Abstract
In 2016, the order Mononegavirales was emended through the addition of two new families (Mymonaviridae and Sunviridae), the elevation of the paramyxoviral subfamily Pneumovirinae to family status (Pneumoviridae), the addition of five free-floating genera (Anphevirus, Arlivirus, Chengtivirus, Crustavirus, and Wastrivirus), and several other changes at the genus and species levels. This article presents the updated taxonomy of the order Mononegavirales as now accepted by the International Committee on Taxonomy of Viruses (ICTV).
Keywords: Anphevirus, Arlivirus, Bornaviridae, Chengtivirus, Crustavirus, Filoviridae, ICTV, International Committee on Taxonomy of Viruses, mononegavirad, Mononegavirales, mononegavirus, Mymonaviridae, Nyamiviridae, Paramyxoviridae, Pneumoviridae, Rhabdoviridae, Sunviridae, virus classification, virus nomenclature, virus taxonomy
INTRODUCTION
The viral order Mononegavirales was established in 1991 to accommodate related viruses with nonsegmented, linear, single-stranded negative-sense RNA genomes. These viruses were initially assigned to three mononegaviral families: Filoviridae, Paramyxoviridae, and Rhabdoviridae [20, 21]. In subsequent years, these families continued to grow through the inclusion of numerous novel species and genera, and the order was therefore emended in 1995 [4], 1997 [23], 2000 [24], 2005 [25], and 2011 [8]. The families Bornaviridae and Nyamiviridae joined the other three mononegaviral families in 1996 [22] and 2014 [1, 11], respectively. In 2015, the Study Groups of the International Committee on Taxonomy of Viruses (ICTV) responsible for the taxonomy of the order and its five families embarked on a joint effort to assign unclassified mononegaviruses to existing or novel taxa and to streamline order nomenclature. Here we present a brief overview of the first round of these efforts, which by now is accepted by the ICTV Executive Committee and, thereby, is official taxonomy.
TAXONOMIC CHANGES AT THE ORDER LEVEL
In recent years, several mononegaviruses have been described that are only distantly related to the members of the families Bornaviridae, Filoviridae, Nyamiviridae, Paramyxoviridae, and Rhabdoviridae. These viruses include Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1) found in an ascomycete in China [16]; Sunshine Coast virus (SunCV; previously called Sunshine virus) isolated from Australian carpet pythons [10]; and Líshí spider virus 2 (LsSV-2), Sānxiá water strider virus 4 (SxWSV-4), Tǎchéng tick virus 6 (TcTV-6), Wēnzhōu crab virus 1 (WzCV-1), and Xīnchéng mosquito virus (XcMV) detected in Chinese arthropods [15]. To accommodate these viruses in the order and to appropriately reflect their phylogenetic relationships to other mononegaviral taxa, two new families and four free-floating genera were established: Mymonaviridae (accommodating SsNSRV-1), Sunviridae (SunCV), Anphevirus (XcMV), Arlivirus (LsSV-2), Chengtivirus (TcTV-6), Crustavirus (WzCV-1), and Wastrivirus (SxWSV-4). In addition, the paramyxoviral subfamily Pneumovirinae was elevated to family status (Pneumoviridae) because the members of this taxon are as closely related to filoviruses as to the members of the paramyxoviral subfamily Paramxyovirinae (now dissolved) (Table 1).
Table 1. Taxonomy of the order Mononegavirales as of 2016.
