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Datasheet

Tomato spotted wilt orthotospovirus
(tomato spotted wilt)

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Datasheet

Tomato spotted wilt orthotospovirus (tomato spotted wilt)

Summary

  • Last modified
  • 18 December 2021
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Tomato spotted wilt orthotospovirus
  • Preferred Common Name
  • tomato spotted wilt
  • Taxonomic Tree
  • Domain: Virus
  •   Group: "Negative sense ssRNA viruses"
  •     Order: Bunyavirales
  •       Family: Tospoviridae
  •         Genus: Orthotospovirus
  • Summary of Invasiveness
  • Spotted wilt disease of tomatoes was first described in 1915 in Australia. The host range of Tomato spotted wilt orthotospovirus (TSWV) includes horticultural and agronomic crops across temperate, subtropical and tropical regions of the w...

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Pictures

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PictureTitleCaptionCopyright
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected Capsicum plant.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected Capsicum plant.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected Capsicum plant.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected Capsicum plant.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected tomato plant.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected tomato plant.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected tomato plant.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected tomato plant.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected aubergine.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected aubergine.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected aubergine.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on a leaf of an infected aubergine.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected tomato plant.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected tomato plant.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected tomato plant.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected tomato plant.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected Capsicum plant.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected Capsicum plant.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected Capsicum plant.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected Capsicum plant.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus damage symptoms to fruit.
TitleDamage symptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus damage symptoms to fruit.
Copyright©Nicola Spence/Horticulture Research International
Tomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus damage symptoms to fruit.
Damage symptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus damage symptoms to fruit.©Nicola Spence/Horticulture Research International
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected aubergine.
TitleSymptoms
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected aubergine.
Copyright©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected aubergine.
SymptomsTomato spotted wilt orthotospovirus (tomato spotted wilt); Tomato spotted wilt orthotospovirus; symptoms on the fruit of an infected aubergine.©Dirk Janssen
Tomato spotted wilt orthotospovirus (tomato spotted wilt); electron micrograph of tomato spotted wilt orthotospovirus particles, in negatively stained preparation, from foliar extract of infected tobacco.
TitleVirus particles
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); electron micrograph of tomato spotted wilt orthotospovirus particles, in negatively stained preparation, from foliar extract of infected tobacco.
Copyright©César M. Chagas
Tomato spotted wilt orthotospovirus (tomato spotted wilt); electron micrograph of tomato spotted wilt orthotospovirus particles, in negatively stained preparation, from foliar extract of infected tobacco.
Virus particlesTomato spotted wilt orthotospovirus (tomato spotted wilt); electron micrograph of tomato spotted wilt orthotospovirus particles, in negatively stained preparation, from foliar extract of infected tobacco.©César M. Chagas
Tomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus particles.
TitleVirus particles
CaptionTomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus particles.
Copyright©Nicola Spence/Horticulture Research International
Tomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus particles.
Virus particlesTomato spotted wilt orthotospovirus (tomato spotted wilt); orthotospovirus particles.©Nicola Spence/Horticulture Research International

Identity

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Preferred Scientific Name

  • Tomato spotted wilt orthotospovirus

Preferred Common Name

  • tomato spotted wilt

Other Scientific Names

  • dahlia oakleaf virus
  • dahlia ringspot virus
  • dahlia yellow ringspot virus
  • mung bean leaf curl virus
  • pineapple yellow spot virus
  • tomato spotted wilt tospovirus
  • Tomato spotted wilt virus

International Common Names

  • English: bronze leaf wilt; tomato bronze leaf virus
  • Spanish: bronceado del tomate
  • French: maladie des taches de bronze de la tomate
  • Portuguese: bronzeamento do tomateiro

