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Mononegavirales is an order of negative-strand RNA viruses which have nonsegmented genomes. Some members that cause human disease in this order include Ebola virus, human respiratory syncytial virus, measles virus, mumps virus, Nipah virus, and rabies virus. Important pathogens of nonhuman animals and plants are also in the group. The order includes eleven virus families: Artoviridae, Bornaviridae, Filoviridae, Lispiviridae, Mymonaviridae, Nyamiviridae, Paramyxoviridae, Pneumoviridae, Rhabdoviridae, Sunviridae, and Xinmoviridae.[1]

Use of term

The order Mononegavirales (pronounced: /ˌmɒnəˌnɛɡəviˈrɑːlɪz/ MON-ə-NEG-ə-vee-RAH-liz) [note 1][2][3] is a virological taxon that was created in 1991[4][5] and amended in 1995,[6] 1997,[7] 2000,[8] 2005,[9] 2011,[2] 2016,[10] 2017,[11] and 2018.[1] The name Mononegavirales is derived from the Ancient Greek adjective μóνος monos (alluding to the monopartite and single-stranded genomes of most mononegaviruses), the Latin verb negare (alluding to the negative polarity of these genomes), and the taxonomic suffix -virales (denoting a viral order).[3]

Order inclusion criteria

The genome organization and RNA synthesis of order Mononegavirales

A virus is a member of the order Mononegavirales if[2][3]

  • its genome is a linear, typically (but not always) nonsegmented, single-stranded, non-infectious RNA of negative polarity; possesses inverse-complementary 3' and 5' termini; and is not covalently linked to a protein;
  • its genome has the characteristic gene order 3'-UTR–core protein genes–envelope protein genes–RNA-dependent RNA polymerase gene–5'-UTR (3'-N-P-M-G-L-5') (there are, however, some exceptions);
  • it produces 5–10 distinct mRNAs from its genome via polar sequential transcription from a single promoter located at the 3' end of the genome; mRNAs are 5' capped and polyadenylated;
  • it replicates by synthesizing complete antigenomes;
  • it forms infectious helical ribonucleocapsids as the templates for the synthesis of mRNAs, antigenomes, and genomes;
  • it encodes an RNA-dependent RNA polymerase (RdRp, L) that is highly homologous to those of other mononegaviruses; and/or
  • it typically (but not always) produces enveloped virions with a molecular mass of 300–1,000×106; an S20W of 550–>1,045; and a buoyant density in CsCl of 1.18–1.22 g/cm3.

Life cycle

Life cycle of vesiculoviruses

The mononegavirus life cycle begins with virion attachment to specific cell-surface receptors, followed by fusion of the virion envelope with cellular membranes and the concomitant release of the virus nucleocapsid into the cytosol. The virus RdRp partially uncoats the nucleocapsid and transcribes the genes into positive-stranded mRNAs, which are then translated into structural and nonstructural proteins.[9]

Mononegavirus RdRps bind to a single promoter located at the 3' end of the genome. Transcription either terminates after a gene or continues to the next gene downstream. This means that genes close to the 3' end of the genome are transcribed in the greatest abundance, whereas those toward the 5' end are least likely to be transcribed. The gene order is therefore a simple but effective form of transcriptional regulation. The most abundant protein produced is the nucleoprotein, whose concentration in the cell determines when the RdRp switches from gene transcription to genome replication.[9]

Replication results in full-length, positive-stranded antigenomes that are in turn transcribed into negative-stranded virus progeny genome copies. Newly synthesized structural proteins and genomes self-assemble and accumulate near the inside of the cell membrane. Virions bud off from the cell, gaining their envelopes from the cellular membrane they bud from. The mature progeny particles then infect other cells to repeat the cycle.[9]


Mononegaviruses have a history that dates back several tens of million of years. Mononegavirus "fossils" have been discovered in the form of mononegavirus genes or gene fragments integrated into mammalian genomes. For instance, bornavirus gene "fossils" have been detected in the genomes of bats, fish, hyraxes, marsupials, primates, rodents, ruminants, and elephants.[12][13][14][15][16] Filovirus gene "fossils" have been detected in the genomes of bats, rodents, shrews, tenrecs, and marsupials.[13][14][17][18] A Midway virus "fossil" was found in the genome of zebrafish.[13] Finally, rhabdovirus "fossils" were found in the genomes of crustaceans, mosquitoes, ticks, and plants.[19][14][20][21]


Mononegavirales phylogenetic tree

The order has eleven families that include numerous genera, which consist of many different species:

