Pepino mosaic virus

Pictures

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Picture

Title

Caption

Copyright

Symptoms

Mosaic and chlorosis on old tomato leaves induced by PepMV.

Piero Roggero

Symptoms

Mosaic on apical tomato leaves induced by PepMV.

Piero Roggero

Symptoms

Symptoms on immature and mature tomato fruits induced by PepMV.

Piero Roggero

Identity

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Preferred Scientific Name

Pepino mosaic virus

Local Common Names

Netherlands: pepino mozaïek virus

EPPO code

PEPMV0 (Pepino mosaic potexvirus)

Taxonomic Tree

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Domain: Virus
Group: "Positive sense ssRNA viruses"
Group: "RNA viruses"
Order: Tymovirales
Family: Alphaflexiviridae
Genus: Potexvirus
Species: Pepino mosaic virus

Notes on Taxonomy and Nomenclature

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Pepino mosaic virus (PepMV) is a definitive species of the genus Potexvirus. This genus, with potato virus X (PVX) as the type species, includes a number of agronomically important viruses. Currently two strains of the virus are recognized; the pepino strain as described by Jones et al (1980) and the tomato strain (Van der Vlugt et al., 2000).

Description

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PepMV particles show typical potexvirus morphology; flexible filaments and measure 510 x 12.5 nm (Jones et al. 1980). Ultrathin sections of infected Nicotiana glutinosa leaves show inclusions containing arrays of filamentous virus-like particles.

Distribution

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Pepino mosaic virus was first described in 1980 (Jones et al., 1980) when isolated from pepino (Solanum muricatum) plants collected in 1974 from the coastal region of Peru. After this initial description no further findings were recorded. In 1999 the virus was again diagnosed from greenhouse-grown tomato plants in the Netherlands. In early 2000 the virus was officially reported from three additional European countries: the UK, France and Germany (EC directive 325). Unofficially the virus has been found in plant material originating from other European and non-European countries. Given its transmission characteristics it is likely that the virus is more widespread then currently officially recognized.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 13 May 2022