Local Common Names

  • Argentina: corovo del tabaco; peste negra del tomate
  • Brazil: vira-cabeca
  • Chile: marchitamiento manchado del tomate
  • Germany: Bronzefleckenkrankheit
  • Italy: avvizzimento maculato del pomodoro
  • Netherlands: Tomatebronsvlekkenvirus
  • South Africa: kat river wilt; kromnek virus

English acronym

  • TSWV

EPPO code

  • TSWV00 (Tomato spotted wilt tospovirus)

Summary of Invasiveness

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Spotted wilt disease of tomatoes was first described in 1915 in Australia. The host range of Tomato spotted wilt orthotospovirus (TSWV) includes horticultural and agronomic crops across temperate, subtropical and tropical regions of the world. Major crops susceptible to TSWV infection are tomato, pepper, lettuce, potato, papaya, groundnut, tobacco and chrysanthemum. Symptoms vary with the host plant, time of year and environmental conditions and include stunting, necrosis, chlorosis, ring spots and ring/line patterns affecting leaves, stems and fruit. Infection rates of 50-90% lead to major losses in commercial vegetable crops and TSWV is now one of the 10 most economically destructive plant viruses with worldwide losses exceeding 1 billion dollars annually. It is a quarantine pest for Mexico, Norway, Morocco and Tunisia. It is categorized in the A2 list of EPPO (European and Mediterranean Plant Protection Organization).

Taxonomic Tree

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  • Domain: Virus
  •     Group: "Negative sense ssRNA viruses"
  •         Order: Bunyavirales
  •             Family: Tospoviridae
  •                 Genus: Orthotospovirus
  •                     Species: Tomato spotted wilt orthotospovirus

Notes on Taxonomy and Nomenclature

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The tomato disease was discovered in 1915 in Australia (Brittlebank, 1919) and its causal agent, a virus, was described by Samuel et al. (1930). This virus with pseudo-isometric or pleomorphic particles was the nominate member of the genus Tospovirus, one of five genera of the Bunyaviridae family (Murphy et al., 1995) and the only genus containing viruses that infect plants. In 2017 the order Bunyavirales was established to accommodate related viruses with segmented, linear, single-stranded, negative-sense or ambisense RNA genomes classified into nine families (Maes et al., 2018). The family Tospoviridae was later recreated for the established genus Tospovirus which was renamed Orthotospovirus (Abudurexiti et al., 2019). The genus Orthotospovirus currently contains 26 species (https://talk.ictvonline.org/taxonomy/).

This datasheet refers to all orthotospoviruses. As the exact host range, geographic distribution and damage caused remain to be established for most of these viruses; the emphasis will be on Tomato spotted wilt orthotospovirus (TSWV). This virus is the most widely encountered and thoroughly studied orthotospovirus. As some orthotospoviruses are potential threats to crops in Europe and the Mediterranean region, a quarantine pest status of TSWV is also attributable to these viruses.

Tomato spotted wilt orthotospovirus (TSWV), Impatiens necrotic spot orthotospovirus (INSV), Iris yellow spot orthotospovirus (IYSV) and Chrysanthemum stem necrosis orthotospovirus (CSNV) have been found in Europe and the Mediterranean region. TSWV occurs most commonly, and INSV occupies a prominent second place and is mainly found in horticultural crops. IYSV has been found a few times in the Netherlands on iris and leek, while CNSV has been detected a few times in plant material imported from Brazil. This group of viruses and their properties have been reviewed by Adam and Kegler (1994) and Mumford et al. (1996).