Listed are all mononegaviruses that have been classified into species. Asterisks denote type species.
| Genus | Species | Virus (Abbreviation) |
|---|---|---|
| Family Bornaviridae | ||
| Bornavirus | Elapid 1 bornavirus | Loveridge’s garter snake virus 1 (LGSV-1) |
| Mammalian 1 bornavirus* | Borna disease virus 1 (BoDV-1) | |
| Borna disease virus 2 (BoDV-2) | ||
| Passeriform 1 bornavirus | canary bornavirus 1 (CnBV-1) | |
| canary bornavirus 2 (CnBV-2) | ||
| canary bornavirus 3 (CnBV-3) | ||
| Passeriform 2 bornavirus | estrildid finch bornavirus 1 (EsBV-1) | |
| Psittaciform 1 bornavirus | parrot bornavirus 1 (PaBV-1) | |
| parrot bornavirus 2 (PaBV-2) | ||
| parrot bornavirus 3 (PaBV-3) | ||
| parrot bornavirus 4 (PaBV-4) | ||
| parrot bornavirus 7 (PaBV-7) | ||
| Psittaciform 2 bornavirus | parrot bornavirus 5 (PaBV-5) | |
| Waterbird 1 bornavirus | aquatic bird bornavirus 1 (ABBV-1) | |
| aquatic bird bornavirus 2 (ABBV-2) | ||
| Family Filoviridae | ||
| Cuevavirus | Lloviu cuevavirus* | Lloviu virus (LLOV) |
| Ebolavirus | Bundibugyo ebolavirus | Bundibugyo virus (BDBV) |
| Reston ebolavirus | Reston virus (RESTV) | |
| Sudan ebolavirus | Sudan virus (SUDV) | |
| Taï Forest ebolavirus | Taï Forest virus (TAFV) | |
| Zaire ebolavirus* | Ebola virus (EBOV) | |
| Marburgvirus | Marburg marburgvirus* | Marburg virus (MARV) |
| Ravn virus (RAVV) | ||
| Family Mymonaviridae | ||
| Sclerotimonavirus | Sclerotinia sclerotimonavirus* | Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1) |
| Family Nyamiviridae | ||
| Nyavirus | Midway nyavirus | Midway virus (MIDWV) |
| Nyamanini nyavirus* | Nyamanini virus (NYMV) | |
| Sierra Nevada nyavirus | Sierra Nevada virus (SNVV) | |
| Socyvirus | Soybean cyst nematode socyvirus* | soybean cyst nematode virus 1 (SbCNV-1) |
| Family Paramyxoviridae | ||
| Aquaparamyxovirus | Atlantic salmon paramyxovirus* | Atlantic salmon paramyxovirus (AsaPV) |
| Avulavirus | Avian paramyxovirus 2 | avian paramyxovirus 2 (APMV-2) |
| Avian paramyxovirus 3 | avian paramyxovirus 3 (APMV-3) | |
| Avian paramyxovirus 4 | avian paramyxovirus 4 (APMV-4) | |
| Avian paramyxovirus 5 | avian paramyxovirus 5 (APMV-5) | |
| Avian paramyxovirus 6 | avian paramyxovirus 6 (APMV-6) | |
| Avian paramyxovirus 7 | avian paramyxovirus 7 (APMV-7) | |
| Avian paramyxovirus 8 | avian paramyxovirus 8 (APMV-8) | |
| Avian paramyxovirus 9 | avian paramyxovirus 9 (APMV-9) | |
| Avian paramyxovirus 10 | avian paramyxovirus 10 (APMV-10) | |
| Avian paramyxovirus 11 | avian paramyxovirus 11 (APMV-11) | |
| Avian paramyxovirus 12 | avian paramyxovirus 12 (APMV-12) | |
| Newcastle disease virus* | avian paramyxovirus 1 (APMV-1)1 | |
| Ferlavirus | Fer-de-Lance paramyxovirus* | Fer-de-Lance virus (FDLV)2 |
| Henipavirus | Cedar henipavirus | Cedar virus (CedV) |
| Ghanaian bat henipavirus | Kumasi virus (KV)3 | |
| Hendra virus* | Hendra virus (HeV) | |
| Mojiang henipavirus | Mòjiâng virus (MojV) | |
| Nipah virus | Nipah virus (NiV) | |
