Virus TaxonomyThe ICTV Report on Virus Classification and Taxon Nomenclature

Potyviridae Chapter

Potyviridae

Stephen J Wylie, Alice Kazuko Inoue-Nagata, Jan Kreuze, Juan José López-Moya, Kristiina Mäkinen, Kazusato Ohshima andAiming Wang

Edited by Hélène Sanfaçon and Michael J. Adams

Corresponding author: Stephen J Wylie (s.wylie@murdoch.edu.au)

Posted March 2017, updated October 2018

PDF created: October 2020

Citation

A summary of this ICTV Report chapter has been published as an ICTV Virus Taxonomy Profile article in the Journal of General Virology, andshould be cited when referencing this online chapter as follows:

Wylie, S.J., Adams, M., Chalam, C., Kreuze, J., López-Moya, J.J., Ohshima, K., Praveen, S., Rabenstein, F., Stenger, D., Wang, A., Zerbini,F.M., and ICTV Report Consortium, ICTV Virus Taxonomy Profile: Potyviridae, Journal of General Virology, 98: 352–354.

Summary

Members of family Potyviridae (termed potyvirids) consist of monopartite and bipartite plant viruses with a single-stranded, positive-senseRNA genome and flexuous, filamentous particles (Table 1.Potyviridae). Genomes have a VPg covalently linked to the 5′- end and the 3′-terminus is polyadenylated. Genomes encode a large polyprotein that is self-cleaved into a set of functional proteins. Gene order is generallyconserved throughout the family (Adams et al., 2005b).

Table 1.Potyviridae. Characteristics of members of the family Potyviridae.

Characteristic Description

Typicalmember potato virus Y-O (U09509), species Potato virus Y, genus Potyvirus

Virion Non-enveloped, flexuous and filamentous capsid, 680–900 nm long and 11–20 nm in diameter with a single core capsidprotein

Genome 8–11 kb of positive-sense, single-stranded, usually monopartite RNA (bipartite in genus Bymovirus)

Replication Cytoplasmic, initiated in virus replication complexes on membranous vesicles at ER exit sites. Replication initiates at 6K2-induced ER-originated vesicles

Translation Directly from genomic RNA.

Host range Plants (all virus genera). Most members are arthropod-borne but those of genus Bymovirus are transmitted byplasmodiophorids

Taxonomy 10 genera including over 200 species

Virion

Morphology

Virions are flexuous filaments with no envelope and are 11–20 nm in diameter, with a helical pitch of about 3.4 nm (Figure 1. Potyviridae).Particle lengths of members of some of the ten genera differ. Members of the genera Potyvirus, Poacevirus, Ipomovirus, Macluravirus,Rymovirus, Tritimovirus, Brambyvirus, Bevemovirus, Roymovirus and of the unassigned virus species Spartina mottle virus are monopartitewith particle modal lengths of 650–950 nm. Members of the genus Bymovirus are bipartite with particles of two modal lengths of 250–300 and500–600 nm.

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Figure 1.Potyviridae. (Left) Schematic diagram of a potyvirus particle. The N-terminus (ca. 30 aa; large rectangle) and C-terminus (ca.19aa; small rectangle) of the coat (capsid) protein is exposed on the surface of the intact virus particle (from (Shukla and Ward 1989)). (Right)Negative-contrast electron micrograph of particles of an isolate of Plum pox virus , stained with 1% PTA, pH 6.0. The bar represents 200 nm(Courtesy of I.M. Roberts.)

Physicochemical and physical properties

Virions of viruses in the genera Potyvirus and Rymovirus have a density in CsCl of about 1.31 g cm and Sedimentation coefficient S of137–160S. Those of the genus Bymovirus have a density in CsCl of about 1.29 g cm .

Nucleic acid

Viruses in all genera except Bymovirus have a single molecule of positive-sense, ssRNA, 8.2 kb (bellflower veinal mottle virus-SW) to 11.0 kb(sweet potato feathery mottle virus-Piu3) in size. Virions are infectious. A VPg of about 24 kDa is covalently linked to the 5′-terminalnucleotide. A polyadenylate tract (20 to 160 adenosines) is present at the 3′-terminus. Bymoviruses have two positive-sense, ssRNAmolecules; RNA1 is 7.3–7.6 kb and RNA2 is 3.5–3.7 kb. Both RNAs have 3′-terminal polyadenylate tracts and probably a VPg at the 5′-termini.

Proteins

Virions contain one type of coat (capsid) protein (CP) of 28.5–47 kDa. N- and C-terminal residues are positioned on the exterior of the virion.Mild trypsin treatment removes N- and C-terminal segments, leaving a trypsin-resistant core of about 24 kDa. Plant proteases may degradethe CP in vivo, as occurs in vitro during purification using some procedures or from certain hosts. All potyvirus CPs display significant aasequence identity in the trypsin-resistant core, but little identity in their N and C-terminal segments.

Genome organization and replication

The genomic RNA (two RNAs for members of the genus Bymovirus) encodes a single major polyprotein. This then undergoes co- and post-translational proteolytic processing by three viral-encoded proteinases to form the mature proteins. Genomic RNA replicates via theproduction of a full-length negative-sense RNA. While there are exceptions noted in the relevant genus descriptions, the polyprotein of themajority of monopartite viruses in the family is cleaved into ten products, which show conservation of sequence and organisation (Kekarainenet al., 2002). As shown in Figure 2.Potyviridae, these products are:

P1 (Protein 1): Of all the potyvirid proteins, P1 is the least conserved in sequence and the most variable in size. It plays a significant rolein virus replication probably due to the stimulation of the gene silencing suppressor HC-Pro. A serine protease domain towards the C-terminus cleaves the P1 from the polyprotein, typically at Tyr/Phe-Ser (Valli et al., 2007).HC-Pro (Helper Component-Protease): the HC-Pro protein has roles in suppression of gene silencing and in vector transmission. Acysteine protease domain towards the C-terminus cleaves it from the remainder of the downstream polyprotein, typically at Gly-Gly.P3 (Protein 3): Involved in virus replication and appears to be significant in host range and symptom development.6K1 (6-kDa peptide 1): Although it is present in the replication complex and required for replication, the function of this small protein isnot known.CI (Cylindrical Inclusion protein): This protein has helicase activity and accumulates in inclusion bodies in the cytoplasm of infected plantcells.6K2 (6-kDa peptide 2): A small transmembrane protein probably anchoring the replication complex to the ER.VPg (Viral Protein genome-linked): Attached to the 5′-terminus of the genome and belongs to a class of intrinsically disordered proteins.It is essential for virus replication and translation, interacting with one or several isoforms of the eIF4E translation initiation factor, and isalso involved in suppression of RNA silencing.NIa-Pro (Nuclear inclusion A-protease): Serine-like cysteine protease responsible for cleavage of most sites in the polyprotein, typicallyat Gln/Glu-Ser/Gly/Ala (Adams et al., 2005a).NIb (Nuclear inclusion B): The RNA-dependent RNA polymerase.CP (Coat (capsid) protein): Viral coat protein that also has roles in virus movement, genome amplification and vector transmission.PIPO (Pretty Interesting Potyvirus ORF): A short ORF embedded within the P3 cistron expressed as the trans-frame P3N-PIPO protein

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by a polymerase slippage mechanism (Chung et al., 2008, Olspert et al., 2015, Rodamilans et al., 2015). PIPO has been identifiedthroughout the family and has been shown to be essential for the intercellular movement of viruses.

The polymerase slippage model for the production of P3N-PIPO is consistent with the theory that potyvirus intercellular movement is coupledto active virus genome replication and translation. Potyvirus replication initiates at the 6K2-induced ER-originated vesicles. The cellular COPIand COPII coating machineries are involved in the biogenesis of the potyvirus 6K2 vesicles (Wei and Wang 2008). The ER-derived 6K2vesicles also target and are associated with chloroplasts for virus replication.

Figure 2.Potyviridae. Genomic map of a typical member of the genus Potyvirus. The ssRNA genome is represented by a line and thepolyprotein ORF by an open box with the mature proteolytic products named. VPg (viral protein genome-linked), the genome-linked viralprotein covalently attached to the 5′-terminal nucleotide is represented by a hexagon; P1-Pro (protein 1 protease), a protein with serineproteolytic activity responsible for cleavage at typically Tyr/Phe-Ser (O); HC-Pro (helper component protease), a protein with aphidtransmission helper-component activity and cysteine proteolytic activity responsible for cleavage at typically Gly-Gly (◆); P3 (protein 3);PIPO (pretty interesting Potyviridae ORF); 6K (six kilodalton peptide); CI (cytoplasmic inclusion); NIa-Pro (nuclear inclusion A protease),cysteine-like proteolytic activity responsible for cleavage at Gln/Glu-(Ser/Gly/Ala) (↓); Nib (nuclear inclusion B), RNA-dependent RNApolymerase; CP (coat protein). Cleavage sites of P1-Pro, (O), HC-Pro (◆) and NIa-Pro (↓) are indicated.

Biology

Inclusion body formation

All members of the family Potyviridae form cytoplasmic cylindrical inclusion (CI) bodies during infection. The CI is an array of a 70 kDa viralprotein that possesses ATPase and helicase activities. Some potyviruses induce nuclear inclusion bodies that are co-crystals of two viral-encoded proteins – NIa and NIb – that are present in equimolar amounts. The small nuclear inclusion (NIa) protein (49 kDa) is a polyproteinconsisting of VPg and NIa-Pro. Amorphous inclusion bodies are also evident in the cytoplasm during certain potyvirus infections andrepresent aggregations of HC-Pro and perhaps other non-structural proteins.