Description

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TSWV virions are pseudo-spherical or pleomorphic enveloped particles, ca 70-110 nm in diameter, which contain at least four protein species: an internal nucleocapsid protein (N); two membrane glycoproteins (G1 and G2) externally associated with the lipid envelope; and a large (L) protein, the putative polymerase. The genome consists of three linear ssRNA molecules which are complexed with N proteins to form circular nucleocapsids. The L (large) RNA molecule has a negative sense character; the M (middle) and S (short) RNA display an ambisense gene arrangement. The L protein is encoded by the L RNA, the two glycoproteins and the N protein are encoded by the viral complementary sense sequence of the M and S RNA, respectively. Two nonstructural proteins (NSs and NSm) are encoded by the viral sense sequence of the S and M RNA. The NSm protein represents the viral movement protein, enabling cell-to-cell spread of the virus and, hence, is essential for systemic infection of the plant. The NSs protein plays a crucial role in TSWV transmission by thrips (Margaria et al., 2014). This protein induces aggregates or filaments in TSWV infected cells and paracrystalline structures in INSV infected cells.

Reassortment of the RNA segments has been demonstrated by co-inoculating TSWV isolates (Qiu et al., 1998). Generation of reassortants in the field can contribute to the diversity of tospoviruses, especially with respect to symptom expression on different hosts, serological properties, transmission efficiency and spread of TSWV to other hosts and regions.

Distribution

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The data given are for TSWV 'sensu lato', since it is difficult to know to which virus earlier (i.e. prior to 1990) records refer. INSV was described in the USA and a few years later in Europe, where it has been recorded in France, Italy, the Netherlands, Portugal, UK, but probably occurs in all regions where ornamentals are cultivated. IYSV was isolated from some species of the Liliaceceae in the Netherlands, and later in Brazil, Israel and the USA.

See also CABI/EPPO (1998, No. 342).

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: 21 Jul 2022
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresent
Burkina FasoPresent
Congo, Democratic Republic of thePresent
Côte d'IvoirePresent
EgyptPresent
KenyaPresent, Localized
LibyaPresent
MadagascarPresent, Localized
MauritiusPresent, Localized
MoroccoAbsent, Unconfirmed presence record(s)
NigerPresent
NigeriaPresent
RéunionPresent, Widespread1991
SenegalPresent
South AfricaPresent, Widespread
SudanPresent
TanzaniaPresent
TunisiaPresent, Localized
UgandaPresent
ZimbabwePresent, Localized

Asia

AfghanistanPresent
ArmeniaPresent
AzerbaijanPresent
BangladeshPresent
ChinaPresent
-BeijingPresent
-ChongqingPresent
-GansuPresent
-GuangdongPresent
-GuizhouPresent
-HeilongjiangPresent
-HubeiPresent
-LiaoningPresent
-NingxiaPresent
-QinghaiPresent
-ShaanxiPresent
-ShandongPresentOriginal citation: Sun et al. (2016)
-SichuanPresent
-TianjinPresent
-YunnanPresent
GeorgiaPresent
IndiaPresent, Localized
-Andhra PradeshPresent
-AssamPresent
-HaryanaPresent
-Himachal PradeshPresent
-KarnatakaPresent
-KeralaPresent
-Madhya PradeshPresent
-MaharashtraPresent
-Tamil NaduPresent
-TelanganaPresent
-Uttar PradeshPresent
IndonesiaPresent
-JavaPresent
IranPresent
IsraelPresent, Localized1992
JapanPresent
-HokkaidoPresent
-HonshuPresent
-Ryukyu IslandsPresent
JordanPresent, Localized
LebanonPresent, Localized
MalaysiaPresent
-Peninsular MalaysiaPresent
NepalPresent
North KoreaPresent
OmanPresent
PakistanPresent
Saudi ArabiaPresent
SingaporePresent, Few occurrences
South KoreaPresent
Sri LankaPresent
SyriaPresent
TaiwanPresent, Few occurrences
ThailandPresent
TurkeyPresent, Localized1981
UzbekistanAbsent, Unconfirmed presence record(s)