| Morbillivirus | Canine distemper virus | canine distemper virus (CDV) |
| Cetacean morbillivirus | cetacean morbillivirus (CeMV) | |
| Feline morbillivirus | feline morbillivirus (FeMV)4 | |
| Measles virus* | measles virus (MeV) | |
| Peste-des-petits-ruminants virus | peste-des-petits-ruminants virus (PPRV) | |
| Phocine distemper virus | phocine distemper virus (PDV) | |
| Rinderpest virus | rinderpest virus (RPV) | |
| Respirovirus | Bovine parainfluenza virus 3 | bovine parainfluenza virus 3 (BPIV-3) |
| Human parainfluenza virus 1 | human parainfluenza virus 1 (HPIV-1) | |
| Human parainfluenza virus 3 | human parainfluenza virus 3 (HPIV-3)5 | |
| Porcine parainfluenza virus 1 | porcine parainfluenza virus 1 (PPIV-1) | |
| Sendai virus* | Sendai virus (SeV)6 | |
| Rubulavirus | Human parainfluenza virus 2 | human parainfluenza virus 2 (HPIV-2) |
| Human parainfluenza virus 4 | human parainfluenza virus 4a (HPIV-4a) | |
| human parainfluenza virus 4b (HPIV-4b) | ||
| Mapuera virus | Mapuera virus (MapV) | |
| Mumps virus* | mumps virus (MuV) | |
| bat mumps virus (BMV)7 | ||
| Parainfluenza virus 5 | parainfluenza virus 5 (PIV-5)8 | |
| Porcine rubulavirus | La Piedad Michoacán Mexico virus (LPMV)9 | |
| Simian virus 41 | simian virus 41 (SV-41) | |
| Family Pneumoviridae | ||
| Metapneumovirus | Avian metapneumovirus* | avian metapneumovirus (AMPV)10 |
| Human metapneumovirus | human metapneumovirus (HMPV) | |
| Orthopneumovirus | Bovine respiratory syncytial virus | bovine respiratory syncytial virus (BRSV) |
| Human respiratory syncytial virus* | human respiratory syncytial virus A2 (HRSV-A2) | |
| human respiratory syncytial virus B1 (HRSV-B1) | ||
| human respiratory syncytial virus S2 (HRSV-S2) | ||
| Murine pneumonia virus | murine pneumonia virus (MPV) | |
| Family Rhabdoviridae | ||
| 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) | |
| Festuca leaf streak cytorhabdovirus | festuca leaf streak virus (FLSV) | |
| Lettuce necrotic yellows cytorhabdovirus* | lettuce necrotic yellows virus (LNYV) | |
| Lettuce yellow mottle cytorhabdovirus | lettuce yellow mottle virus (LYMoV) | |
| Northern cereal mosaic cytorhabdovirus | northern cereal mosaic virus (NCMV) | |
| Sonchus cytorhabdovirus 1 | sonchus virus (SonV) | |
| Strawberry crinkle cytorhabdovirus | strawberry crinkle virus (SCV) | |
| Wheat American striate mosaic cytorhabdovirus | wheat American striate mosaic virus (WASMV) | |
| Dichorhavirus | Coffee ringspot dichorhavirus | coffee ringspot virus (CoRSV) |
| Orchid fleck dichorhavirus* | orchid fleck virus (OFV)11 | |
| Ephemerovirus | Adelaide River ephemerovirus | Adelaide River virus (ARV) |
| Berrimah ephemerovirus | Berrimah virus (BRMV) | |
| Bovine fever ephemerovirus* | bovine ephemeral fever virus (BEFV)12 | |
| Kotonkan ephemerovirus | kotonkan virus (KOTV) | |
| Obodhiang ephemerovirus | Obodhiang virus (OBOV) | |
| 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) | |