Host range

Some members have a narrow host range, most members infect an intermediate number of plants, and a few members infect species in up to30 families. Transmission to most hosts is readily accomplished by mechanical inoculation. Many viruses are widely distributed. Distribution isaided by seed transmission in some cases.

Transmission

Potyvirids are vectored by a variety of organisms. Members of the genera Potyvirus and Macluravirus have aphid vectors that transmit in anon-persistent, non-circulative manner. Two helper component protease aa motifs (R/KITC and PTK) and a CP motif (DAG for manypotyviruses) are highly conserved and required for aphid transmission. Viruses in other genera in the Potyviridae lacking these motifs are nottransmitted by aphids, and include rymoviruses, poaceviruses and tritimoviruses which are transmitted by eriophyid mites in a semi-persistent manner. The vector of rose yellow mosaic virus (RoYMV), the type species of the monotypic genus Roymovirus, has not beenidentified. The conserved potyvirus HC-Pro motifs are not present. Instead, a putative C-2x-C eriophyid mite transmission motif occurs at theN-terminus of the HC-Pro. The DAG motif of the CP is lacking. Together this information suggests RoYMV may also be transmitted byeriophyid mites, although this has not been proven experimentally. Bymoviruses are transmitted by root-infecting vectors in the orderPlasmodiophorales, once described as fungi but now classified as Cercozoa. Ipomoviruses appear to be transmitted by whiteflies, and thevector of the one species within Brambyvirus is unknown. The vector of bellflower veinal mottle virus (BVMV), the type species of themonotypic genus Bevemovirus, has not been identified. The HC-Pro of BVMV lacks the conserved potyvirus aphid transmission motifs, butthe CP has a DTG near its N-terminus, possibly analogous to the DAG motif involved in aphid transmission.

Antigenicity

An epitope of the CP in the conserved internal trypsin-resistant core has been identified that is similar in most members of the family.

Derivation of names

Potyviridae: from the type species of the genus Potyvirus, Potato virus Y.

Genus demarcation criteria

The ten genera are differentiated by biological criteria, mainly transmission by specific vectors, and by molecular data (Figure 3. Potyviridae)(Adams et al., 2005b, Gibbs and Ohshima 2010). For the entire open reading frame, genus demarcation criteria are <46% nucleotide identity,

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but this does not separate rymoviruses from potyviruses, which have different vectors.

Figure 3.Potyviridae. Pairwise unrooted neighbor-joining tree of complete polyprotein sequences of representative viruses within thefamily Potyviridae. The tree was produced in MEGA 7 (Kumar et al., 2016) using the JTT and gamma rate variation options from aCLUSTAL (Larkin et al., 2007). alignment of polyprotein sequences. Branches supported by > 70% of 100 bootstrap replicates areindicated. This phylogenetic tree and corresponding sequence alignment are available to download from the Resources page.

Species demarcation criteria

Species demarcation criteria for the complete ORF are <76% nucleotide identity and <82% amino acid identity. The thresholds for speciesdemarcation using nucleotide identity values for the individual coding regions range from 58% for the P1 coding region to 74–78% for otherregions. For the coat protein, the optimal species demarcation criterion is 76–77% nucleotide and 80% amino acid identity (Adams et al.,2005b).

Relationships with other taxa

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Members of the family Potyviridae are related to viruses in the order Picornavirales in their genome structure, in particular the presence of asingle ORF translated into a polyprotein and the block of replication genes (helicase, VPg, 3C-like protease and polymerase). However, theydiffer from picornaviruses in virion morphology, the size of VPg and the type of helicase.

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Genus: Bevemovirus

Distinguishing features

Like macluraviruses, members of the type species, Bellflower veinal mottle virus , lack the P1 region of most potyvirids. The HC-Pro lacks theconserved potyvirus aphid transmission motifs R/KITC and PTK. Isolates of bellflower veinal mottle virus share low (22–27%) nucleotide andamino acid sequence identities to macluraviruses.

Virion

Nucleic acid

Bellflower veinal mottle virus has a positive-sense RNA genome of 8,259 nt.

Genome organization and replication

Genome organization resembles that of members of the genus Macluravirus.

Biology

Isolates of Bellflower veinal mottle virus were obtained from bellflower (Campanula takesimana) that exhibited veinal mottle symptoms inSouth Korea. The virus is probably not aphid transmitted because the HC-Pro lacks the conserved potyvirus aphid transmission motifsR/KITC and PTK.

Derivation of names

Bevemovirus: from bellflower veinal mottle virus

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Bellflower veinal mottle virus bellflower veinal mottle virus SW KY491536 NC_039002 Complete genome BVMoV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Brambyvirus

Distinguishing features

The genus includes a single species, members of which are distinguished from all other members of the family in that they encode a verylarge P1 protein (83.6 k Da) containing an AlkB domain, and are also phylogenetically distinct (Susaimuthu et al., 2008).

Virion

Morphology

Virions are flexuous filaments 800×11–15 nm in size.

Nucleic acid

Virions contain a single molecule of linear, positive-sense ssRNA of about 11 kb.

Proteins

There is a single coat (capsid) protein of 40.9 kDa.

Genome organization and replication

Apart from the size of the P1 coding region, the genome organization is identical to that of most monopartite viruses in the family Potyviridae(Figure 2.Potyviridae).

Biology

Host range

The virus has been reported only from wild and cultivated blackberry ( Rubus sp.) where it is often symptomless but is also a component of acomplex of viruses. It is not known to cause symptoms in any herbaceous test host.

Transmission

The virus is presumed to be transmitted by an aerial vector that has not yet been identified.

Antigenicity

The virus could not be detected by a universal potyvirus monoclonal antibody but there are no additional data.

Derivation of names

Brambyvirus: from bramble, the host of blackberry virus Y, a member of the type species Blackberry virus Y

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Blackberry virus Y blackberry virus Y Ark3 AY994084 NC_008558 Complete genome BlVY

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Bymovirus

Distinguishing features

Compared with other viruses in the family, members of the genus Bymovirus are distinct in having a divided (bipartite) genome and in beingtransmitted by the root-infecting parasite, Polymyxa graminis (Plasmodiophorales) a fungoid protist.

Virion

Morphology

Virions are flexuous filaments of two modal lengths, 250–300 nm and 500-600 nm; both are 13 nm in width (Figure 1.Bymovirus).

Figure 1.Bymovirus. Virions of an isolate of barley yellow mosaic virus, stained with 1% PTA, pH 7.0. The bar represents 200 nm (from D.Lesemann).

Physicochemical and physical properties

Virion density in CsCl is 1.28–1.30 g cm .

Nucleic acid

Virions contain two molecules of linear positive-sense, ssRNA. RNA1 is 7.5–8.0 kb and RNA2 is 3.5–4.0 kb; RNA makes up 5% by weight ofparticles. There is little sequence identity between the two RNAs except in the 5′-untranslated region.

Proteins

Virions have a single coat (capsid) protein (CP) of 28.5–33 kDa. The CP of barley yellow mosaic virus isolates has 297 aa.

Genome organization and replication

The two RNA molecules are translated initially into precursor polypeptides from which functional proteins are derived by proteolyticprocessing (Figure 2.Bymovirus). The organization of RNA1 is similar to that of other potyviruses but without the P1 and HC-Pro proteins. TheRNA2 polyprotein is unique to bymoviruses although the first protein encoded by RNA2 (P1, ca. 28 kDa) has aa domains with sequencesimilarities to the potyvirus protein HC-Pro. The larger protein of RNA2 (P2) is believed to have a role in vector transmission (You and Shirako2010).

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Figure 2.Bymovirus. Genomic map of the bymovirus bipartite genome, using, as an example an isolate of barley yellow mosaic virus.Conventions are as for potyvirus genome organization map (Figure 2.Potyviridae). Function of most mature proteins are postulated byanalogy with members of the genus Potyvirus. P1 corresponds to the C-terminal protease of HC-Pro.

Biology

Cytology

There are characteristic pinwheel-like inclusions and membranous network structures are formed in the cytoplasm of infected plant cells. Nonuclear inclusions are found.

Host range

The host range of bymoviruses is narrow, restricted to the host family Gramineae. Each member has a very restricted host range; forexample, the barley-infecting viruses do not infect wheat and vice versa.

Transmission

Bymoviruses are transmitted by Polymyxa graminis in a persistent manner, surviving in resting spores as long as these remain viable; theyare transmissible experimentally by mechanical inoculation, sometimes with difficulty.

Antigenicity

The viral proteins are moderately immunogenic; apart from barley mild mosaic virus, most members of the genus are serologically related.The CP aa sequence identity among members is 35–74%.