Continent/Country/Region	Distribution	First Reported	Reference	Notes
Africa
Egypt	Present		EPPO (2022) Amer and Mahmoud (2020) Amer et al. (2020)
Madagascar	Absent, Unconfirmed presence record(s)		EPPO (2022)
Morocco	Present		CABI and EPPO (2015) Souiri et al. (2017) Souiri et al. (2019) EPPO (2022)
South Africa	Present, Few occurrences		CABI and EPPO (2015) Carmichael et al. (2011) EPPO (2022)
Asia
China	Present, Few occurrences		CABI and EPPO (2015) Zhang YaoLiang et al. (2003) EPPO (2022)
-Jiangsu	Absent, Invalid presence record(s)		EPPO (2022)
-Shanghai	Present		Zhang YaoLiang et al. (2003) EPPO (2022)
Israel	Present, Localized		EPPO (2020) Klap et al. (2020) Klap et al. (2020a) EPPO (2022)
South Korea	Present		Genbank (2022)	Genbank accession no. LC656470. Direct submission in 2021.
Syria	Present		Fakhro et al. (2010) CABI and EPPO (2015) EPPO (2022)	Latakia.
Taiwan	Absent, Unconfirmed presence record(s)		EPPO (2022)
Turkey	Present, Few occurrences		CABI and EPPO (2015) Șevİk and Delİgöz (2016) Kılıçgözde Ürgen and Yardımcı (2017) EPPO (2022)
Europe
Austria	Present, Few occurrences		EPPO (2022) CABI and EPPO (2015)
Belgium	Present, Few occurrences		Hanssen et al. (2008) Hanssen (2010) CABI and EPPO (2015) EPPO (2022)
Bulgaria	Absent, Formerly present		CABI and EPPO (2015) EPPO (2022)
Croatia	Absent, Eradicated		Milanović et al. (2011) Novak et al. (2011) Novak et al. (2012) CABI and EPPO (2015) EPPO (2022)
Cyprus	Present, Widespread		CABI and EPPO (2015) Papayiannis et al. (2012) EPPO (2022)
Czechia	Absent, Eradicated		EPPO (2022) IPPC (2008) Kůdela et al. (2010) CABI and EPPO (2015)
Denmark	Present, Few occurrences		CABI and EPPO (2015) EPPO (2022)
Estonia	Absent, Confirmed absent by survey		EPPO (2022)
Finland	Absent, Confirmed absent by survey	2001	CABI and EPPO (2015) EPPO (2022)
France	Present, Few occurrences		CABI and EPPO (2015) Bosseur et al. (2004) EPPO (2022)
Germany	Present, Few occurrences	1999	Schmatz et al. (2003) CABI and EPPO (2015) EPPO (2022)
Greece	Present, Localized		Efthimiou et al. (2011) CABI and EPPO (2015) EPPO (2022)
Hungary	Present, Few occurrences		Horváth et al. (2007) Takács et al. (2012) CABI and EPPO (2015) EPPO (2022)
Ireland	Present, Few occurrences		CABI and EPPO (2015) EPPO (2022)
Italy	Present, Few occurrences	2001	Roggero et al. (2001) Parrella and Crescenzi (2005) Davino et al. (2010) CABI and EPPO (2015) EPPO (2022)
-Sardinia	Present, Few occurrences		CABI and EPPO (2015) Tiberini et al. (2011) EPPO (2022)
-Sicily	Present, Localized		Davino et al. (2009) Panno et al. (2012) CABI and EPPO (2015) Davino et al. (2017) EPPO (2022)
Lithuania	Present, Few occurrences		EPPO (2022) Šneideris et al. (2013) Žižytė et al. (2013) CABI and EPPO (2015)
Malta	Absent, Confirmed absent by survey		EPPO (2022)
Netherlands	Present, Localized		Vlugt et al. (2000) Vlugt et al. (2002) Stijger and Vlugt (2006) NPPO of the Netherlands (2013) CABI and EPPO (2015) EPPO (2022) CABI (Undated)
Norway	Absent, Eradicated		CABI and EPPO (2015) EPPO (2022)
Poland	Present, Few occurrences		Hasiów-Jaroszewska et al. (2009) CABI and EPPO (2015) Minicka et al. (2020) EPPO (2022)
Portugal	Absent, Confirmed absent by survey		EPPO (2022)
