Order Mononegavirales
Virus classification
Group: Group V ((-)ssRNA)
Order: Mononegavirales


The order Mononegavirales is the taxonomic home of numerous related viruses. Members of the order that are commonly known are, for instance, Ebola virus, human respiratory syncytial virus, measles virus, mumps virus, Nipah virus, and rabies virus. All of these viruses cause significant disease in humans. Many very important pathogens of nonhuman animals and plants are also members of this order.


Use of term

The order Mononegavirales is a virological taxon (i.e. a concept) that was created in 1991[1][2] and emended in 1995,[3] 1997,[4] 2000,[5] and 2005.[6] The name Mononegavirales is derived from the Greek adjective μóνος [monos] (alluding to the single-stranded genomes of mononegaviruses), the Latin verb negare (alluding to the negative polarity of these genomes], and the taxonomic suffix -virales (denoting a viral order).[7] The order currently includes the four virus families Bornaviridae, Filoviridae, Paramyxoviridae, and Rhabdoviridae.[6][7][8]


Mononegavirales is pronounced ˌmɒnəˌnɛgəviː’rɑ:lɨz (IPA) or mo-nuh-ne-guh-vee-rah-liz in English phonetic notation.[7] 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, never abbreviated and to be preceded by the word "order".[6] The names of its physical members (mononegaviruses/mononegavirads) are to be written in lower case, are not italicized and used without articles.[6][7]

Order inclusion criteria

A virus is a member of the order Mononegavirales if[6][7]

  • its genome is: a linear, nonsegmented, single-stranded, non-infectious RNA of negative polarity; possesses inverse-complementary 3' and 5' termini; and does not possess a 5' cap, is not polyadenylated, 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
  • 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
  • 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) that is highly homologous to those of other mononegaviruses
  • it forms enveloped virions with a molecular mass of 300–1,000 x 106; an S20W of 550–>1,045; and a buoyant density in CsCl of 1.18–1.22 g/cm3

Order organization

The order includes four accepted families that include numerous genera, consisting of many different species. Subfamilies have only been established for the mononegavirus family Paramyxoviridae. The order has expanded considerably during recent years due to the discovery of many novel agents that were found to be phylogenetically diverse from already known mononegaviruses. Novel taxa (genera and/or species) had to be proposed, some of which have by now been accepted by the ICTV, and others that are in various stages of the suggestion/proposal/consideration process. The table below provides an overview of the current composition of the order. Note that this table only lists taxa (concepts), but not their virus members (physical entities).