Derivation of names

Bymovirus: from barley yellow mosaic virus

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available

sequenceVirus

Abbrev.★ Barley mild mosaic virus barley mild mosaic virus UK-F RNA1: Y10973; RNA2: X90904 RNA1: NC_003483; RNA2:

NC_003482 Complete genome BaMMV

★ Barley yellow mosaic virus barley yellow mosaic virus Yancheng

RNA1: AJ132268; RNA2:AJ132269

RNA1: NC_002990; RNA2:NC_002991 Complete genome BaYMV

★ Oat mosaic virus oat mosaic virus Cranbrook

RNA1: AJ306718; RNA2:AJ306719

RNA1: NC_004016; RNA2:NC_004017 Complete genome OMV

★ Rice necrosis mosaic virus rice necrosis mosaic virus Ka-1 RNA1: LC055681; RNA2:LC060925

RNA1: NC_028144; RNA2:NC_028145 Complete genome RNMV

★Wheat spindle streak mosaicvirus

wheat spindle streak mosaicvirus FR X73883 Partial genome WSSMV

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★Wheat yellow mosaic virus wheat yellow mosaic virus JA RNA1: D86634; RNA2: D86635RNA1: NC_002350; RNA2:NC_002349 Complete genome WYMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name Accessionnumber

Virusabbreviation

Soybean leaf rugose mosaicvirus AB560671* SLRMV

Virus names and virus abbreviations are not official ICTV designations.

* incomplete RNA1 genome sequence

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Genus: Ipomovirus

Distinguishing features

Ipomoviruses are distinguished from members of other genera by their mode of transmission being by whiteflies, and by their separatebranching on phylogenetic analyses.

Virion

Morphology

Virions are flexuous filaments 800–950 nm long.

Physicochemical and physical properties

Virion Sedimentation coefficient S is 155S for sweet potato mild mottle virus (SPMMV).

Nucleic acid

Virions contain a single molecule of linear, positive-sense ssRNA of about 9.7 kb; virions contain 5% RNA by weight.

Proteins

The viral coat (capsid) protein (CP) is a single polypeptide of 302–378 aa (35–41 kDa).

Genome organization and replication

Ipomoviruses exhibit unusual structural variability. The structure and organization of the SPMMV genome is similar to that of members of thegenus Potyvirus (Figure 2.Potyviridae), but some motifs of HC-Pro and CP characteristic of members of the genus Potyvirus are incompleteor missing, which may account for its vector relations. The unusually large P1 protein (83 kDa) of SPMMV contains no obvious AlkB domainand hence differs from that found in members of the genus Brambyvirus. Cucumber vein yellowing virus (CVYV) and squash vein yellowingvirus (SqVYV) differ from SPMMV by encoding two P1-like serine proteases (P1a and P1b) but no HC-Pro. P1b functions as a suppressor ofRNA silencing (Li et al., 2008). Cassava brown streak virus (CBSV) differs from SPMMV by having no HC-Pro and differs from CVYV andSqVYV by having only P1b which suppresses silencing (Figure 1.Ipomovirus). Additionally, cassava brown streak virus (CBSV) contains aMaf/HAM1-like sequence recombined into the NIb/CP junction, which can accommodate heterologous genes in engineered infectiouspotyvirus clones. Homology of HAM1h with cellular Maf/HAM1 NTP pyrophosphatases suggests that HAMh1 might intercept non-canonicalNTPs to reduce mutation rates of viral RNA.

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Figure 1.Ipomovirus. Genomic maps of the ipomoviruses sweet potato mild mottle virus (SPMMV), squash vein yellowing virus (SqVYV)and cassava brown streak virus (CBSV). The ssRNA genome is represented by a line and the polyprotein ORF by an open box.Conventions are as for the potyvirus genome organization map (Figure 2.Potyviridae). Activities of most mature proteins are postulated byanalogy with genus Potyvirus. CVYV and SqVYV contain two P1-like serine proteases (P1a and P1b), of which P1b functions as asuppressor of RNA silencing. CBSV also contain P1b which suppresses silencing and, additionally, carries a Maf/HAM1-like sequencerecombined into the NIb/CP junction. HAMh1 might intercept non-canonical NTPs to reduce mutation rates of viral RNA.

Biology

Host range

The natural host range of SPMMV is wide, with more than nine families susceptible, whereas the host range of CBSV, CVYV and SqVYV isless known apart from the hosts which they have been found to infect in the field (Winter et al., 2010).

Transmission

CBSV, CVYV and SqVYV are transmitted by the whitefly Bemisia tabaci in a non-persistent manner. B. tabaci may also be the vector ofSPMMV, but this is not fully confirmed. All ipomoviruses are transmissible experimentally by mechanical inoculation and by grafting.

Antigenicity

Moderately immunogenic. No serological relationships with other members of the family Potyviridae have been found.

Derivation of names

Ipomovirus: from Ipomea and mosaic, the symptoms induced on the host of members of the type species Sweet potato mild mottle virus .

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Cassava brown streak virus cassava brown streak virus KOR6 GU563327 NC_012698 Complete genome CBSV★ Coccinia mottle virus Coccinia mottle virus Su12-25 KU935732 NC_030840 Complete genome CocMoV★ Cucumber vein yellowing virus cucumber vein yellowing virus ALM32 AY578085 NC_006941 Complete genome CVYV★ Squash vein yellowing virus squash vein yellowing virus FL EU259611 NC_010521 Complete genome SqVYV★ Sweet potato mild mottle virus sweet potato mild mottle virus EA Z73124 NC_003797 Complete coding genome SPMMV★ Tomato mild mottle virus tomato mild mottle virus Ethiopia HE600072 NC_038920 Complete genome TMMoV

Tomato mild mottle virus eggplant mild leaf mottle virus Israel HQ840786 Complete genome EMLMV★ Ugandan cassava brown streak virus Ugandan cassava brown streak virus UG FJ185044 Complete genome UCBSV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Macluravirus

Distinguishing features

Macluraviruses resemble members of the genus Potyvirus in their transmission by aphids but virions are slightly shorter. They form a distinctgroup in phylogenetic analyses and have different polyprotein consensus cleavage sites.

Virion

Morphology

Virions are flexuous filaments mostly 650–675 nm×13–16 nm.

Physicochemical and physical properties

Virion Sedimentation coefficient S is 155–158S; density in CsCl is 1.31–1.33 g cm .

Nucleic acid

Virions contain one molecule of linear positive-sense, ssRNA of about 8.0 kb.

Proteins

Macluraviruses have a single coat (capsid) protein (CP) species of 33–34 kDa.

Genome organization and replication

The complete genome sequences of four different macluraviruses are available. The aa sequences of macluravirus CPs show limited (14–23%) identity with CP sequences of some aphid-transmitted potyviruses. Macluraviruses show significant aa sequence identity in portions ofthe replicase protein with viruses in other genera of the family Potyviridae. The macluraviruses seem to have a genome organization andreplication strategy typical of viruses in the family Potyviridae (Figure 2.Potyviridae).

Biology

Host range

Current information suggests that most viruses have a narrow host range, infecting species in up to nine host families.

Transmission

The viruses are transmitted by aphids in a non-persistent manner and experimentally by mechanical inoculation.

Antigenicity

Moderately immunogenic. No serological relationships to members of the genus Potyvirus have been found except for a weak reactionbetween maclura mosaic virus and bean yellow mosaic virus.

Derivation of names

Macluravirus: from the genus name of the host (Maclura) of members of the type species Maclura mosaic virus.

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Alpinia mosaic virus Alpinia mosaic virus TW AF499025 NC_043132 Partial genome AlpMV★ Alpinia oxyphylla mosaic virus Alpinia oxyphylla mosaic virus Alpo MG978107 Complete genome AloMV★ Artichoke latent virus artichoke latent virus FR37 KP405232 NC_026759 Complete genome ArLV

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Artichoke latent virus Ranunculus latent virus RN128 DQ152193 Partial genome RanLV★ Broad-leafed dock virus A broad-leafed dock virus A ab032 Auckland KU053507 NC_038560 Complete genome BDVA★ Cardamom mosaic virus cardamom mosaic virus KS MF622947 NC_039088 Complete genome CdMV★ Chinese yam necrotic mosaic

virusChinese yam necrotic mosaicvirus PES3 AB710145 NC_018455 Complete genome CYNMV

★ Maclura mosaic virus Maclura mosaic virus UK U58771 NC_043133 Partial genome MacMV★ Narcissus latent virus Narcissus latent virus NZ DQ450199 NC_043134 Partial genome NLV★ Yam chlorotic mosaic virus yam chlorotic mosaic virus YS KT724961 NC_038561 Complete genome YCMV★ Yam chlorotic necrosis virus yam chlorotic necrosis virus Y-Jish MG755240 Complete genome YCNV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name Accessionnumber

Virusabbreviation

Large cardamom chirkevirus JN257715* LCCV

Virus names and virus abbreviations are not official ICTV designations.

* incomplete genome sequence

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Genus: Poacevirus

Distinguishing features

Triticum mosaic virus (TriMV) is transmitted by the wheat curl mite and has an unusually long 5´-untranslated region. Vectors are unknown forthe other members of the genus.

Virion

Morphology

Virions of sugarcane streak mosaic virus (SCSMV) are flexuous filaments 890 nm×15 nm in size.

Nucleic acid

Virions contain a positive-sense ssRNA of 9.7–10.2 kb with a 3′-poly(A) terminus.