Serbia	Present		Stankovic et al. (2020) EPPO (2022)	Jablanica district.
Slovakia	Absent, Eradicated		CABI and EPPO (2015) EPPO (2022)
Slovenia	Absent, Confirmed absent by survey		EPPO (2022)
Spain	Present, Widespread		Jordá et al. (2001) Jordá et al. (2001a) Córdoba et al. (2004) Córdoba-Sellés et al. (2007) Alfaro-Fernández et al. (2008) CABI and EPPO (2015) Verbeek et al. (2015) Alcaide et al. (2020) EPPO (2022)
-Balearic Islands	Absent, Confirmed absent by survey		EPPO (2022)
-Canary Islands	Present		Jordá et al. (2001) Alfaro-Fernández et al. (2007) CABI and EPPO (2015) Verbeek et al. (2015) Gómez-Aix et al. (2019) EPPO (2022)
Sweden	Absent, Formerly present	2001	CABI and EPPO (2015) EPPO (2022)
Switzerland	Present, Localized		CABI and EPPO (2015) EPPO (2022)
Ukraine	Present		CABI and EPPO (2015) Verhoeven et al. (2003) EPPO (2022)
United Kingdom	Absent		Mumford et al. (2004) Spence et al. (2006) CABI and EPPO (2015) Skelton et al. (2019) EPPO (2022)
-Channel Islands	Absent, Confirmed absent by survey		EPPO (2022)
-England	Absent, No presence record(s)		Skelton et al. (2019) EPPO (2022)	The PepMV isolate in this sample was a mild strain deliberately used by the grower to give cross protection. It is not a finding for England or the UK.
North America
Canada	Present, Few occurrences		French et al. (2001) Ling and Bledsoe (2009) CABI and EPPO (2015) EPPO (2022)
-Alberta	Present		CABI and EPPO (2015) French et al. (2005) EPPO (2022)
-British Columbia	Present		Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
-Ontario	Present, Few occurrences		French et al. (2001) Stobbs et al. (2009) CABI and EPPO (2015) EPPO (2022)
-Quebec	Present		CABI and EPPO (2015) Gilbert et al. (2010) EPPO (2022)
Guatemala	Absent, Invalid presence record(s)		CABI and EPPO (2015)
Mexico	Present, Few occurrences		CABI and EPPO (2015) Ling and Zhang (2011) Ling et al. (2011) EPPO (2022)
United States	Present, Localized		French et al. (2001) CABI and EPPO (2015) EPPO (2022)
-Alabama	Present		Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
-Arizona	Present		Maroon-Lango et al. (2005) Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
-California	Present		Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
-Colorado	Present		French et al. (2001) Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
-Florida	Present		CABI and EPPO (2015) EPPO (2022)
-Maryland	Present		Maroon-Lango et al. (2005) CABI and EPPO (2015) EPPO (2022)
-Minnesota	Present		Lockhart (2007) CABI and EPPO (2015) EPPO (2022)
-Oklahoma	Present		Maroon-Lango et al. (2003) CABI and EPPO (2015) EPPO (2022)
-Texas	Present		French et al. (2001) Ling et al. (2008) CABI and EPPO (2015) EPPO (2022)
Oceania
New Zealand	Present		EPPO (2021) EPPO (2022)	EPPO Reporting Service No. 2021/138. Detected in four commercial tomato glasshouses in Auckland.
South America
Chile	Present, Localized		Pospieszny et al. (2007) CABI and EPPO (2015) EPPO (2022)
Ecuador	Present		CABI and EPPO (2015) Soler et al. (2005) EPPO (2022)
Peru	Present, Widespread		Jones et al. (1980) Soler et al. (2002) Soler et al. (2002a) Moreno-Pérez et al. (2014) CABI and EPPO (2015) EPPO (2022)