Table of the order showing all families, genera, species, and their viruses:[1]

Family Genus Species Virus (Abbreviation) Bornaviridae Carbovirus Queensland carbovirus* jungle carpet python virus (JCPV) Southwest carbovirus southwest carpet python virus (SWCPV) Orthobornavirus Elapid 1 orthobornavirus Loveridge’s garter snake virus 1 (LGSV-1) Mammalian 1 orthobornavirus* Borna disease virus 1 (BoDV-1) Borna disease virus 2 (BoDV-2) Mammalian 2 orthobornavirus variegated squirrel bornavirus 1 (VSBV-1) Passeriform 1 orthobornavirus canary bornavirus 1 (CnBV-1) canary bornavirus 2 (CnBV-2) canary bornavirus 3 (CnBV-3)) Passeriform 2 orthobornavirus estrildid finch bornavirus 1 (EsBV-1) Psittaciform 1 orthobornavirus parrot bornavirus 1 (PaBV-1) parrot bornavirus 2 (PaBV-2) parrot bornavirus 3 (PaBV-3) parrot bornavirus 4 (PaBV-4) parrot bornavirus 7 (PaBV-7) Psittaciform 2 orthobornavirus parrot bornavirus 5 (PaBV-5) Waterbird 1 orthobornavirus aquatic bird bornavirus 1 (ABBV-1) aquatic bird bornavirus 2 (ABBV-2) Filoviridae Cuevavirus Lloviu cuevavirus* Lloviu virus (LLOV) Dianlovirus Měnglà virus (MLAV) Ebolavirus 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) Mymonaviridae Sclerotimonavirus Sclerotinia sclerotimonavirus* Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1) Nyamiviridae Nyavirus Midway nyavirus Midway virus (MIDWV) Nyamanini nyavirus* Nyamanini virus (NYMV) Sierra Nevada nyavirus Sierra Nevada virus (SNVV) Peropuvirus Pteromalus puparum peropuvirus* Pteromalus puparum negative-strand RNA virus 1 (PpNSRV-1) Socyvirus Soybean cyst nematode socyvirus* soybean cyst nematode virus 1 (SbCNV-1) Paramyxoviridae Aquaparamyxovirus Salmon aquaparamyxovirus* Atlantic salmon paramyxovirus (AsaPV) Avulavirus Avian avulavirus 1* avian paramyxovirus 1 (APMV-1) Avian avulavirus 2 avian paramyxovirus 2 (APMV-2) Avian avulavirus 3 avian paramyxovirus 3 (APMV-3) Avian avulavirus 4 avian paramyxovirus 4 (APMV-4) Avian avulavirus 5 avian paramyxovirus 5 (APMV-5) Avian avulavirus 6 avian paramyxovirus 6 (APMV-6) Avian avulavirus 7 avian paramyxovirus 7 (APMV-7) Avian avulavirus 8 avian paramyxovirus 8 (APMV-8) Avian avulavirus 9 avian paramyxovirus 9 (APMV-9) Avian avulavirus 10 avian paramyxovirus 10 (APMV-10) Avian avulavirus 11 avian paramyxovirus 11 (APMV-11) Avian avulavirus 12 avian paramyxovirus 12 (APMV-12) Avian avulavirus 13 avian paramyxovirus 13 (APMV-13) Avian avulavirus 14 avian paramyxovirus 14 (APMV-14) Avian avulavirus 15 avian paramyxovirus 15 (APMV-15) Avian avulavirus 16 avian paramyxovirus 16 (APMV-16) Avian avulavirus 17 Antarctic penguin virus A (APV-A) Avian avulavirus 18 Antarctic penguin virus B (APV-B)) Avian avulavirus 19 Antarctic penguin virus C (APV-C) Ferlavirus Reptilian ferlavirus* Fer-de-Lance virus (FDLV) Henipavirus Cedar henipavirus Cedar virus (CedV) Ghanaian bat henipavirus Kumasi virus (KV) Hendra henipavirus* Hendra virus (HeV) Mojiang henipavirus Mòjiāng virus (MojV) Nipah henipavirus Nipah virus (NiV) Morbillivirus Canine morbillivirus canine distemper virus (CDV) Cetacean morbillivirus cetacean morbillivirus (CeMV) Feline morbillivirus feline morbillivirus (FeMV) Measles morbillivirus* measles virus (MeV) Small ruminant morbillivirus peste-des-petits-ruminants virus (PPRV) Phocine morbillivirus phocine distemper virus (PDV) Rinderpest morbillivirus rinderpest virus (RPV) Respirovirus Bovine