Proteins

The TriMV coat (capsid) protein (CP) is a single peptide of 293 aa with a MW of about 32 kDa. For SCSMV the viral CP is 281 aa with theMW estimated to be 36 kDa. Caladenia virus A (CalVA) has a CP of 285 aa and MW of 31

Genome organization and replication

The TriMV genome consists of 10,266 nt. The genome encodes a polyprotein of 3,122 aa. TriMV has an unusually long 5´-untranslatedregion, but this is not shared by members of the other two species. The order of coding regions is typical of that of the potyviruses (Figure2.Potyviridae). Most known potyvirus motifs are present in the polyprotein of poaceviruses. The aphid transmission amino acid sequencemotifs present in the HC-Pros (KITC, PTK) and CPs (DAG) of potyviruses are absent in poaceviruses. The zinc-finger-like motif H-(X )-C-(X )-C-(X )-C in the tritimovirus wheat streak mosaic virus (WSMV) HC-Pro is essential for its transmission by mites. A similar motif in thepoacevirus CalVA (unknown vector) is present as H-(X )-H-(X )-C-(X )-C (Wylie et al., 2012). In TriMV (mite transmitted) and SCSMV(unknown vector), the C-(X )-C motif exists, but the remainder of the motif is incomplete. P1 rather than the HC-Pro suppresses the hostRNA-suppression response in TriMV (Tatineni et al., 2012). Members of the genus Poacevirus are phylogenetically closest to those of anothermite-transmitted genus, Tritimovirus.

Biology

Host range

TriMV and SCSMV only infect hosts in the Gramineae, while CalVA infects orchids (Wylie et al., 2012).

Transmission

TriMV is transmitted by the wheat curl mite (Aceria tosichella).

Antigenicity

A Western blot revealed that antiserum raised to SCSMV weakly cross-reacts with wheat streak mosaic virus capsid protein. SCSMV doesnot cross react with other sugarcane-infecting members of the Potyviridae.

Derivation of names

Poacevirus: from Poaceae, the family name of the hosts of members of two species in the genus.

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Caladenia virus A Caladenia virus A KP1 JX156425 NC_018572 Complete genome CalVA

229 2

2 29 22

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★ Sugarcane streak mosaic virus sugarcane streak mosaic virus PAK GQ388116 NC_014037 Complete genome SCSMV★ Triticum mosaic virus Triticum mosaic virus U06-123 FJ263671 NC_012799 Complete genome TriMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Potyvirus

Distinguishing features

The largest genus in the family contains viruses transmitted by aphids in a non-persistent manner.

Virion

Morphology

Virions are flexuous filaments, 680–900 nm long and 11–13 nm wide, with helical symmetry and a pitch of about 3.4 nm. Particles of someviruses are longer in the presence of divalent cations than in the presence of EDTA.

Physicochemical and physical properties

Virion Sedimentation coefficient S is 137–160S; density in CsCl is 1.31 g cm ; Extinction coefficient E =2.4–2.7.

Nucleic acid

Virions contain a single molecule of linear, positive-sense ssRNA of about 9.7 kb; virions contain 5% RNA by weight.

Proteins

Virions contain a single coat (capsid) protein (CP) of 30–47 kDa. The CP of most isolates of the type species, Potato virus Y, contains 267 aa.

Genome organization and replication

The genome is organized as described in Figure 2.Potyviridae.

Biology

Many individual viruses have a narrow host range, but a few infect plant species in up to 30 host families. The viruses are transmitted byaphids in a non-persistent manner and are transmissible experimentally by mechanical inoculation. Some isolates are inefficiently transmittedby aphids and others are not transmissible by aphids at all. This is apparently due to mutations within the helper component and/or CPcistrons. Some viruses are seed-transmitted.

Antigenicity

Virions are moderately immunogenic; there are serological relationships among many members. Some monoclonal antibodies react withmost aphid-transmitted potyviruses. The CP aa sequence identity among aphid-transmitted viruses is 40–70%. Some viruses areserologically related to viruses in the genera Rymovirus and Bymovirus.

Derivation of names

Poty: from potato virus Y, an isolate of the type species Potato virus Y

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession

numberRefSeqnumber Available sequence Virus

Abbrev.★ African eggplant mosaic virus African eggplant mosaic virus 2013-281 MF997470 NC_043537 Complete genome AEMV★ Algerian watermelon mosaic

virus Algerian watermelon mosaic virus H4 EU410442 NC_010736 Complete genome AWMV

★ Alstroemeria mosaic virus Alstroemeria mosaic virus; Alstroemeria streak virus O1 AB158522 NC_043135 Partial genome AlMV★ Alternanthera mild mosaic virus Alternanthera mild mosaic virus BR EF442668 NC_043136 Partial genome AltMMV★ Amaranthus leaf mottle virus Amaranthus leaf mottle virus I AJ580095 NC043420 Partial genome AmLMV★ Amazon lily mosaic virus Amazon lily mosaic virus JP AB158523 NC_043137 Partial genome ALiMV★ Angelica virus Y Angelica virus Y g EF488741 NC_043138 Partial genome AVY★ Apium virus Y Apium virus Y; parsley virus Y Ce HM363516 NC_014905 Complete genome ApVY★ Araujia mosaic virus Araujia mosaic virus ARG1973 EF710625 NC_043139 Partial genome ArjMV★ Arracacha mottle virus Arracacha mottle virus C-17 DQ925486 NC_018176 Complete genome AMoV

20,w −3 0.1%1 cm, 260 nm

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★ Asparagus virus 1 Aasparagus virus 1 DSMZ PV-0954 KJ830760 NC_025821 Complete genome AV1

★ Banana bract mosaic virus banana bract mosaic virus PH DQ851496 NC_009745 Complete genome BBrMV★ Barbacena virus Y Barbacena virus Y KLL097 KU685505 NC_030847 Complete genome BarVY★ Basella rugose mosaic virus Basella rugose mosaic virus AC DQ821938 NC_009741 Complete genome BaRMV

Basella rugose mosaic virus peace lily mosaic virus Haiphong DQ851494 Complete genome PcLMV★ Bean common mosaic necrosis

virusbean common mosaic necrosis virus; bean common mosaic virusserotype A NL-3 U19287 NC_004047 Complete coding

genome BCMNV

★ Bean common mosaic virus blackeye cowpea mosaic virus R AJ312437 NC_003397 Complete genome BlCMVBean common mosaic virus peanut stripe virus blotch U34972 Complete genome PStVBean common mosaic virus bean common mosaic virus NL1 AY112735 Complete genome BCMV

★ Bean yellow mosaic virus bean yellow mosaic virus MB4 D83749 NC_003492 Complete genome BYMVBean yellow mosaic virus white lupin mosaic virus W DQ641248 Complete genome WhLMV

★ Beet mosaic virus beet mosaic virus Wa AY206394 NC_005304 Complete genome BtMV★ Bidens mosaic virus Bidens mosaic virus SP01 KF649336 NC_023014 Complete genome BiMV★ Bidens mottle virus Bidens mottle virus; sunflower chlorotic spot virus SF-1 AF538686 NC_014325 Complete genome BiMoV★ Blue squill virus A blue squill virus A SW3 JQ807999 NC_019415 Complete genome BSVA★ Brugmansia mosaic virus Brugmansia mosaic virus SK JX867236 NC_020105 Complete genome BruMV★ Brugmansia suaveolens mottle

virus Brugmansia suaveolens mottle virus Bs-Campinas AB551370 NC_014536 Complete genome BsMoV

★ Butterfly flower mosaic virus butterfly flower mosaic virus Hangzhou AM774001 NC_043421 Partial genome BFMV★ Calanthe mild mosaic virus Calanthe mild mosaic virus JP AB011404 NC_043140 Partial genome CalMMV★ Calla lily latent virus calla lily latent virus m19 No entry in Genbank CLLV★ Callistephus mottle virus Callistephus mottle virus DJ KX013584 NC_030794 Complete genome CalMoV★ Canna yellow streak virus Canna yellow streak virus UK GQ421689 NC_013261 Complete genome CaYSV★ Carnation vein mottle virus carnation vein mottle virus JP AB017630 NC_043141 Partial genome CVMoV★ Carrot thin leaf virus carrot thin leaf virus Cs JX156434 NC_025254 Complete genome CTLV★ Carrot virus Y carrot virus Y Vic AF203537 NC_043142 Partial genome CarVY★ Catharanthus mosaic virus Catharanthus mosaic virus Mandevilla-

US KP343681 NC_027210 Complete genome CatMV

★ Celery mosaic virus celery mosaic virus California HQ676607 NC_015393 Complete genome CeMV★ Ceratobium mosaic virus Ceratobium mosaic virus 13 AF022442 NC_043143 Partial genome CerMV★ Chilli ringspot virus chilli ringspot virus HN/14 JN008909 NC_016044 Complete coding

genome CRSV

★ Chilli veinal mottle virus chilli veinal mottle virus PVB AJ237843 NC_005778 Complete genome ChiVMV★ Chinese artichoke mosaic virus Chinese artichoke mosaic virus JP AB099711 NC_043144 Partial genome ChAMV★ Clitoria virus Y Clitoria virus Y QD AF228515 NC_043145 Partial genome ClVY★ Clover yellow vein virus clover yellow vein virus No.30 AB011819 NC_003536 Complete coding

genome ClYVV

★ Cocksfoot streak virus cocksfoot streak virus DE AF499738 NC_003742 Complete genome CSV★ Colombian datura virus Columbian datura virus; Petunia flower mottle virus UK JQ801448 NC_020072 Complete genome CDV★ Commelina mosaic virus Commelina mosaic virus FL No entry in Genbank ComMV★ Cowpea aphid-borne mosaic

virus cowpea aphid-borne mosaic virus; South African passiflora virus ZM AF348210 NC_004013 Complete codinggenome CABMV