Risk of Introduction

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PepMV is easily mechanically transmissible. Since symptoms are not always readily recognized, there is a danger that the virus can spread rapidly and unnoticed. Host plant studies show that other Solanaceous crop plants like aubergine and potato can be infected.

Hosts/Species Affected

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PepMV has a relatively narrow host range and primarily infects Solanaceous plants. The primary host of the tomato strain of PepMV is tomato (Lycopersicon esculentum) but the virus can also cause symptoms in pepino (Solanum muricatum), aubergine (Solanum melongena) and potato (Solanum tuberosum). There are indications that the host ranges of both strains may differ slightly.

Host Plants and Other Plants Affected

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Plant name	Family	Context	References
Amaranthus (amaranth)	Amaranthaceae	Wild host	Jordá et al. (2001); Jordá et al. (2001)
Bassia scoparia	Chenopodiaceae	Unknown	Córdoba et al. (2004)
Calystegia sepium (great bindweed)	Convolvulaceae	Unknown	Córdoba et al. (2004)
Chenopodiastrum murale (nettleleaf goosefoot)	Chenopodiaceae	Unknown	Córdoba et al. (2004)
Convolvulus althaeoides		Unknown	Córdoba et al. (2004)
Convolvulus arvensis (bindweed)	Convolvulaceae	Unknown	Córdoba et al. (2004)
Conyza sumatrensis (tall fleabane)	Asteraceae	Unknown	Córdoba et al. (2004)
Coronopus		Unknown	Córdoba et al. (2004)
Datura innoxia (downy thorn apple)	Solanaceae	Unknown	Hasiów-Jaroszewska et al. (2009)
Datura stramonium (jimsonweed)	Solanaceae	Other
Diplotaxis erucoides	Brassicaceae	Unknown	Córdoba et al. (2004)
Echium creticum		Unknown	Córdoba et al. (2004)
Echium humile		Unknown	Córdoba et al. (2004)
Heliotropium europaeum (common heliotrope)	Boraginaceae	Unknown	Córdoba et al. (2004)
Lycopersicon peruvianum	Solanaceae	Wild host	Soler et al. (2002); Soler et al. (2002)
Lycopersicon pimpinellifolium (currant tomato)	Solanaceae	Other	Soler et al. (2005); Soler et al. (2002); Soler et al. (2002)
Malva parviflora		Wild host
Malva parviflora (pink cheeseweed)	Malvaceae	Unknown	Jordá et al. (2001); Jordá et al. (2001)
Moricandia arvensis	Brassicaceae	Unknown	Córdoba et al. (2004)
Nicotiana glauca (tree tobacco)	Solanaceae	Wild host	Jordá et al. (2001); Jordá et al. (2001)
Nicotiana rustica (wild tobacco)	Solanaceae	Other
Ocimum basilicum (basil)	Lamiaceae	Other	Davino et al. (2009)
Onopordum		Unknown	Córdoba et al. (2004)
Piptatherum miliaceum (smilograss)	Poaceae	Unknown	Córdoba et al. (2004)
Plantago arenaria	Plantaginaceae	Unknown	Córdoba et al. (2004)
Rumex (Dock)	Polygonaceae	Unknown	Córdoba et al. (2004)
Sisymbrium irio	Brassicaceae	Unknown	Córdoba et al. (2004)
Solanum americanum	Solanaceae	Unknown	Jordá et al. (2001); Jordá et al. (2001); Stobbs et al. (2009)
Solanum chilense		Wild host	Soler et al. (2002); Soler et al. (2002)
Solanum chmielewskii		Wild host	Soler et al. (2002); Soler et al. (2002)
Solanum lycopersicum (tomato)	Solanaceae	Main	Alfaro-Fernández et al. (2007); Córdoba-Sellés et al. (2007); Fakhro et al. (2005); French et al. (2001); Jordá et al. (2001); Jordá et al. (2001); Mansilla et al. (2003); Martínez-Culebras et al. (2002); Soler et al. (2002); Vlugt et al. (2002); Vlugt et al. (2000); Hasiów-Jaroszewska et al. (2009); Ling (2008); Ling et al. (2008); Verbeek et al. (2015); Iacono et al. (2015); Souiri et al. (2017); Sneideris et al. (2013); Novak et al. (2012); Skelton et al. (2019); Amer and Mahmoud (2020); Amer et al. (2020)
Solanum melongena (aubergine)	Solanaceae	Main
Solanum muricatum (melon pear)	Solanaceae	Other	Jones et al. (1980); Martínez-Culebras et al. (2002); Soler et al. (2002)
Solanum neorickii		Wild host	Soler et al. (2002); Soler et al. (2002)
Solanum nigrum (black nightshade)	Solanaceae	Wild host
Solanum tuberosum (potato)	Solanaceae	Main
Sonchus oleraceus (common sowthistle)	Asteraceae	Wild host	Jordá et al. (2001); Jordá et al. (2001)
Sonchus tenerrimus (field sowthistle)	Asteraceae	Unknown	Córdoba et al. (2004)
Taraxacum officinale complex (dandelion)	Asteraceae	Unknown	Córdoba et al. (2004)

Growth Stages

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Flowering stage, Fruiting stage, Vegetative growing stage

Symptoms

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Initial symptoms of PepMV on tomato plants include small yellow leaf spots that may resemble symptoms caused by Potato virus X (PVX). At a later stage, older leaves may show mottling and top leaves may show slight curling. Symptoms may depend on climatic conditions and become more visible under (relative) low light conditions. Sometimes fruits of infected plants may show an orange mottling but these symptoms can be very erratic and may differ between the trusses of one plant.