Europe

AlbaniaPresent, Localized
AustriaPresent, Few occurrences
BelgiumPresent, Localized
Bosnia and HerzegovinaPresent, Few occurrences
BulgariaPresent, Widespread1952
CroatiaPresent, Localized1977
CyprusPresent, Localized
CzechiaPresent, Localized1992
DenmarkAbsent, Eradicated1990
EstoniaAbsent, Formerly present
FinlandPresent, Transient under eradication1989
FrancePresent, Localized1987
GermanyPresent, Localized
GreecePresent, Widespread1972
-CretePresent
GuernseyPresent, Widespread
HungaryPresent, Widespread
IrelandPresent, Localized
ItalyPresent, Widespread1989
-SardiniaPresent
-SicilyPresent
LatviaAbsent, Eradicated
LithuaniaPresent, Few occurrences
MaltaPresent
MoldovaPresent, Widespread
MontenegroPresent
NetherlandsPresent1987
North MacedoniaPresentOriginal citation: Trajcbreve~evski (2002)
NorwayAbsent, Eradicated
PolandPresent
PortugalPresent, Localized1991
-MadeiraPresent
RomaniaPresent, Few occurrences
RussiaPresent
-Russian Far EastPresent
-Southern RussiaPresent
SerbiaPresent
Serbia and MontenegroPresent
SlovakiaAbsent, Unconfirmed presence record(s)1993
SloveniaPresent, Localized
SpainPresent, Widespread
-Balearic IslandsPresent, Localized
-Canary IslandsPresent
SwedenPresent, Few occurrences1990
SwitzerlandPresent, Localized
UkrainePresent
United KingdomPresent, Widespread1986
-Channel IslandsPresent
-EnglandPresent, Widespread
-ScotlandPresent, Widespread

North America

CanadaPresent, Localized
-AlbertaPresent
-British ColumbiaPresent
-ManitobaPresent
-Nova ScotiaPresent
-OntarioPresent
-QuebecPresent
-SaskatchewanPresent
Costa RicaPresent
Dominican RepublicPresent
HaitiPresent, Localized
JamaicaPresent
MartiniqueAbsent, Intercepted only
MexicoPresent, Localized
Puerto RicoPresent
United StatesPresent, Widespread
-AlabamaPresent
-ArkansasPresent
-CaliforniaPresent
-ConnecticutPresent
-DelawarePresent
-FloridaPresent
-GeorgiaPresent
-HawaiiPresent
-IdahoPresent
-IndianaPresent
-IowaPresent
-KansasPresent
-KentuckyPresent
-LouisianaPresent
-MainePresent
-MassachusettsPresent
-MichiganPresent
-MinnesotaPresent
-MississippiPresent
-MissouriPresent
-MontanaPresent
-NebraskaPresent
-NevadaPresent
-New HampshirePresent
-New MexicoPresent
-New YorkPresent
-North CarolinaPresent
-North DakotaPresent
-OhioPresent
-OklahomaPresent
-OregonPresent
-PennsylvaniaPresent
-South CarolinaPresent
-South DakotaPresent
-TennesseePresent
-TexasPresent
-UtahPresent
-VermontPresent
-VirginiaPresent
-WashingtonPresent
-WisconsinPresent
-WyomingPresent

Oceania

AustraliaPresent, Widespread
-New South WalesPresent
-Northern TerritoryPresent
-QueenslandPresent
-South AustraliaPresent
-TasmaniaPresent
-VictoriaPresent
-Western AustraliaPresent
Cook IslandsPresent
New ZealandPresent, Widespread
Papua New GuineaPresent

South America

ArgentinaPresent
BoliviaPresent
BrazilPresent, Localized
-BahiaPresent
-GoiasPresent
-Minas GeraisPresent
-ParanaPresent
-PernambucoPresent
-Sao PauloPresent
ChilePresent, Widespread
ColombiaPresent
EcuadorPresent, Few occurrences
GuyanaPresent
ParaguayPresent, Localized
SurinamePresent
UruguayPresent, Widespread
VenezuelaPresent

History of Introduction and Spread

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TSWV is present in temperate, tropical and subtropical climate zones and its distribution can be considered to be worldwide. The disease caused by TSWV was first described in 1915 in Australia and in the 1930s it was detected in Hawaii and in different European countries. The effective control of its main vector, Thrips tabaci, caused a decrease in its incidence in western Europe, although it remained present in eastern Europe and northern Greece in tobacco plants. In the 1980s, TSWV was once again a major threat to global horticulture, due to the emergence in the western USA of a new, more efficient vector, Frankliniella occidentalis, which spread the virus in both hemispheres.