★ Cucurbit vein banding virus cucurbit vein banding virus 43468 KY657266 NC_035134 Complete genome CVBV★ Cypripedium virus Y Cypripedium virus Y CP AF185954 NC_043146 Partial genome CypVY★ Cyrtanthus elatus virus A Cyrtanthus elatus virus A; Vallota speciosa virus Marijiniup 7 JQ723475 NC_017977 Complete genome CEVA★ Daphne mosaic virus Daphne mosaic virus CZ DQ299908 NC_008028 Complete genome DapMV★ Daphne virus Y Daphne virus Y SK KU556609 NC_038984 Complete genome DVY★ Dasheen mosaic virus dasheen mosaic virus M13 AJ298033 NC_003537 Complete genome DsMV

Dasheen mosaic virus vanilla mosaic virus CI KX505964 Complete genome VanMV★ Datura shoestring virus Datura shoestring virus IN No entry in Genbank DSSV★ Dendrobium chlorotic mosaic

virus Dendrobium chlorotic mosaic virus 98-De-31 MK241979 Complete genome DeCMV

★ Dioscorea mosaic virus Dioscorea mosaic virus FL MH206616 Complete genome DMV★ Diuris virus Y Diuris virus Y AU AF203527 NC_043147 Partial genome DiVY★ Donkey orchid virus A donkey orchid virus A SW3.1 JX156422 NC_021197 Complete coding

genome DOVA

★ East Asian Passiflora distortionvirus East Asian Passiflora distortion virus AK LC379162 Complete genome EAPDV

★ East Asian Passiflora virus East Asian passiflora virus AO AB246773 NC_007728 Complete genome EAPV★ Endive necrotic mosaic virus endive necrotic mosaic virus 31048 AJ223827 Partial genome ENMV★ Euphorbia ringspot virus Euphorbia ringspot virus US AY697300 Partial genome EuRSV★ Freesia mosaic virus Freesia mosaic virus KO FM206346 NC_014064 Complete genome FreMV★ Fritillary virus Y fritillary virus Y Pan'an AM039800 NC_010954 Complete genome FVY★ Gloriosa stripe mosaic virus Gloriosa stripe mosaic virus; christmas bell potyvirus CB EF427894 NC_038562 Complete coding

genome GSMV

★ Gomphocarpus mosaic virus Gomphocarpus mosaic virus CM532 LC228573 Complete genome GoMV★ Habenaria mosaic virus Habenaria mosaic virus Ha-1 AB818538 NC_021786 Complete genome HaMV★ Hardenbergia mosaic virus Hardenbergia mosaic virus 57.2 HQ161081 NC_015394 Complete genome HarMV★ Henbane mosaic virus henbane mosaic virus PHYS/H AM184113 NC_043422 Partial genome HMV★ Hibbertia virus Y Hibbertia virus Y Kioloa AF228516 NC_043148 Partial genome HiVY★ Hippeastrum mosaic virus Hippeastrum mosaic virus Marijiniup 1 JQ395040 NC_017967 Complete genome HiMV★ Hyacinth mosaic virus hyacinth mosaic virus Nannup

BC28 KY828925 NC_037051 Complete genome HyaMV

★ Impatiens flower break virus impatiens flower break virus Asan KU981084 NC_030236 Complete genome IFBV★ Iris fulva mosaic virus iris fulva mosaic virus US No entry in Genbank IFMV★ Iris mild mosaic virus iris mild mosaic virus WA-1 JF320812 NC_043149 Partial genome IMMV★ Iris severe mosaic virus iris severe mosaic virus BJ KT692938 NC_029076 Complete genome ISMV★ Japanese yam mosaic virus Japanese yam mosaic virus mild AB027007 NC_000947 Complete genome JYMV★ Jasmine virus T Jasmine virus T FZ KT222674 NC_029051 Complete genome JVT★ Johnsongrass mosaic virus Johnsongrass mosaic virus; sugarcane mosaic virus

Johnsongrass strain AU Z26920 NC_003606 Complete codinggenome JGMV

★ Kalanchoe mosaic virus Kalanchoë mosaic virus F39 GQ497731 NC_043150 Partial genome KMV★ Keunjorong mosaic virus keunjorong mosaic virus Cheongwon JF838187 NC_016159 Complete genome KjMV★ Konjac mosaic virus konjac mosaic virus; Japanese hornwort mosaic virus F AB219545 NC_007913 Complete genome KoMV★ Leek yellow stripe virus leek yellow stripe virus Yuhang GYH AJ307057 NC_004011 Complete coding

genome LYSV

★ Lettuce Italian necrotic virus lettuce Italian necrotic virus I234 KP769852 NC_027706 Complete genome LINV★ Lettuce mosaic virus lettuce mosaic virus E X97705 NC_003605 Complete coding

genome LMV

★ Lily mottle virus lily mottle virus; tulip band breaking virus; tulip breaking virus lilystrain Sb AJ564636 NC_005288 Complete coding

genome LMoV

★ Lily virus Y lily yellow mosaic virus lily-bua MF543013 NC_040802 Complete genome LVY★ Lupinus mosaic virus lupine mosaic virus LU2 EU847625 NC_014898 Complete genome LuMV★ Lycoris mild mottle virus Lycoris mild mottle virus TW AF399672 NC_043151 Partial genome LyMMoV★ Maize dwarf mosaic virus maize dwarf mosaic virus Bulgaria AJ001691 NC_003377 Complete genome MDMV

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★ Malva vein clearing virus Malva vein clearing virus DS-Ba-01 FM212972 NC_043423 Partial genome MVCV★ Mashua virus Y mashua virus Y Cam MH680824 Complete genome MasVY★ Meadow saffron breaking virus meadow saffron breaking virus FR AY388995 NC_043152 Partial genome MSBV★ Mediterranean ruda virus Mediterranean ruda virus ParP17 MF953305 NC_043536 Complete genome MeRV★ Moroccan watermelon mosaic

virus Moroccan watermelon mosaic virus TN05-76 EF579955 NC_009995 Complete genome MWMV

★ Narcissus degeneration virus Narcissus degeneration virus Zhangzhou AM182028 NC_008824 Complete genome NDV★ Narcissus late season yellows

virus Narcissus late season yellows virus Marijiniup8 KC691259 NC_023628 Complete genome NLSYV

★ Narcissus yellow stripe virus Narcissus yellow stripe virus Zhangzhou AM158908 NC_011541 Complete codinggenome NYSV

★ Nerine yellow stripe virus Nerine yellow stripe virus Ne800 EF362621 NC_043153 Partial genome NeYSV★ Nothoscordum mosaic virus Nothoscordum mosaic virus CAY No entry in Genbank NoMV★ Onion yellow dwarf virus onion yellow dwarf virus Yuhang AJ510223 NC_005029 Complete coding

genome OYDV

★ Ornithogalum mosaic virus Ornithogalum stripe mosaic virus KP JQ807997 NC_019409 Complete genome OrMV★ Ornithogalum virus 2 Ornithogalum virus 2 Japanese AB271783 NC_043154 Partial genome OrV2★ Ornithogalum virus 3 Ornithogalum virus 3 JP AB282754 NC_043155 Partial genome OrV3★ Panax virus Y Panax virus Y Yunnan GQ916624 NC_014252 Complete genome PanVY★ Papaya leaf distortion mosaic

virus papaya leaf distortion mosaic virus P BD171712 NC_005028 Complete genome PLDMV

★ Papaya ringspot virus papaya ringspot virus HA X67673 NC_001785 Complete genome PRSVPapaya ringspot virus papaya ringspot virus; watermelon mosaic virus 1 CI AY027810 Complete genome PRSV

★ Paris mosaic necrosis virus Paris mosaic necrosis virus PMNV-cn MF509898 NC_043532 Complete genome PMNV★ Parsnip mosaic virus parsnip mosaic virus UK No entry in Genbank ParMV★ Passiflora chlorosis virus Passiflora chlorosis virus Florida DQ860147 NC_043156 Partial genome PaCV★ Passion fruit woodiness virus passion fruit woodiness virus MU2 HQ122652 NC_014790 Complete genome PWV★ Pea seed-borne mosaic virus pea seed-borne mosaic virus DPD1 D10930 NC_001671 Complete coding

genome PSbMV

★ Peanut mottle virus peanut mottle virus M AF023848 NC_002600 Complete genome PeMoV★ Pecan mosaic-associated virus pecan mosaic-associated virus LA KT633868 NC_030293 Complete genome PMaV★ Pennisetum mosaic virus Pennisetum mosaic virus B AY642590 NC_007147 Complete genome PenMV★ Pepper mottle virus pepper mottle virus California M96425 NC_001517 Complete genome PepMoV★ Pepper severe mosaic virus pepper severe mosaic virus KO AM181350 NC_008393 Complete genome PepSMV★ Pepper veinal mottle virus pepper veinal mottle virus P DQ645484 NC_011918 Complete genome PVMV★ Pepper yellow mosaic virus pepper yellow mosaic virus Pi-15 AB541985 NC_014327 Complete genome PepYMV★ Peru tomato mosaic virus Peru tomato mosaic virus PPK13 AJ437280 NC_004573 Complete genome PTV★ Pfaffia mosaic virus Pfaffia mosaic virus BR AY485276 NC_043157 Partial genome PfMV★ Platycodon mild mottle virus Platycodon mild mottle virus Okcheon MH779625 Complete genome PlaMMV★ Pleione virus Y Pleione virus Y AU AF185958 NC_043158 Partial genome PlVY★ Plum pox virus plum pox virus M AJ243957 Complete genome PPV