Symptoms on other Solanaceous crop plants can differ. Some potato cultivars showed no visible symptoms upon mechanical inoculation while others reacted strongly with necrosis (Jones et al., 1980).

List of Symptoms/Signs

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Sign	Life Stages	Type
Fruit / discoloration
Leaves / abnormal forms
Leaves / abnormal patterns

Means of Movement and Dispersal

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Natural dispersal (non-biotic)

Leaves and fruits of infected plants can contain high concentrations of virus. Spread of the virus is by mechanical contact. Roots of infected tomato plants also show significant concentrations of virus. Upon damage or death of these roots, virus particles are likely to be released in the soil or drainage water.

Vector transmission

In glasshouse experiments, the virus was shown to be transmitted by contact through bumblebees (Bombus terrestris). It is not transmitted by aphids.

Seedborne spread

The virus can be present on the outside of seeds collected from infected fruits. Seed lots should therefore be disinfected. There are no indications that the virus is transmitted inside the seed.

Agricultural practices

The virus is easily mechanically transmissible and symptoms can be weak under certain conditions. When not recognized in time the virus is likely to spread rapidly through tomato crops by normal cultural practices through hands, tools, clothing and machinery. The use of skimmed milk to disinfect hands and tools is recommended. Experimental evidence (unpublished) suggests that PepMV is less stable then PVX. Standard disinfection and hygiene protocols that apply for PVX are also expected to be sufficient for PepMV.

Movement in trade

Fruits may contain high virus concentrations. Crates and packaging materials are likely to get contaminated following fruit damage. Reusable crate material should be thoroughly cleaned and disinfected.

Planting material originating from nurseries should be tested before transfer to production facilities.

Seedborne Aspects

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Incidence

PepMV was detected by serology in tomato seeds (Salomone and Roggero, 2002).

Pathogen Transmission

Seed

In 2002, PepMV was detected by serology in tomato seeds but was not found to be seed transmissible to 50 seedlings (Salomone and Roggero, 2002). More recently, Hanssen et al. (2010) demonstrated that PepMV can be transmitted by tomato seeds, highlighting the risk of using seeds from PepMV-infected plants and further spreading the disease. PepMV has been identified in the seed coat of both immature and mature tomato seeds, but not in the embryo (Ling, 2008). Córdoba-Sellés et al. (2007) estimate that at least 25% of seeds from infected tomato plants carry PepMV. PepMV is neither seed-transmitted or seedborne in Nicotiana benthamiana.

Other sources

PepMV is a very contagious pathogen, easily spread mechanically via contaminated tools, shoes, clothing, hands and plant-to-plant contact (Pospieszny and Borodynko, 2002).

Seed Health Tests

Seeds lots can be tested for the presence of the virus either by ELISA (Salomone and Roggero, 2002) or by PCR using Potex group-specific or PepMV primers sets.

Seed Treatments

Immersion of seeds in 10% trisodium phosphate for 3 h mostly eradicated the virus and was not detrimental to germination (Córdoba-Sellés et al. 2007). Ling (2010) found that a seed treatment of commercial bleach solution (0.5 and 1.0% sodium hypochlorite) was the most effective at reducing the incidence of PepMV, followed by trisodium phosphate. Hydorchloric acid treatment was not found to be effective. In the same study, dry heat baking seeds at 72 or 80°C for 48 to 72 h was effective in lowering PepMV infection rates, whereas soaking seeds in water at 55°C for two hours had a detrimental effect on seed germination.