Risk of Introduction

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TSWV is not at present listed as a quarantine pest by EPPO, but is of quarantine significance. EPPO at one time considered potato strains as A1 quarantine pests under the heading non-European potato viruses, but this idea has now been overtaken by events: TSWV is much more immediately threatening on hosts other than potato than as a theoretical non-European strain on potato. The recent introduction and widespread occurrence of the vector Frankliniella occidentalis in the EPPO region has provided a means of rapid dissemination of the virus. Those affected parts of the region have shown the potential importance of the virus throughout, especially for glasshouse cultivation of ornamentals and vegetables. However, it should be noted that both vector and virus are now present in the majority of EPPO countries. Only a rather small group of countries is apparently free from TSWV.

Hosts/Species Affected

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Capsicum annuum, lettuce, pea, tobacco, potato, tomato, and a large number of ornamental plant species are the main hosts of TSWV. Groundnut is a host of TSWV in the USA (causing groundnut ring mosaic), whereas two other orthospoviruses, GBNV and PYSV, are common on groundnut in South-East Asia. TSWV is a real threat to the tobacco industry in the Balkans, to tomato in some regions in Brazil, South Africa and other countries.

TSWV has an extremely wide host range with more than 1300 plants including agricultural crops, wild and weed species (Parrella et al., 2003). This include plant species in 15 monocotyledonous and 69 dicotyledonous families and one family of the Pteridophyta (Turina et al., 2012).

Groundnut ringspot orthotospovirus (GRSV), Tomato chlorotic spot orthotospovirus (TCSV), Chrysanthemum stem necrosis orthotospovirus (CSNV) and TSWV were identified in tomato, lettuces, endives and pepper crops in Sao Paulo, Brazil, and TCSV was the most prevalent virus (Colariccio et al., 2001). In a study of the geographical distribution of orthotospoviruses affecting tomato crops in Argentina, 63% of samples were found to be GRSV, 28.2% were TCSV and corresponding to a restricted area only 8.8% were TSWV (Williams et al., 2001). In tomato and pepper fields in San Francisco Valley and Federal District, USA, 53.5% of samples analysed by ELISA were positive to GRSV, 11.6% to TSWV, 1.9% to TCSV; Impatiens necrotic spot orthotospovirus (INSV) was not detected in this survey (Lima et al., 2000). With 189 recorded susceptible host species, INSV is a specialist pathogen of ornamental plants (Daughtrey et al., 1997). Double infections with TSWV and INSV have been detected in several ornamentals (Daughtrey, 1996; Kaminska et al., 1997). CSNV is probably the main orthotospovirus species infecting chrysanthemum in Brazil, whereas TSWV is the most frequently found orthotospovirus in chrysanthemum in Europe and the USA. Watermelon bud necrosis orthotospovirus (WBNV) can cause severe yield losses in India in watermelon. Infections of Iris yellow spot orthotospovirus (IYSV) occur in onion in Iowa, USA, and in Bahia and Pernambuco, Brazil, where it causes a disease known as 'sapeca' (Ávila et al., 1998).

Weeds and cultivated species are hosts (Jorda et al., 1995); the role of weeds as possible inoculum sources has been stressed (Cho et al., 1987; Bautista et al., 1995; Lavina et al., 1996; Ochoa et al., 1999; Groves et al., 2001).