Plum pox virus plum pox virus D X16415 Complete genome PPVPlum pox virus plum pox virus Rec JQ794501 Complete genome PPVPlum pox virus plum pox virus W3174 AY912055 Complete genome PPVPlum pox virus plum pox virus El Amar AM157175 Complete genome PPVPlum pox virus plum pox virus Volk143 KJ787006 Complete genome PPV

★ Pokeweed mosaic virus pokeweed mosaic virus MS-FRO3 JX291161 Complete codinggenome PkMV

★ Potato virus A potato virus A B11 AJ296311 NC_004039 Complete genome PVAPotato virus A Tamarillo mosaic virus FN AJ131403 Complete genome TamMV

★ Potato virus V potato virus V DV 42 AJ243766 NC_004010 Complete codinggenome PVV

★ Potato virus Y potato virus Y O U09509 Complete genome PVYPotato virus Y potato virus Y N X97895 Complete genome PVYPotato virus Y potato virus Y NTN AY166866 Complete genome PVYPotato virus Y potato virus Y C AJ890348 Complete genome PVYPotato virus Y potato virus Y Wilga 156 AJ889867 Complete genome PVY

★ Potato yellow blotch virus potato yellow blotch virus QV276 JX294310 Complete genome PYBV★ Ranunculus leaf distortion virus Ranunculus leaf distortion virus RN122 DQ152190 NC_043159 Partial genome RanLDV★ Ranunculus mild mosaic virus Ranunculus mild mosaic virus RN129 DQ152191 NC_043160 Partial genome RanMMV★ Ranunculus mosaic virus Ranunculus mosaic virus RN136 DQ152192 NC_043161 Partial genome RanMV★ Rhopalanthe virus Y Rhopalanthe virus Y AU AF185956 NC_043162 Partial genome RhVY★ Saffron latent virus saffron latent virus Ir-Kh1 KY562565 NC_036802 Complete genome SaLV★ Sarcochilus virus Y Sarcochilus virus Y AU AF185957 NC_043163 Partial genome SaVY★ Scallion mosaic virus scallion mosaic virus HZ AJ316084 NC_003399 Complete genome ScaMV★ Shallot yellow stripe virus shallot yellow stripe virus; Welsh onion yellow stripe virus ZQ2 AJ865076 NC_007433 Complete genome SYSV★ Sorghum mosaic virus sorghum mosaic virus Xiaoshan AJ310197 NC_004035 Complete coding

genome SrMV

★ Soybean mosaic virus soybean mosaic virus N D00507 NC_002634 Complete genome SMVSoybean mosaic virus Agropyron mosaic virus Pinella AJ507388 Complete genome SMV

★ Spiranthes mosaic virus 3 Spiranthes mosaic virus 3 USA AY685218 NC_043164 Partial genome SpMV3★ Sudan watermelon mosaic

virus Sudan watermelon mosaic virus Su94-54 KY623505 NC_035459 Complete genome SuWMV

★ Sugarcane mosaic virus sugarcane mosaic virus Zhejiang AJ297628 NC_003398 Complete genome SCMV★ Sunflower chlorotic mottle virus sunflower chlorotic mottle virus Common GU181199 NC_014038 Complete genome SCMoV★ Sunflower mild mosaic virus sunflower mild mosaic virus Entre Rios JQ350738 NC_021065 Complete genome SMMV★ Sunflower mosaic virus sunflower mosaic virus TX AF465545 NC_043165 Partial genome SuMV★ Sunflower ring blotch virus sunflower ring blotch virus Chaco KX856009 NC_034208 Complete genome SuRBV★ Sweet potato feathery mottle

virus sweet potato feathery mottle virus S D86371 NC_001841 Complete genome SPFMV

★ Sweet potato latent virus sweet potato latent virus TW KC443039 NC_020896 Complete genome SPLV★ Sweet potato mild speckling

virus sweet potato mild speckling virus AR U61228 NC_038563 Partial genome SPMSV

★ Sweet potato virus 2 sweet potato virus 2; Ipomoea vein mosaic virus; sweet potatovirus Y GWB-2 JN613807 NC_017970 Complete genome SPV2

★ Sweet potato virus C sweet potato virus C C1 GU207957 NC_014742 Complete genome SPVC★ Sweet potato virus G sweet potato virus G Jesus Maria JQ824374 NC_018093 Complete genome SPVG★ Tamarillo leaf malformation

virus Tamarillo leaf malformation virus A KM523548 NC_026615 Complete genome TLMV

★ Telfairia mosaic virus Telfairia mosaic virus No entry in Genbank TeMV★ Telosma mosaic virus Telosma mosaic virus Hanoi DQ851493 NC_009742 Complete genome TelMV★ Thunberg fritillary mosaic virus Thunberg fritillary mosaic virus Ningbo AJ851866 NC_007180 Complete genome TFMV★ Tobacco etch virus tobacco etch virus HAT M11458 NC_001555 Complete genome TEV★ Tobacco mosqueado virus tobacco mosqueado virus RS-01 KT834407 NC_030118 Complete genome TMosqV

★Tobacco vein banding mosaicvirus tobacco vein banding mosaic virus YND EF219408 NC_009994 Complete genome TVBMV

★ Tobacco vein mottling virus tobacco vein mottling virus KY X04083 NC_001768 Complete genome TVMV

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★ Tomato necrotic stunt virus tomato necrotic stunt virus MX9354 JQ314463 NC_017824 Complete genome TNSV★ Tradescantia mild mosaic virus Tradescantia mild mosaic virus IFA195 AY861351 NC_043166 Partial genome TraMMV★ Tuberose mild mosaic virus tuberose mild mosaic virus TW AF062926 NC_043167 Partial genome TuMMV★ Tuberose mild mottle virus tuberose mild mottle virus Hangzhou AJ581528 NC_043424 Partial genome TuMMoV★ Tulip breaking virus tulip breaking virus Texas Flame KF826466 NC_043168 Partial genome TBV★ Tulip mosaic virus tulip mosaic virus JP X63630 NC_043425 Partial genome TulMV★ Turnip mosaic virus turnip mosaic virus UK1 AF169561 NC_002509 Complete genome TuMV★ Twisted-stalk chlorotic streak

virus twisted-stalk chlorotic streak virus Denali 2001 AY954248 NC_043169 Partial genome TSCSV

★ Vallota mosaic virus Vallota mosaic virus NZ FJ618540 NC_043170 Partial genome ValMV★ Vanilla distortion mosaic virus vanilla distortion mosaic virus Cor KF906523 NC_025250 Complete genome VDMV★ Verbena virus Y Verbena virus Y Michigan EU564817 NC_010735 Complete genome VVY★Watermelon leaf mottle virus watermelon leaf mottle virus FL AF028004 NC_043171 Partial genome WLMV★Watermelon mosaic virus watermelon mosaic virus; watermelon mosaic virus 2 Fr AY437609 NC_006262 Complete genome WMV★Wild melon banding virus wild melon banding virus Su03-07 KY623506 NC_035458 Complete genome WMVBV★Wild onion symptomless virus wild onion symptomless virus TUR256-1 LC159494 NC_030391 Complete coding

genome WoSV

★Wild potato mosaic virus wild potato mosaic virus Type AJ437279 NC_004426 Complete genome WPMV★Wild tomato mosaic virus wild tomato mosaic virus Laichau DQ851495 NC_009744 Complete genome WTMV★Wisteria vein mosaic virus Wisteria vein mosaic virus Beijing AY656816 NC_007216 Complete genome WVMV★ Yam mild mosaic virus yam mild mosaic virus Brazil JX470965 NC_019412 Complete genome YMMV★ Yam mosaic virus yam mosaic virus Ivory Coast U42596 NC_004752 Complete genome YMV★ Yambean mosaic virus yambean mosaic virus SR JN190431 NC_016441 Complete genome YBMV★ Zantedeschia mild mosaic virus Zantedeschia mild mosaic virus TW AY626825 NC_011560 Complete genome ZaMMV★ Zea mosaic virus Iranian johnsongrass mosaic virus Shz JQ692088 NC_018833 Complete genome IJGMV

Zea mosaic virus Zea mosaic virus Israel AF228693 Partial genome ZeMV★ Zucchini shoestring virus zucchini shoestring virus RSA Patty

pan KU355553 NC_043172 Complete codinggenome ZSV

★ Zucchini tigre mosaic virus zucchini tigre mosaic virus Re01-25 KC345607 NC_023175 Complete genome ZTMV★ Zucchini yellow fleck virus zucchini yellow fleck virus It DQ641510 NC_043173 Partial genome ZYFV★ Zucchini yellow mosaic virus zucchini yellow mosaic virus TN3 AF127929 NC_003224 Complete genome ZYMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name Accessionnumber

Virusabbreviation

Ammi majus latent virus AB361564* AMLVAnemone mosaic virus EU042755* AnMVArisaema potyvirus 1 FJ546415* ArV1Arisaema potyvirus 2 FJ546416* ArV2Begonia flower breaking virus FJ539085* BFBVBermuda grass mosaic virus GU552269* BGMVBermuda grass southern mosaicvirus KU372146 BGSMV

Chickpea yellow mosaic virus AF527879* CpYMVClitoria chlorosis virus JF427593* ClCVCommelina mild mosaic virus HQ225836* CMMVCotyledon virus Y JN572103* CotVYDelphinium vein-clearing virus FJ349327* DeVCVLily virus A JN127335* LVAMelon vein-banding mosaic virus HQ423412* MVbMVMuscari mosaic virus EU042752* MMVOmphalodes virus Y AY974328* OmVYOrnamental onion stripe mosaicvirus EU042750* OOSMV

Ornithogalum necrotic mosaicvirus FJ159380* ONMV

Ornithogalum virus 4 EU042753* OV4Passiflora foetida virus Y AB679294* PfVYPecan mosaic-associated virus KT633868 PMaVpotato virus B JX294310Siratro 1 virus Y DQ098900* S1VYSiratro 2 virus Y DQ098901* S2VYSnowdrop virus Y EU927399* SVYStenomesson mosaic virus EU042757* StMV Tricyrtis virus Y AY864850* TrVY Trillium crinkled leaf virus FJ648825* TCLV Triteleia mosaic virus GU270649* TrMV

Veltheimia mosaic virus EF203686* VelMVVeltheimia virus Y EU684971* VelVY

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Virus names and virus abbreviations are not official ICTV designations.