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Containers and packaging - wood		Yes
Soil, sand and gravel	Drainage water.	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Borne internally	Borne externally	Visibility of pest or symptoms
Fruits (inc. pods)	Yes		Pest or symptoms usually invisible
Leaves	Yes		Pest or symptoms usually invisible
Roots	Yes		Pest or symptoms usually invisible
Stems (above ground)/Shoots/Trunks/Branches	Yes		Pest or symptoms usually invisible
True seeds (inc. grain)		Yes	Pest or symptoms usually invisible

Impact

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Infection of tomato crops with PepMV does not necessarily result in significant economic damage since fruit symptoms may be absent. However, experimental data (unpublished) suggest that yield may be affected and fruit setting may be delayed.

Diagnosis

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The presence of PepMV in infected plant material and fruits can be assayed by serological methods like ELISA, using commercially available antisera. Antisera are specific and react equally well with both strains. No cross-reaction to PVX or Potato aucuba mosaic virus (PAMV), the only other potexviruses known to infect Solanaceous crops, was observed.

PepMV is mechanically transmissible to different experimental host plants. Several test plants allow discrimination between the pepino and tomato strains of PepMV for example, Nicotiana glutinosa and Datura stramonium.

The use of RT PCR in diagnosing PepMV is described in Hasiów et al. (2008). A polyprobe method of detection is detailed in Aparicio et al. (2009). An immunocapture-retrotranscription-PCR (IC-RT-PCR) approach is detailed in Mansilla et al. (2003).

Detection and Inspection

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Detection of the virus in commercial tomato and other Solanaceous crops depends on field inspection for typical symptoms. There may be small yellow spots, especially on young leaves at the top of the plant. At a later stage a more clear mosaic may be seen on older leaves. Fruits may sometimes show an orange mottling. Depending on climatic conditions, virus symptoms on the leaves can sometimes be weak and hard to see, and the virus may therefore escape attention.

Similarities to Other Species/Conditions

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Initial, weak symptoms on tomato may resemble symptoms of PVX infection. A weak serological relationship to Narcissus mosaic virus is reported (Jones et al. 1980).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

The virus should be controlled by the use of virus-free seed and planting material and strict hygienic measures.

References

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Alfaro-Fernández, A., Córdoba-Sellés, C., Cebrián, M. C., Sánchez-Navarro, J. A., Espino, A., Martín, R., Jordá, C., 2007. First report of Tomato torrado virus in tomato in the Canary Islands, Spain. Plant Disease, 91(8), 1060. doi: 10.1094/PDIS-91-8-1060B

Amer, M. A., Ibrahim, Y. E., Kheder, A. A., Hamed, A. H., Farrag, A. A., Al-Saleh, M. A., 2020. Confirmation incidence of Tomato chlorosis virus naturally infecting tomato crop in Egypt. International Journal of Agriculture and Biology, 23(5), 963-969. doi: 10.17957/IJAB/15.1375

Amer, M. A., Mahmoud, S. Y., 2020. First report of Tomato brown rugose fruit virus on tomato in Egypt. New Disease Reports, 41 doi: 10.5197/j.2044-0588.2020.041.024

Aparicio F; Soler S; Aramburu J; Galipienso L; Nuez F; Pallás V; López C, 2009. Simultaneous detection of six RNA plant viruses affecting tomato crops using a single digoxigenin-labelled polyprobe. European Journal of Plant Pathology, 123(1):117-123. http://springerlink.metapress.com/link.asp?id=100265

CABI/EPPO, 2015. Pepino mosaic virus. [Distribution map]. Distribution Maps of Plant Diseases, No.October. Wallingford, UK: CABI, Map 856 (Edition 3).

Carmichael DJ; Rey MEC; Naidoo S; Cook G; Heerden SWvan, 2011. First report of Pepino mosaic virus infecting tomato in South Africa. Plant Disease, 95(6):767. http://apsjournals.apsnet.org/loi/pdis

Commission decision 2000/325/EC, 2000. Official journal of the European Communities. Brussels.

