMedicine, Mohammed Bin Rashid
University ofMedicine andHealth Sciences, Dubai,
UnitedArabEmirates
117 Institute forBiosecurity andMicrobial Forensics. Stillwater,
Oklahoma State University, Oklahoma, USA
118 Department ofMicrobiology, Icahn School ofMedicine
atMount Sinai, NewYork, NY, USA
119 French Agency forFood, Environmental
andOccupational Heath Safety ANSES, Laboratory
ofPloufragan-Plouzané-Niort, Ploufragan, France
120 Instituto de Biología Molecular y Celular de Plantas (IBMCP),
Consejo Superior de Investigaciones Cientificas-Universidat
Politècnica de Valencia, Valencia, Spain
121 National Reference Centre forArboviruses
andHaemorrhagic Fever viruses, Department
ofMicrobiology, Medical School, Aristotle University
ofThessaloniki, Thessaloniki, Greece
122 Methods Development andResearch Infrastructure,
Bioinformatics andSystems Biology, Robert Koch Institute,
Berlin, Germany
123 College ofVeterinary Medicine, Baker Institute forAnimal
Health, Cornell University, Ithaca, NY, USA
124 Departamento de Veterinária, Universidade Federal de
Viçosa, Viçosa, MG, Brazil
125 Center forEmerging Zoonotic andParasitic Diseases,
National Institute forCommunicable Diseases oftheNational

J. H. Kuhn etal.
1 3
Health Laboratory Service, Sandringham-Johannesburg,
Gauteng, SouthAfrica
126 Department ofPopulation Health, College ofVeterinary
Medicine, University ofGeorgia, Athens, GA, USA
127 Institute ofDiagnostic Virology, Friedrich-Loeffler-Institut,
Greifswald-InselRiems, Germany
128 Department ofMicrobiology andImmunology, Vagelos
College ofPhysicians andSurgeons, Columbia University
Irving Medical Center, NewYork, NY, USA
129 United States Army Medical Research Institute ofInfectious
Diseases (USAMRIID), Frederick, MD, USA
130 Instituto Biológico de São Paulo, SãoPaulo, SP, Brazil
131 Departamento de Biologia Celular, Universidade de Brasília,
Brasília, Brazil
132 Instituto de Biotecnología y Biología Molecular,
CONICET-UNLP, Facultad de Ciencias Exactas, Unversidad
Nacional de La Plata, BuenosAires, Argentina
133 Instituto de Biotecnología y Biología Molecular,
CONICET-UNLP, Facultad de Ciencias Exactas, Unversidad
Nacional de La Plata, BuenosAires, Argentina
134 Institute ofDiagnostic Virology, Friedrich-Loeffler-Institut,
Greifswald-InselRiems, Germany
135 Department ofInfectious Disease & Global Health, Tufts
University Cummings School ofVeterinary Medicine,
NorthGrafton, MA, USA
136 Institute forFish andWildlife Health, Vetsuisse Faculty,
University ofBern, Bern, Switzerland
137 Department ofBiochemistry, Molecular Biology,
Entomology andPlant Pathology, Mississippi State
University, MississippiState, MS, USA
138 Institute forPlant Protection, National Agriculture andFood
Research Organization, Tsukuba, Japan
139 Institute forFish andWildlife Health, Vetsuisse Faculty,
University ofBern, Bern, Switzerland
140 Faculty ofMedicine, University Medical Center-University
Freiburg, Freiburg, Germany
141 Division ofNeurological Sciences, Vetsuisse Faculty,
University ofBern, Bern, Switzerland
142 Hawkesbury Institute fortheEnvironment, Western Sydney
University, Sydney, NSW, Australia
143 Sun Yat-sen University, Shenzhen, China
144 Bioinformatics Unit, Scientific Institute IRCCS “E. Medea”,
BosisioParini, Italy
145 CBR Division, Dstl, Porton Down, Salisbury, Wiltshire, UK
146 Department ofMicrobiology, College ofMedicine, Korea
University, Seoul, RepublicofKorea
147 School ofBiomedical Sciences, Faculty ofHealth,
Queensland University ofTechnology, Brisbane, QLD,
Australia
148 Viral Special Pathogens Branch, Division
ofHigh-Consequence Pathogens andPathology, Centers
forDisease Control andPrevention, Atlanta, GA, USA
149 Department ofMicrobiology, Immunology, andPathology,
College ofVeterinary Medicine andBiomedical Sciences,
Colorado State University, FortCollins, CO, USA
150 Division ofGlobal Epidemiology, International Institute
forZoonosis Control, Hokkaido University, Sapporo, Japan
151 Department ofBotany andZoology, Faculty ofScience,
Masaryk University, Brno, CzechRepublic
152 Centers forDisease Control andPrevention, Atlanta, GA, USA
153 Laboratorio de Virología Molecular, Centro Ciencia & Vida,
Fundación Ciencia & Vida andFacultad de Medicina y
Ciencia, Universidad San Sebastián, Santiago, Chile
154 Institute forPlant Protection, National Agriculture andFood
Research Organization, Tsukuba, Japan
155 Institute forLife andMedical Sciences (LiMe), Kyoto
University, Kyoto, Japan
156 Institut Pasteur de Guinée, BP 4416, Conakry, Guinea
157 Aichi Agricultural Research Center, Nagakute, Aichi, Japan
158 Institute forSustainable Plant Protection, National Research
Council ofItaly (IPSP-CNR), Torino, Italy
159 Department ofEntomology andPlant Pathology, Division
ofAgriculture, University ofArkansas System, Fayetteville,
AR, USA
160 Department ofViroscience, Erasmus MC, University
Medical Centre Rotterdam, Rotterdam, TheNetherlands
161 Department ofMicrobiology, Immunology
andTransplantation, KU Leuven, Leuven, Belgium
162 Wageningen University andResearch, Biointeractions
andPlant Health, Wageningen, TheNetherlands
163 Integrated Research Facility atFort Detrick, National
Institute ofAllergy andInfectious Diseases, National
Institutes ofHealth, Frederick, MD, USA
164 National Biodefense Analysis andCountermeasures Center,
Fort Detrick, Frederick, MD, USA
165 School ofChemistry andMolecular Biosciences, University
ofQueensland, St.Lucia, QLD, Australia
166 Department ofEntomology andPlant Pathology, North
Carolina State University, Raleigh, NC, USA
167 School ofMedicine, University ofPittsburgh, Pittsburgh, PA, USA
168 National Center forBiotechnology Information, National
Library ofMedicine, National Institutes ofHealth, Bethesda,
MD, USA
169 Environmental Agriculture Promotion Division, Department
ofAgricultural Development, Kochi Prefectural Government,
Kochi, Kochi, Japan
170 Yokohama Plant Protection Station, Yokohama, Kanagawa, Japan
171 Institute ofInsect Sciences, Zhejiang University, Hangzhou,
China
172 National Institute forCommunicable Disease Control
andPrevention, China Center forDisease Control
andPrevention, Beijing, China
173 Pharmaq Analytiq, Bergen, Norway