* incomplete genome sequence

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Genus: Roymovirus

Distinguishing features

The amino acid motifs KITC and PTK are not present in HC-Pro. Roymoviruses have low nucleotide and amino acid identities with otherpotyvirids. The nucleotide and deduced amino acid sequences of the polyprotein share greatest identities with other viruses in the familyPotyviridae, but they are not clearly closer to members of any particular genera in the family. Comparison of the deduced amino acidsequence of the large ORF revealed identities with other viruses of the family to be low, ranging from 13% (some bymoviruses) to 23% (somepotyviruses).

Virion

Morphology

Particles are 720 to 750 nm in size.

Nucleic acid

Rose yellow mosaic virus has a genome of 9,508 nt.

Proteins

The large polyprotein is probably processed in the same manner as potyviruses. PIPO is present.

Genome organization and replication

Genome organization resembles that of members of the genus Potyvirus.

Biology

Isolates of the type species, Rose yellow mosaic virus, were isolated from several rose cultivars in New York and Minnesota.

The conserved amino acid motifs KITC and PTK are not present in the HC-Pro of rose yellow mosaic virus. Instead, a putative C-2x-Ceriophyid mite transmission motif is found at amino acid residues 48–51 at the N-terminus of the protein. The coat protein amino acid motifDAG found in most aphid-transmitted potyvirids is also lacking. Together this information suggests that rose yellow mosaic virus may betransmitted by eriophyid mites, although this has not been proven experimentally.

Phylogenetic analysis of the coat protein and ORF places rose yellow mosaic virus basal to viruses belonging to the genera Tritimovirus,Poeacevirus and Ipomovirus (Figure 3.Potyviridae).

Derivation of names

Roymovirus: from rose yellow mosaic virus

Species demarcation criteria

See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Passiflora edulis symptomless virus Passiflora edulis symptomless virus PeSV-Rehovot MH379332 Complete genome PeSV★ Rose yellow mosaic virus rose yellow mosaic virus Minnesota JF280796 NC_019031 Complete coding genome RYMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Rymovirus

Distinguishing features

This genus includes three species. Rymoviruses are presumably transmitted by host-adapted populations of eriophyid mite species in a semi-persistent manner. Rymoviruses share a reciprocal monophyletic relationship with members of the genus Potyvirus (French and Stenger2005).

Virion

Morphology

Virions are flexuous filaments 690–720 nm×11–15 nm in size.

Physicochemical and physical properties

Virion density in CsCl is 1.325 g cm (for ryegrass mosaic virus (RGMV)). Virion Sedimentation coefficient S is 165–166S for mostmembers.

Nucleic acid

Virions contain a single molecule of linear positive-sense ssRNA of about 9.5 kb with a 3′-poly(A) terminus.

Proteins

Rymoviruses encode one type of coat (capsid) protein (CP), with a theoretical Mr of 35,482 Da and an apparent Mr estimated by Westernblots of 45 kDa for RGMV.

Genome organization and replication

The complete genome sequences available for isolates of RGMV, Agropyron mosaic virus (AgMV), and Hordeum mosaic virus (HoMV)indicate that rymoviruses have a genome organization and replication strategy similar to other members of the Potyviridae with monopartitegenomes (Figure 2.Potyviridae).

Biology

Host range

Most rymoviruses have limited but widespread host ranges within the family Gramineae but some have relatively narrow host ranges.

Transmission

Transmission by eriophyid mites and mechanical transmission have been reported for most members. The eriophyid mites transmittingrymoviruses are different from those transmitting tritimoviruses. The cereal rust mite Abacarus hystrix transmits both RGMV and AgMV, butonly the former is efficiently transmitted. No vector is known for HoMV. Recent studies have revealed that host-associated populations of A.hystrix represent a species complex.

Antigenicity

Particles of most rymoviruses are moderately immunogenic. HoMV and AgMV are serologically related.

Derivation of names

Rymovirus: from ryegrass mosaic virus, a member of the type species Ryegrass mosaic virus.

Species demarcation criteria

See discussion under family description.

Member species

-3 20,w

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★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Agropyron mosaic virus Agropyron mosaic virus ND402 AY623626 NC_005903 Complete genome AgMV★ Hordeum mosaic virus Hordeum mosaic virus ATCC PV81 AY623627 NC_005904 Complete genome HoMV★ Ryegrass mosaic virus ryegrass mosaic virus Danish Y09854 NC_001814 Complete genome RGMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Genus: Tritimovirus

Distinguishing features

Tritimoviruses are transmitted by mites of a different species from those that transmit rymoviruses. They form a separate cluster inphylogenetic analyses.

Virion

Morphology

Virions are flexuous filaments 690–700 nm long.

Physicochemical and physical properties

Virion Sedimentation coefficient S is 166S for wheat streak mosaic virus (WSMV).

Nucleic acid

Virions contain a positive-sense ssRNA of about 9.4–9.6 kb with a 3′-poly(A) terminus.

Proteins

The viral coat (capsid) protein (CP) is a single peptide of about 349 aa for WSMV and 320 aa for brome streak mosaic virus (BrSMV). The Mrestimated by electrophoresis is 42 kDa.

Genome organization and replication

The WSMV genome consists of 9,384 nt excluding the 3′-terminal poly(A) tail. Sequence analysis reveals an ORF of 3,035 aa. The structureand organization of the WSMV genome is similar to those of other members of the family Potyviridae except the bymoviruses (Figure2.Potyviridae). Most known potyvirus motifs are present in the polyprotein of WSMV. However, motifs in the putative helper-component andCP of BrSMV are incomplete or missing, which may account for different vector relations of the tritimoviruses. The WSMV CP sequenceshows limited (22-25%) identity with CP sequences of some aphid-transmitted potyviruses. WSMV shows significant aa sequence identitywith aphid-transmitted potyviruses in the cylindrical inclusion protein and portions of the nuclear inclusion proteins. WSMV RNA is translatedin vitro into several large proteins that can be immunoprecipitated with WSMV CP antiserum, suggesting that WSMV uses a proteolyticprocessing strategy to express functional proteins such as the CP. Antiserum to tobacco etch virus (TEV) 58 kDa nuclear inclusion proteinalso reacts with in vitro translation products of WSMV. An in vitro translation product is precipitated with antiserum to HC-Pro helpercomponent of an isolate of the species Tobacco vein mottling virus (genus Potyvirus). Comparative sequence analyses show similarities withother members of the family Potyviridae, but these are limited to the nine mature proteins. WSMV is especially susceptible to proteinases inplanta and has CP molecules of 42, 36 and 32 kDa; the two smaller proteins are parts of the 42 kDa protein.

Biology

Host range

The viruses only affect hosts in the Gramineae but while WSMV has a wide host range BrSMV and ONMV have narrow ones.

Transmission

WSMV and BrSMV are transmitted by eriophyid mites in a semi-persistent manner. The HC-Pro of WSMV is required for mite transmission.All tritimoviruses are transmissible experimentally by mechanical inoculation.

Antigenicity

Moderately immunogenic. WSMV and oat necrotic mottle virus (ONMV) are serologically related to each other, but not to the other membersof the family Potyviridae.

Derivation of names

Tritimovirus: from the host genus (Triticum) and induced symptoms (mosaic) of isolates of the type species Wheat streak mosaic virus.

Species demarcation criteria

20,w

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See discussion under family description.

Member species

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available sequence Virus Abbrev.

★ Brome streak mosaic virus brome streak mosaic virus FR Z48506 NC_003501 Complete coding genome BrSMV★ Oat necrotic mottle virus oat necrotic mottle virus Type-NE AY377938 NC_005136 Complete genome ONMV★ Tall oatgrass mosaic virus tall oatgrass mosaic virus Benesov KF260962 NC_022745 Complete genome TOgMV★Wheat eqlid mosaic virus wheat Eqlid mosaic virus IR EF608612 NC_009805 Complete genome WEqMV★Wheat streak mosaic virus wheat streak mosaic virus Sidney 81 AF057533 NC_001886 Complete genome WSMV★ Yellow oat grass mosaic virus yellow oat-grass mosaic virus Sb KF984546 NC_024471 Complete genome YOgMV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Unassigned species

Summary

Strawberry mottle virus, black raspberry virus and chocolate lily virus A are related to satsuma dwarf virus (SDV) in phylogenetic trees usingthe conserved Pro-Pol region (Figure 4.Secoviridae). Dioscorea mosaic associated virus, recently isolated from yam, is most closely relatedto chocolate lily virus A (Hayashi et al., 2016). These viruses also have a bipartite genome. However, the nature of their capsid protein(s) andtheir genomic organization are not known. For this reason, they are unassigned species in the family Secoviridae. Strawberry latent ringspotvirus was formerly considered a sadwavirus because it has two capsid proteins (CP) and some distant relation with SDV in phylogenetictrees using the Pro-Pol sequence (Figure 4.Secoviridae). However, its genomic organization is more related to that of cheraviruses (with theexception of the number of CPs, Figure 3.Secoviridae) and it branches more closely with cheraviruses than with sadwaviruses in thephylogenetic trees using the Pro-Pol sequence (Figure 4.Secoviridae). For these reasons, it is not considered a sadwavirus anymore, and isnow an unassigned species in the family Secoviridae.