Córdoba, M. C., Martínez-Priego, L., Jordá, C., 2004. New natural hosts of Pepino mosaic virus in Spain. Plant Disease, 88(8), 906. doi: 10.1094/PDIS.2004.88.8.906D

Córdoba-Sellés Mdel C; García-Rández A; Alfaro-Fernández A; Jordá-Gutiérrez C, 2007. Seed transmission of Pepino mosaic virus and efficacy of tomato seed disinfection treatments. Plant Disease, 91(10):1250-1254. HTTP://www.apsnet.org

Davino, S., Accotto, G. P., Masenga, V., Torta, L., Davino, M., 2009. Basil (Ocimum basilicum), a new host of Pepino mosaic virus. Plant Pathology, 58(2), 407. doi: 10.1111/j.1365-3059.2009.02026.x

Efthimiou KE; Gatsios AP; Aretakis KC; Papayiannis LC; Katis NI, 2011. First report of Pepino mosaic virus infecting greenhouse cherry tomatoes in Greece. Plant Disease, 95(1):78. http://apsjournals.apsnet.org/loi/pdis

EPPO, 2011. EPPO Reporting Service. EPPO Reporting Service. Paris, France: EPPO. http://archives.eppo.org/EPPOReporting/Reporting_Archives.htm

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Fakhro, A., Paschek, U., Bargen, S. von, Büttner, C., Schwarz, D., 2005. Distribution and spread of Pepino mosaic virus (PepMV) in tomatoes cultivated in a re-circulating hydroponic system. In: Plant protection and plant health in Europe: introduction and spread of invasive species, held at Humboldt University, Berlin, Germany, 9-11 June 2005, [ed. by Alford, D. V., Backhaus, G. F.]. Alton, UK: British Crop Protection Council. 223-224.

French, C. J., Bouthillier, M., Bernardy, M., Ferguson, G., Sabourin, M., Johnson, R. C., Masters, C., Godkin, S., Mumford, R., 2001. First report of Pepino mosaic virus in Canada and the United States. Plant Disease, 85(10), 1121. doi: 10.1094/PDIS.2001.85.10.1121B

Hanssen IM; Mumford R; Blystad DR; Cortez I; Hasiów-Jaroszewska B; Hristova D; Pagán I; Pereira AM; Peters J; Pospieszny H; Ravnikar M; Stijger I; Tomassoli L; Varveri C; Vlugt Rvan der; Nielsen SL, 2010. Seed transmission of Pepino mosaic virus in tomato. European Journal of Plant Pathology, 126(2):145-152. http://springerlink.metapress.com/link.asp?id=100265

Hasiów B; Borodynko N; Pospieszny H, 2008. Development of a real-time RT-PCR assay for detection of Pepino mosaic virus different strains. (Zastosowanie techniki real time RT-PCR w diagnostyce róznych szczepów wirusa mozaiki pepino.) Progress in Plant Protection, 48(2):458-462. http://www.progress.plantprotection.pl/pliki/2008/48-2-22.pdf

Hasiów-Jaroszewska, B., Pospieszny, H., Borodynko, N., 2009. New necrotic isolates of Pepino mosaic virus representing the Ch2 genotype. Journal of Phytopathology, 157(7/8), 494-496. doi: 10.1111/j.1439-0434.2008.01496.x

Iacono, G., Hernandez-Llopis, D., Alfaro-Fernandez, A., Davino, M., Font, M. I., Panno, S., Galipenso, L., Rubio, L., Davino, S., 2015. First report of Southern tomato virus in tomato crops in Italy. New Disease Reports, 32, 27. http://www.ndrs.org.uk/article.php?id=032027

IPPC, 2008. First outbreak of Pepino mosaic virus in the Czech Republic. IPPC Official Pest Report, No. CZ-4/1. Rome, Italy: FAO. https://www.ippc.int/IPP/En/default.jsp

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