respirovirus 3 bovine parainfluenza virus 3 (BPIV-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) Rubulavirus Achimota rubulavirus 1 Achimota virus 1 (AchPV-1) Achimota rubulavirus 2 Achimota virus 2 (AchPV-2) Bat mumps rubulavirus bat mumps virus (BMV) Canine rubulavirus parainfluenza virus 5 (PIV-5) Human rubulavirus 2 human parainfluenza virus 2 (HPIV-2) Human rubulavirus 4 human parainfluenza virus 4a (HPIV-4a) human parainfluenza virus 4b (HPIV-4b) Mapuera rubulavirus Mapuera virus (MapV) Menangle rubulavirus Menangle virus (MenPV) Mumps rubulavirus* mumps virus (MuV) Porcine rubulavirus La Piedad Michoacán Mexico virus (LPMV) Simian rubulavirus simian virus 41 (SV-41) Sosuga rubulavirus Sosuga virus Teviot rubulavirus Teviot virus (TevPV) Tioman rubulavirus Tioman virus (TioPV) Tuhoko rubulavirus 1 Tuhoko virus 1 (ThkPV-1) Tuhoko rubulavirus 2 Tuhoko virus 2 (ThkPV-2) Tuhoko rubulavirus 3 Tuhoko virus 3 (ThkPV-3) 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 A2 (HRSV-A2) human respiratory syncytial virus B1 (HRSV-B1) Murine orthopneumovirus murine pneumonia virus (MPV) Rhabdoviridae Almendravirus Arboretum almendravirus Arboretum virus (ABTV) Balsa almendravirus Balsa virus (BALV) Coot Bay almendravirus Coot Bay virus (CBV) Puerto Almendras almendravirus* Puerto Almendras virus (PTAMV) Rio Chico almendravirus Rio Chico virus (RCHV) Curiovirus Curionopolis curiovirus* Curionopolis virus (CURV) Iriri curiovirus Iriri virus (IRIRV) Itacaiunas curiovirus Itacaiunas virus (ITAV) Rochambeau curiovirus Rochambeau virus (RBUV) Cytorhabdovirus Alfalfa dwarf cytorhabdovirus alfalfa dwarf virus (ADV) Barley yellow striate mosaic cytorhabdovirus barley yellow striate mosaic virus (BYSMV) Broccoli necrotic yellows cytorhabdovirus broccoli necrotic yellows virus (BNYV) Colocasia bobone disease-associated cytorhabdovirus Colocasia bobone disease-associated virus (CBDaV) Festuca leaf streak cytorhabdovirus festuca leaf streak virus (FLSV) Lettuce necrotic yellows cytorhabdovirus* lettuce necrotic yellows virus (LNYV) Lettuce yellow mottle cytorhabdovirus lettuce yellow mottle virus (LYMoV) Northern cereal mosaic cytorhabdovirus northern cereal mosaic virus (NCMV) Sonchus cytorhabdovirus sonchus virus (SonV) Strawberry crinkle cytorhabdovirus strawberry crinkle virus (SCV) Wheat American striate mosaic cytorhabdovirus wheat American striate mosaic virus (WASMV) Dichorhavirus Coffee ringspot dichorhavirus coffee ringspot virus (CoRSV) Orchid fleck dichorhavirus* orchid fleck virus (OFV) Ephemerovirus Adelaide River ephemerovirus Adelaide River virus (ARV) Berrimah ephemerovirus Berrimah virus (BRMV) Bovine fever ephemerovirus* bovine ephemeral fever virus (BEFV) Kimberley ephemerovirus Kimberley virus (KIMV) Malakal virus (MALV) Koolpinyah ephemerovirus Koolpinyah virus (KOOLV) Kotonkan ephemerovirus kotonkan virus (KOTV) Obodhiang ephemerovirus Obodhiang virus (OBOV) Yata ephemerovirus Yata virus (YATV) Hapavirus Flanders hapavirus Flanders virus (FLAV) Hart Park hapavirus Hart Park virus (HPV) Gray Lodge hapavirus Gray Lodge virus (GLOV) Joinjakaka hapavirus Joinjakaka virus (JOIV) La Joya hapavirus La Joya virus (LJV) Kamese hapavirus Kamese virus (KAMV) Landjia hapavirus Landjia virus (LANV = LJAV) Manitoba hapavirus Manitoba virus (MANV = MNTBV) Marco hapavirus Marco virus (MCOV) Mosqueiro hapavirus Mosqueiro virus (MQOV) Mossuril hapavirus