| Ikoma lyssavirus | Ikoma lyssavirus (IKOV) | |
| Irkut lyssavirus | Irkut virus (IRKV) | |
| Khujand lyssavirus | Khujand virus (KHUV) | |
| Lagos bat lyssavirus | Lagos bat virus (LBV) | |
| Mokola lyssavirus | Mokola virus (MOKV) | |
| Rabies lyssavirus* | rabies virus (RABV) | |
| Shimoni bat lyssavirus | Shimoni bat virus (SHIBV) | |
| West Caucasian bat lyssavirus | West Caucasian bat virus (WCBV) | |
| Novirhabdovirus | Hirame novirhabdovirus | Hirame rhabdovirus (HIRV) |
| Oncorhynchus 1 novirhabdovirus* | infectious hematopoietic necrosis virus (IHNV) | |
| Oncorhynchus 2 novirhabdovirus | viral hemorrhagic septicemia virus (VHSV)13 | |
| 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) | ||
| Tibrovirus | Coastal Plains tibrovirus | Coastal Plains virus (CPV) |
| Tibrogargan tibrovirus* | Bivens Arm virus (BAV) | |
| Tibrogargan virus (TIBV) | ||
| Tupavirus | Durham tupavirus* | Durham virus (DURV) |
| Tupaia tupavirus | tupaia virus (TUPV) | |
| Varicosavirus | Lettuce big-vein associated varicosavirus* | lettuce big-vein associated virus (LBVaV)14 |
| Vesiculovirus | Alagoas vesiculovirus | vesicular stomatitis Alagoas virus (VSAV) |
| Carajas vesiculovirus | Carajás virus (CJSV) | |
| Chandipura vesiculovirus | Chandipura virus (CHPV) | |
| Cocal vesiculovirus | Cocal virus (COCV) | |
| Indiana vesiculovirus* | vesicular stomatitis Indiana virus (VSIV) | |
| Isfahan vesiculovirus | Isfahan virus (ISFV) | |
| Maraba vesiculovirus | Maraba virus (MARAV) | |
| New Jersey vesiculovirus | vesicular stomatitis New Jersey virus (VSNJV) | |
| Piry vesiculovirus | Piry virus (OIRYV) | |
| Unassigned | Flanders virus | Flanders virus (FLAV) |
| Ngaingan virus | Ngaingan virus (NGAV) | |
| Wongabel virus | Wongabel virus (WONV) | |
| Family Sunviridae | ||
| Sunshinevirus | Reptile sunshinevirus 1* | Sunshine Coast virus (SunCV) |
| Unassigned | ||
| Anphevirus | Xinchng anphevirus* | Xînchéng mosquito virus (XcMV) |
| Arlivirus | Lishi arlivirus* | Líshí spider virus 2 (LsSV-2) |
| Chengtivirus | Tacheng chengtivirus* | Tãchéng tick virus 6 (TcTV-6) |
| Crustavirus | Wenzhou crustavirus* | Wçnzhôu crab virus 1 (WzCV-1) |
| Wastrivirus | Sanxia wastrivirus* | Sânxiá water strider virus 4 (SxWSV-4) |
Includes: Newcastle disease virus (NDV) and pigeon paramyxovirus;
synonym: anaconda paramyxovirus;
synonym: GH-M74a virus;
abbreviation as recently introduced in [26];
historically, an isolate from a samango monkey (Cercopithecus mitis) in 1963 was long classified as a distinct species called simian agent 10 (SA-10), but was shown later to be HPIV-3. SA-10 was sometimes called SV-10;
synonym: murine parainfluenza virus 1;
synonym: bat paramyxovirus;
synonym: simian virus 5;
synonym: porcine rubulavirus;
synonyms: avian pneumovirus, turkey rhinotracheitis virus;
synonyms: citrus leprosis virus nuclear type, citrus necrotic spot virus;
synonym Tzipori virus;
synonyms: Egtved virus, Paralichthys olivaceus rhabdovirus;
synonym: tobacco stunt virus.