In Europe and the Mediterranean region, the principal vegetable and industrial host crops, susceptible to TSWV, are artichoke (Cynara scolymus), aubergine (Solanum melongena), Capsicum annuum, chicory (Cichorium spp.), cucurbit, faba bean (Vicia faba), lettuce, potato (Solanum tuberosum), tobacco and tomato. Some of the principal ornamental species are Alstroemeria, Anemone, Aster, Begonia, Calceolaria, Callistephus, chrysanthemum (Dendranthema morifolium), cineraria (Senecio cruentus), Cyclamen, Dahlia, Gerbera, gloxinia (Sinningia spp.), Pelargonium, Ranunculus, Tagetes and Zinnia. Impatiens hybrids are important hosts of TSWV and INSV in New Guinea. The latter is usually found on other ornamentals (for example, cyclamen, kalanchoe) although not on vegetable crops. In Portugal, it has only been recorded on ornamentals (Louro, 1996) and in the Netherlands on almost 40 ornamentals and only on one vegetable, C. annuum (Verhoeven and Roenhorst, 1998). In addition to several ornamentals, three vegetables: Cichorium endivia, Ocimum basilicum and Valerianella olitoria have been recorded as hosts in France (Llamas-Bousquet and Berling, 1993). This virus has been found occasionally in chrysanthemum in the USA (Daughtrey et al., 1997). It rarely infects and spreads in solanaceous greenhouse and field crops. Infections by INSV in sweet pepper in the Netherlands caused symptoms that were restricted to the fruits. Restriction of symptoms to fruits has also been observed in tomato in Brazil (Pavan et al., 1996). Ruter and Gitaitis (1993) report INSV from a number of woody ornamentals (TSWV is not known to infect any woody plants); it has also, unusually, been reported from a fern (Asplenium nidus-avis; Lavina and Battle, 1994) and from prickly pear cactus (Opuntia microdasys; Blockley and Mumford, 2001). Wild species, including Arctium lappa, Datura stramonium, Emilia sonchifolia, Senecio vulgaris, Solanum nigrum, Sonchus spp. and Stellaria media may be important reservoirs of TSWV (Bautista et al., 1995).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Acalypha australisEuphorbiaceaeWild host
Adenium obesumApocynaceaeUnknown
Agapanthus praecoxUnknown
Agapanthus praecox subsp. orientalisLiliaceaeMain
Agastache foeniculumLamiaceaeUnknown
Ageratum conyzoides (billy goat weed)AsteraceaeMain
Alcea rosea (Hollyhock)MalvaceaeWild host
Allium cepa (onion)LiliaceaeOther
Allium cepa var. aggregatum (shallot)LiliaceaeMain
Allium porrum (leek)LiliaceaeOther
Nischwitz et al. (2006)
Allium sativum (garlic)LiliaceaeOther
Alstroemeria (Inca lily)AlstroemeriaceaeMain
Amaranthus (amaranth)AmaranthaceaeWild host
Amaranthus blitum (livid amaranth)AmaranthaceaeWild host
Amaranthus retroflexus (redroot pigweed)AmaranthaceaeOther
Amaranthus thunbergiiAmaranthaceaeOther
Amaranthus viridis (slender amaranth)AmaranthaceaeUnknown
Ambrosia artemisiifolia (common ragweed)AsteraceaeWild host
Ananas comosus (pineapple)BromeliaceaeMain
Anemone (windflower)RanunculaceaeMain
Anemone coronaria (Poppy anemone)RanunculaceaeMain
Anthemis (chamomile)AsteraceaeUnknown
Anthemis arvensisUnknown