Mossuril virus (MOSV) Ngaingan hapavirus Ngaingan virus (NGAV) Ord River hapavirus Ord River virus (ORV) Parry Creek hapavirus Parry Creek virus (PCV) Wongabel hapavirus'*' Wongabel virus (WONV) Ledantevirus Barur ledantevirus Barur virus (BARV) Fikirini ledantevirus Fikirini virus (FKRV) Fukuoka ledantevirus Fukuoka virus (FUKV) Kanyawara ledantevirus Kanyawara virus (KYAV) Kern Canyon ledantevirus Kern Canyon virus (KCV) Keuraliba ledantevirus Keuraliba virus (KEUV) Kolente ledantevirus Kolente virus (KOLEV) Kumasi ledantevirus Kumasi rhabdovirus (KRV) Le Dantec ledantevirus* Le Dantec virus (LDV) Mount Elgon bat ledantevirus Mount Elgon bat virus (MEBV) Nkolbisson ledantevirus Nkolbisson virus (NKOV) Nishimuro ledantevirus Nishimuro virus (NISV) Oita ledantevirus Oita virus (OITAV) Wuhan ledantevirus Wǔhàn louse fly virus 5 (WLFV-5) Yongjia ledantevirus Yǒngjiā tick virus 2 (YTV-2) Lyssavirus Aravan lyssavirus Aravan virus (ARAV) Australian bat lyssavirus Australian bat lyssavirus (ABLV) Bokeloh bat lyssavirus Bokeloh bat lyssavirus (BBLV) Duvenhage lyssavirus Duvenhage virus (DUVV) European bat 1 lyssavirus European bat lyssavirus 1 (EBLV-1) European bat 2 lyssavirus European bat lyssavirus 2 (EBLV-2) Gannoruwa bat lyssavirus Gannoruwa bat lyssavirus (GBLV) Ikoma lyssavirus Ikoma lyssavirus (IKOV) Irkut lyssavirus Irkut virus (IRKV) Khujand lyssavirus Khujand virus (KHUV) Lagos bat lyssavirus Lagos bat virus (LBV) Lleida bat lyssavirus Lleida bat virus (LLEBV) Mokola lyssavirus Mokola virus (MOKV) Rabies lyssavirus* rabies virus (RABV) Shimoni bat lyssavirus Shimoni bat virus (SHIBV) West Caucasian bat lyssavirus West Caucasian bat virus (WCBV) Novirhabdovirus Hirame novirhabdovirus hirame rhabdovirus (HIRV) Piscine novirhabdovirus viral hemorrhagic septicemia virus (VHSV) Salmonid novirhabdovirus* infectious hematopoietic necrosis virus (IHNV) Snakehead novirhabdovirus snakehead rhabdovirus (SHRV) Nucleorhabdovirus Datura yellow vein nucleorhabdovirus datura yellow vein virus (DYVV) Eggplant mottled dwarf nucleorhabdovirus eggplant mottled dwarf virus (EMDV) Maize fine streak nucleorhabdovirus maize fine streak virus (MSFV) Maize Iranian mosaic nucleorhabdovirus maize Iranian mosaic virus (MIMV) Maize mosaic nucleorhabdovirus maize mosaic virus (MMV) Potato yellow dwarf nucleorhabdovirus* potato yellow dwarf virus (PYDV) Rice yellow stunt nucleorhabdovirus rice yellow stunt virus (RYSV) rice transitory yellowing virus (RTYV) Sonchus yellow net nucleorhabdovirus sonchus yellow net virus (SYNV) Sowthistle yellow vein nucleorhabdovirus sowthistle yellow vein virus (SYVV) Taro vein chlorosis nucleorhabdovirus taro vein chlorosis virus (TaVCV) Perhabdovirus Anguillid perhabdovirus eel virus European X (EVEX) Perch perhabdovirus* perch rhabdovirus (PRV) Sea trout perhabdovirus lake trout rhabdovirus (LTRV) Sigmavirus Drosophila affinis sigmavirus Drosophila affinis sigmavirus (DAffSV) Drosophila ananassae sigmavirus Drosophila ananassae sigmavirus (DAnaSV) Drosophila immigrans sigmavirus Drosophila immigrans sigmavirus (DImmSV) Drosophila melanogaster sigmavirus* Drosophila melanogaster sigmavirus (DMelSV) Drosophila obscura sigmavirus Drosophila obscura sigmavirus (DObsSV) Drosophila tristis sigmavirus Drosophila tristis sigmavirus (DTriSV) Muscina stabulans sigmavirus Muscina stabulans sigmavirus (MStaSV) Sprivivirus Carp sprivivirus* spring