Order Mononegavirales: families, genera, and species
Family name Subfamily name Genus name Species name
Unassigned "Nyavirus" (suggested)[9] "Midway virus" (suggested)[9]
"Nyamanini virus"* (suggested)[9]
Bornaviridae Bornavirus "Avian bornavirus 1" (suggested)[10]
"Avian bornavirus 2" (suggested)[10]
"Avian bornavirus 3" (suggested)[10]
"Avian bornavirus 4" (suggested)[10]
"Avian bornavirus 5" (suggested)[10]
"Canada geese bornavirus" (suggested)[10]
"Canary bornavirus" (suggested)[10]
Borna disease virus*
Filoviridae "Cuevavirus" (suggested)[7] "Lloviu cuevavirus"* (suggested)[7]
Ebolavirus Bundibugyo ebolavirus (accepted)[7]
Reston ebolavirus
Sudan ebolavirus
Taï Forest ebolavirus
Zaire ebolavirus*
Marburgvirus Marburg marburgvirus* (accepted)[7]
Paramyxoviridae Paramyxovirinae Aquaparamyxovirus (accepted)[11][12] Atlantic salmon paramyxovirus* (accepted)[11][12]
Avulavirus Avian paramyxovirus 2
Avian paramyxovirus 3
Avian paramyxovirus 4
Avian paramyxovirus 5
Avian paramyxovirus 6
Avian paramyxovirus 7
Avian paramyxovirus 8
Avian paramyxovirus 9
"Avian paramyxovirus 10" (suggested)[13]
Newcastle disease virus*
Ferlavirus (accepted)[14] Fer-de-Lance paramyxovirus* (accepted)[14]
Henipavirus Hendra virus*
Nipah virus
"Jeilongvirus" (suggested)[15] "Beilong virus" (suggested)[15]
"J virus"* (suggested)[15]
Morbillivirus Canine distemper virus
Cetacean morbillivirus
Measles virus*
Peste-des-petits-ruminants virus
Phocine distemper virus
Rinderpest virus
Respirovirus Bovine parainfluenza virus 3
Human parainfluenza virus 1
Human parainfluenza virus 3
Sendai virus*
Simian virus 10
Rubulavirus Human parainfluenza virus 2
Human parainfluenza virus 4
Mapuera virus
"Menangle virus" (tentative)
Mumps virus*
Parainfluenza virus 5
Porcine rubulavirus
Simian virus 41
"Tioman virus" (tentative)
"Tuhoko paramyxovirus 1" (suggested)[16]
"Tuhoko paramyxovirus 2" (suggested)[16]
"Tuhoko paramyxovirus 3" (suggested)[16]
"TPMV-like viruses" (suggested)[17] "Tupaia paramyxovirus"* (suggested)[17]
Unassigned "'Mossman virus" (suggested)[18]
"Nariva virus" (suggested)[19]
"Pacific salmon paramyxovirus" (suggested)[20]
"Salem virus" (suggested)[21]
Pneumovirinae Pneumovirus Human respiratory syncytial virus*
Bovine respiratory syncytial virus
Murine pneumonia virus
Metapneumovirus Avian metapneumovirus*
Human metapneumovirus
Rhabdoviridae Cytorhabdovirus Barley yellow striate mosaic virus
Broccoli necrotic yellows virus
Festuca leaf streak virus
Lettuce necrotic yellows virus*
Lettuce yellow mottle virus
Northern cereal mosaic virus
Sonchus virus
Strawberry crinkle virus
Wheat American striate mosaic virus
"Dichorhabdovirus" (suggested)[22] "Orchid fleck virus"* (suggested)[22]
Ephemerovirus Adelaide River virus
Berrimah virus
Bovine ephemeral fever virus*
"Kimberley virus" (tentative)
Kotonkan virus (accepted)
"Malakal virus" (tentative)
"Obodhiang virus" (proposed)
"Puchong virus" (tentative)
"Hart Park-like viruses" (suggested) "Flanders virus"* (suggested)[23]
"Wongabel virus" (suggested)[23]
Lyssavirus Aravan virus
Australian bat lyssavirus
Duvenhage virus
European bat lyssavirus 1
European bat lyssavirus 2
Irkut virus
Khujand virus
Lagos bat virus
Mokola virus
Rabies virus*
"Rochambeau virus" (tentative)
"Shimoni bat virus" (proposed)[24]
West Caucasian bat virus
Novirhabdovirus "Eel virus B12" (tentative)
"Eel virus C26" (tentative)
Hirame rhabdovirus
Infectious hematopoietic necrosis virus*
Snakehead rhabdovirus
Viral hemorrhagic septicemia virus
Nucleorhabdovirus Datura yellow vein virus
Eggplant mottled dwarf virus
Maize fine streak virus
Maize Iranian mosaic virus (accepted)[25]
Maize mosaic virus
Potato yellow dwarf virus*
Rice yellow stunt virus
Sonchus yellow net virus
Sowthistle yellow vein virus
Taro vein chlorosis virus
"Perch rhabdovirus group" (suggested)[26] "Genotype A/Perch rhabdovirus"* (suggested)[26]
"Genotype B" (suggested)[26]
"Genotype C" (suggested)[26]
"Genotype D" (suggested)[26]
"Sigmavirus" (proposed)[27] "Culex tritaeniorhynchus rhabdovirus" (suggested)[28]
"Drosophila affinis sigmavirus" (proposed)[27]
"Drosophila ananassae sigmavirus" (proposed)[27]
"Drosophila immigrans sigmavirus" (proposed)[27]
"Drosophila melanogaster sigmavirus"* (proposed)[27]
"Drosophila obscura sigmavirus" (proposed)[27]
"Drosophila tristis sigmavirus" (proposed)[27]
"Muscina stabulans sigmavirus" (proposed)[27]
"Sinistar-like viruses" (suggested)[26][29] "Siniperca chuatsi rhabdovirus"* (suggested)[29]
"TIBV/CPV-like viruses" (suggested)[23] "Coastal Plains virus"* (suggested)[23]
"Tibrogargan virus" (suggested)[23]
Tupaia virus
Vesiculovirus "BeAn 157575 virus"" (tentative)
Calchaqui virus
Carajas virus
Chandipura virus
Cocal virus
"Eel virus American" (tentative)
"eel virus European X" (suggested)
"Grey Lodge virus" (tentative)
Isfahan virus*
"Jurona virus" (tentative)
"Klamath virus" (tentative)
"Kwatta virus" (tentative)
"La Joya virus" (tentative)
"Lake trout rhabdovirus" (suggested)
"Malpais Spring virus" (tentative)
Maraba virus
"Mount Elgon bat virus" (tentative)
"Perinet virus" (tentative)
"Pike fry rhabdovirus" (tentative)
Piry virus
"Porton virus" (tentative)
"Radi virus" (tentative)
"Sea trout rhabdovirus" (suggested)
Spring viremia of carp virus
"Ulcerative disease rhabdovirus" (tentative)
Vesicular stomatitis Alagoas virus
Vesicular stomatitis Indiana virus*
Vesicular stomatitis New Jersey virus
"Yug Bogdanovac virus" (tentative)
Unassigned "Durham virus" (suggested)[30]
Moussa virus (accepted)[31]
Ngaingan virus

Table legend: "*" denotes type species; "suggested" refers to taxa that have been suggested by individual researchers but that have not been formally proposed to the ICTV; "proposed" refers to taxa that have been formally proposed; "accepted" refers to taxa that have been accepted by the Executive Committee of the ICTV but that have yet to be ratified; and "tentative" refers to tentative taxa listed in the most recent (Eighth) ICTV Report.

Life cycle

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. 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. 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.[6]


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.[32][33][34] Filovirus gene "fossils" have been detected in the genomes of bats, rodents, shrews, tenrecs, and marsupials.[35][33][34] A Midway virus "fossil" was found in the genome of zebrafish.[33]. Finally, rhabdovirus "fossils" were found in the genomes of mosquitoes and ticks.[34]


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