TAXONOMIC CHANGES AT THE FAMILY LEVEL
The monogeneric family Bornaviridae was reorganized in 2015 by establishing five distinct species in the genus Bornavirus [2, 12] following a non-Latinized binomial species name format [29]. These efforts were continued in 2016 by expanding the genus by an additional two species (Elapid 1 bornavirus for Loveridge’s garter snake virus 1 [27] and Psittaciform 2 bornavirus for parrot bornavirus 5 [9, 18]) (Table 1).
The monogeneric family Nyamiviridae was expanded to include a second genus (Socyvirus) for the until-then free-floating nyamivirus species Soybean cyst nematode virus. This species name was changed to Soybean cyst nematode socyvirus to adhere to the non-Latinized binomial species name format [29] (Table 1).
The family Paramyxoviridae was emended by expanding the genus Avulavirus by three species (Avian paramyxovirus 10–12 for avian paramyxoviruses 10–12, respectively [5, 19, 28]), the genus Henipavirus by three species (Cedar henipavirus for Cedar virus [17], Ghanaian bat henipavirus for Kumasi virus [GH-M74a] [7], and Mojiang henipavirus for Mòjiāng virus [31]), the genus Morbillivirus by one species (Feline morbillivirus for feline morbillivirus [30]) and the genus Respirovirus by one species (Porcine parainfluenza virus 1 for porcine parainfluenza virus 1 [14]). The species Simian Virus 10 was dissolved on the evidence that simian virus 10 is an isolate of human parainfluenzavirus 3 rather than a distinct virus [13]. The genus Pneumovirus, now included in the new family Pneumoviridae, was renamed Orthopneumovirus to avoid confusion between family and genus members (Table 1).
The family Rhabdoviridae was expanded by two genera: Dichorhavirus (new; [6]) and Varicosavirus (previously free-floating outside of the order) to accommodate bisegmented plant viruses (coffee ringspot virus and orchid fleck virus; lettuce big-vein associated virus). The species Alfalfa dwarf cytorhabdovirus (for alfalfa dwarf virus [3]) was added to the genus Cytorhabdovirus. Finally, the non-Latinized binomial species name format [29] was applied throughout the family (Table 1).
A summary of the current, ICTV-accepted taxonomy of the order Mononegavirales is presented in Table 1.
Acknowledgments
We thank Laura Bollinger (NIH/NIAID Integrated Research Facility at Fort Detrick, Frederick, MD, USA) for critically editing the manuscript.
Funding
This work was supported in part through Battelle Memorial Institute’s prime contract with the US National Institute of Allergy and Infectious Diseases (NIAID) under Contract No. HHSN272200700016I. A subcontractor to Battelle Memorial Institute who performed this work is: J.H.K., an employee of Tunnell Government Services, Inc. This work was also funded in part under Contract No. HSHQDC-07-C-00020 awarded by DHS S&T for the management and operation of the National Biodefense Analysis and Countermeasures Center (NBACC), a Federally Funded Research and Development Center (V.W.-J.); and National Institutes of Health (NIH) contract HHSN272201000040I/HHSN27200004/D04 (N.V., R.B.T.). Y.B. was supported by the Intramural Research Program of the NIH, National Library of Medicine.
Footnotes
COMPLIANCE WITH ETHICAL STANDARDS
The views and conclusions contained in this document are those of the authors and should not be interpreted as necessarily representing the official policies, either expressed or implied, of the US Department of the Army, the US Department of Defense, the US Department of Health and Human Services, the Department of Homeland Security (DHS) Science and Technology Directorate (S&T) or of the institutions and companies affiliated with the authors. In no event shall any of these entities have any responsibility or liability for any use, misuse, inability to use, or reliance upon the information contained herein. The US departments do not endorse any products or commercial services mentioned in this publication.
Conflict of Interest
The authors have no conflicts of interest.
Ethical approval
This article does not contain any studies with human participants or animals performed by any of the authors.
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