viremia of carp virus (SVCV) Pike fry sprivivirus grass carp rhabdovirus (GrCRV) pike fry rhabdovirus (PFRV) tench rhabdovirus (TenRV) Sripuvirus Almpiwar sripuvirus Almpiwar virus (ALMV) Chaco sripuvirus Chaco virus (CHOV) Niakha sripuvirus* Niakha virus (NIAV) Sena Madureira sripuvirus Sena Madureira virus (SMV) Sripur sripuvirus Sripur virus (SRIV) Tibrovirus Bas-Congo tibrovirus Bas-Congo virus (BASV) Beatrice Hill tibrovirus Beatrice Hill virus (BHV) Coastal Plains tibrovirus Coastal Plains virus (CPV) Ekpoma 1 tibrovirus Ekpoma virus 1 (EKV-1) Ekpoma 2 tibrovirus Ekpoma virus 2 (EKV-2) Sweetwater Branch tibrovirus Sweetwater Branch virus (SWBV) Tibrogargan tibrovirus* Bivens Arm virus (BAV) Tibrogargan virus (TIBV) Tupavirus Durham tupavirus* Durham virus (DURV) Klamath tupavirus* Klamath virus (KLAV) Tupaia tupavirus tupaia virus (TUPV) Varicosavirus Lettuce big-vein associated varicosavirus* lettuce big-vein associated virus (LBVaV) Vesiculovirus Alagoas vesiculovirus vesicular stomatitis Alagoas virus (VSAV) American bat vesiculovirus American bat vesiculovirus (ABVV) Carajas vesiculovirus Carajás virus (CJSV) Chandipura vesiculovirus Chandipura virus (CHPV) Cocal vesiculovirus Cocal virus (COCV) Indiana vesiculovirus* vesicular stomatitis Indiana virus (VSIV) Isfahan vesiculovirus Isfahan virus (ISFV) Jurona vesiculovirus Jurona virus (JURV) Malpais Spring vesiculovirus Malpais Spring virus (MSPV) Maraba vesiculovirus Maraba virus (MARAV) Morreton vesiculovirus Morreton virus (MORV) New Jersey vesiculovirus vesicular stomatitis New Jersey virus (VSNJV) Perinet vesiculovirus Perinet virus (PERV) Piry vesiculovirus Piry virus (PIRYV) Radi vesiculovirus Radi virus (RADV) Yug Bogdanovac vesiculovirus Yug Bogdanovac virus (YBV) Unassigned Moussa virus Moussa virus (MOUV) Sunviridae Sunshinevirus Reptile sunshinevirus 1* Sunshine Coast virus (SunCV) Unassigned Anphevirus Xincheng anphevirus* Xīnchéng mosquito virus (XcMV) Unassigned Arlivirus Lishi arlivirus* Líshí spider virus 2 (LsSV-2) Unassigned Chengtivirus Tacheng chengtivirus* Tǎchéng tick virus 6 (TcTV-6) Unassigned Crustavirus Wenzhou crustavirus* Wēnzhōu crab virus 1 (WzCV-1) Unassigned Wastrivirus Sanxia wastrivirus* Sānxiá water strider virus 4 (SxWSV-4)

Table legend: "*" denotes type species.


  1. ^ According to the rules for taxon naming established by the International Committee on Taxonomy of Viruses (ICTV), the name Mononegavirales is always to be capitalized, italicized, and never abbreviated. The names of the order's physical members ("mononegaviruses" or "mononegavirads") are to be written in lower case, are not italicized, and used without articles.


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Mononegavirales: Brief Summary

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Mononegavirales is an order of negative-strand RNA viruses which have nonsegmented genomes. Some members that cause human disease in this order include Ebola virus, human respiratory syncytial virus, measles virus, mumps virus, Nipah virus, and rabies virus. Important pathogens of nonhuman animals and plants are also in the group. The order includes eleven virus families: Artoviridae, Bornaviridae, Filoviridae, Lispiviridae, Mymonaviridae, Nyamiviridae, Paramyxoviridae, Pneumoviridae, Rhabdoviridae, Sunviridae, and Xinmoviridae.

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