Unassigned species in family Secoviridae

★ Exemplar isolate of the speciesSpecies Virus name Isolate Accession number RefSeq number Available

sequenceVirus

Abbrev.★ Strawberry latent ringspot

virusstrawberry latent ringspotvirus

NCGR MEN454.001

RNA-1: AY860978; RNA-2:AY860979

RNA-1: NC_006964; RNA-2:NC_006965 Complete genome SLRSV

Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

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Authors: Potyviridae

Stephen J Wylie*Potyviridae Study Group ChairState Agricultural Biotechnology CentreMurdoch UniversityPerthWestern Australia 6150AustraliaE-mail: s.wylie@murdoch.edu.au

Alice Kazuko Inoue-NagataEmbrapa Hortaliças Empresa Brasileira de Pesquisa Agropecuária (Embrapa) BrasíliaBrazilE-mail: alice.nagata@embrapa.br Jan KreuzeInternational Potato Center (CIP)Apartado 1558Lima 12PeruE-mail: j.kreuze@cgiar.org>

Juan José López-MoyaCentre for Research in Agricultural Genomics CRAGCSIC-IRTA-UAB-UBCampus Bellaterra08193-BarcelonaSpainE-mail: juanjose.lopez@cragenomica.es

Kristiina MäkinenDepartment of MicrobiologyUniversity of HelsinkiFinlandE-mail: kristiina.mainen@helsinki.fi

Kazusato OhshimaLaboratory of Plant VirologyDepartment of Applied Biological SciencesFaculty of AgricultureSaga University1-banchiHonjo-machiSaga 840-8502JapanE-mail: ohshimak@cc.saga-u.ac.jp

Aiming WangLondon Research and Development CentreAgriculture and Agri-Food Canada1391 Sandford STLondonOntarioN5V 4T3CanadaE-mail: Aiming.Wang@AGR.GC.CA

* to whom correspondence should be addressed

Authors of a previous version of this Report

Mike AdamsMineheadSomersetTA24 5DYUnited KingdomE-mail: mike.adams.ictv@gmail.com

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Celia ChalamDivision of Plant QuarantineICAR-National Bureau of Plant Genetic ResourcesPusa CampusNew Delhi 110012India.E-mail: mailcelia@gmail.com

Shelly PraveenDivision of BiochemistryIndian Agricultural Research InstituteNew Delhi-110012IndiaE-mail: shellypraveen@hotmail.com

Frank RabensteinInstitute for Epidemiological and Pathogen DiagnosisJulius Kuhn InstitutQuedlinburgSaxony-AnhaltGermanyE-mail: frank.rabenstein@jki.bund.de

Drake StengerCrop Diseases, Pests, Genetics, ARS, USDA9611 S. Riverbend AveParlierCA 93648USAE-mail: dstenger@fresno.ars.usda.gov

Murilo Zerbini.Dep. de Fitopatologia/BIOAGROUniversidade Federal de ViçosaViçosaMG, 36570-900BrazilE-mail: zerbini@ufv.br

The chapter in the Ninth ICTV Report, which served as the template for this chapter, was contributed by Adams, M.J., Zerbini, F.M., French, R.,Rabenstein, F., Stenger, D.C. and Valkonen, J.P.T.

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Resources: Potyviridae

Sequence alignments and tree files:

Figure 3.Potyviridae

Tree file (nexus format)

Alignment file (fasta format)

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References: Potyviridae

Adams, M. J., Antoniw, J. F. & Beaudoin, F. (2005a). Overview and analysis of the polyprotein cleavage sites in the family Potyviridae. Mol PlantPathol 6, 471-487. [PubMed]

Adams, M. J., Antoniw, J. F. & Fauquet, C. M. (2005b). Molecular criteria for genus and species discrimination within the family Potyviridae. ArchVirol 150, 459-479. [PubMed]

Chung, B. Y., Miller, W. A., Atkins, J. F. & Firth, A. E. (2008). An overlapping essential gene in the Potyviridae. Proceedings of the NationalAcademy of Sciences, USA 105, 5897-5902. [PubMed]

French, R. & Stenger, D. C. (2005). Genome sequences of Agropyron mosaic virus and Hordeum mosaic virus support reciprocal monophyly ofthe genera Potyvirus and Rymovirus in the family Potyviridae. Arch Virol 150, 299-312. [PubMed]

Gibbs, A. & Ohshima, K. (2010). Potyviruses and the digital revolution. Annu Rev Phytopathol 48, 205-223. [PubMed]

Kekarainen, T., Savilahti, H. & Valkonen, J. P. (2002). Functional genomics on potato virus A: virus genome-wide map of sites essential for viruspropagation. Genome Res 12, 584-594. [PubMed]

Kumar, S., Stecher, G. & Tamura, K. (2016). MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Mol Biol Evol 33,1870-1874. [PubMed]

Larkin, M. A., Blackshields, G., Brown, N. P., Chenna, R., McGettigan, P. A., McWilliam, H., Valentin, F., Wallace, I. M., Wilm, A., Lopez, R.,Thompson, J. D., Gibson, T. J. & Higgins, D. G. (2007). Clustal W and Clustal X version 2.0. Bioinformatics 23, 2947-2948. [PubMed]

Li, W., Hilf, M. E., Webb, S. E., Baker, C. A. & Adkins, S. (2008). Presence of P1b and absence of HC-Pro in Squash vein yellowing virussuggests a general feature of the genus Ipomovirus in the family Potyviridae. Virus Res 135, 213-219. [PubMed]

Olspert, A., Chung, B. Y., Atkins, J. F., Carr, J. P. & Firth, A. E. (2015). Transcriptional slippage in the positive-sense RNA virus familyPotyviridae. EMBO Rep 16, 995-1004. [PubMed]

Rodamilans, B., Valli, A., Mingot, A., San Leon, D., Baulcombe, D., Lopez-Moya, J. J. & Garcia, J. A. (2015). RNA polymerase slippage as amechanism for the production of frameshift gene products in plant viruses of the Potyviridae family. J Virol 89, 6965-6967. [PubMed]

Shukla, D. D. & Ward, C. W. (1989). Structure of potyvirus coat proteins and its application in the taxonomy of the potyvirus group. Adv Virus Res36, 273-314. [PubMed]

Susaimuthu, J., Tzanetakis, I. E., Gergerich, R. C. & Martin, R. R. (2008). A member of a new genus in the Potyviridae infects Rubus. VirusRes 131, 145-151. [PubMed]

Tatineni, S., Qu, F., Li, R., Morris, T. J. & French, R. (2012). Triticum mosaic poacevirus enlists P1 rather than HC-Pro to suppress RNAsilencing-mediated host defense. Virology 433, 104-115. [PubMed]

Valli, A., Lopez-Moya, J. J. & Garcia, J. A. (2007). Recombination and gene duplication in the evolutionary diversification of P1 proteins in thefamily Potyviridae. J Gen Virol 88, 1016-1028. [PubMed]

Wei, T. & Wang, A. (2008). Biogenesis of cytoplasmic membranous vesicles for plant potyvirus replication occurs at endoplasmic reticulum exitsites in a COPI- and COPII-dependent manner. J Virol 82, 12252-12264. [PubMed]

Winter, S., Koerbler, M., Stein, B., Pietruszka, A., Paape, M. & Butgereitt, A. (2010). Analysis of cassava brown streak viruses reveals thepresence of distinct virus species causing cassava brown streak disease in East Africa. J Gen Virol 91, 1365-1372. [PubMed]

Wylie, S. J., Tan, A. J., Li, H., Dixon, K. W. & Jones, M. G. (2012). Caladenia virus A, an unusual new member of the family Potyviridae fromterrestrial orchids in Western Australia. Arch Virol 157, 2447-2452. [PubMed]

You, Y. & Shirako, Y. (2010). Bymovirus reverse genetics: requirements for RNA2-encoded proteins in systemic infection. Mol Plant Pathol 11,383-394. [PubMed]

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Citation: Potyviridae

A summary of this ICTV Report chapter has been published as an ICTV Virus Taxonomy Profile article in the Journal of General Virology, andshould be cited when referencing this online chapter as follows:

Wylie, S.J., Adams, M., Chalam, C., Kreuze, J., López-Moya, J.J., Ohshima, K., Praveen, S., Rabenstein, F., Stenger, D., Wang, A., Zerbini,F.M., and ICTV Report Consortium, ICTV Virus Taxonomy Profile: Potyviridae, Journal of General Virology, 98: 352–354.

Funding support

Support for the preparation of this ICTV Report chapter and associated Journal of General Virology taxonomy profile, was funded by a grant fromthe Wellcome Trust (WT108418AIA).

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