Anthemis tinctoriaAsteraceaeUnknown
Antirrhinum majus (snapdragon)ScrophulariaceaeUnknown
Apium graveolens (celery)ApiaceaeMain
Arachis hypogaea (groundnut)FabaceaeMain
Cho et al. (2020); Golnaraghi et al. (2001)
Arctium lappa (burdock)AsteraceaeWild host
Arctotheca calendula (capeweed)AsteraceaeWild host
Arctotis x hybridaAsteraceaeOther
Argyranthemum frutescensAsteraceaeUnknown
Aristolochia clematitis (Birthwort)AristolochiaceaeOther
Artemisia princeps (Japanese mugwort)AsteraceaeWild host
Artemisia vulgaris (mugwort)AsteraceaeUnknown
Arum maculatumAraceaeUnknown
Asclepias curassavica (bloodflower)AsclepiadaceaeOther
Asplenium nidus (bird's nest fern)AspleniaceaeWild host
AsterAsteraceaeMain
Atriplex patula (common orache)ChenopodiaceaeUnknown
Avena fatua (wild oat)PoaceaeUnknown
Ballota nigraLamiaceaeUnknown
BegoniaBegoniaceaeMain
Benincasa hispida (wax gourd)CucurbitaceaeMain
Bidens pilosa (blackjack)AsteraceaeMain
Boerhavia erectaUnknown
Brassica juncea (mustard)BrassicaceaeWild host
Brassica napus var. oleiferaBrassicaceaeUnknown
Brassica rapa (field mustard)BrassicaceaeWild host
Brassica rapa subsp. campestrisBrassicaceaeMain
BrugmansiaSolanaceaeOther
Brugmansia suaveolens (white angel's trumpet)SolanaceaeOther
Calceolaria (pouch flower)ScrophulariaceaeMain
Calendula officinalis (Pot marigold)AsteraceaeMain
CallistephusAsteraceaeMain
Callistephus chinensis (China aster)AsteraceaeMain
Calystegia sepium (great bindweed)ConvolvulaceaeWild host
Campanula mediumCampanulaceaeOther
Canavalia gladiata (sword bean)FabaceaeMain
Canna indica (canna lilly)CannaceaeMain
Capsella bursa-pastoris (shepherd's purse)BrassicaceaeWild host
Capsicum (peppers)SolanaceaeMain
Capsicum annuum (bell pepper)SolanaceaeMain
Capsicum chinense (habanero pepper)SolanaceaeUnknown
Capsicum frutescens (chilli)SolanaceaeUnknown
Cardamine flexuosa (wavy bittercress)BrassicaceaeWild host
Cardamine hirsuta (hairy bittercress)BrassicaceaeUnknown
Cardamine parvifloraBrassicaceaeWild host
Carduus nutans (nodding thistle)AsteraceaeUnknown
Carica papaya (pawpaw)CaricaceaeMain
Catharanthus roseus (Madagascar periwinkle)ApocynaceaeMain
Centaurea (Knapweed)AsteraceaeUnknown
Cerastium glomeratumCaryophyllaceaeWild host
Chaerophyllum temulumApiaceaeUnknown
Chamomilla recutita (common chamomile)AsteraceaeUnknown
Chamomilla suaveolens (Rounded chamomile)AsteraceaeUnknown
Chenopodiastrum murale (nettleleaf goosefoot)ChenopodiaceaeUnknown
Chenopodium album (fat hen)ChenopodiaceaeUnknown
Chenopodium ficifolium (Fig-leaved goosefoot)ChenopodiaceaeWild host
Chenopodium giganteum (large lambsquarters)ChenopodiaceaeUnknown
Gera et al. (2000)
Chenopodium glaucum (Oak-leaved goosefoot)ChenopodiaceaeUnknown
Chenopodium quinoa (quinoa)ChenopodiaceaeUnknown
Gera et al. (2000)
Chondrilla juncea (rush skeletonweed)AsteraceaeUnknown
Chrysanthemum (daisy)AsteraceaeOther
Chrysanthemum coronarium (garland chrysanthemum)AsteraceaeMain
Chrysanthemum frutescens (marguerite)AsteraceaeUnknown
Chrysanthemum morifolium (chrysanthemum (florists'))AsteraceaeMain
Cicer arietinum (chickpea)FabaceaeMain
Cichorium (chicory)AsteraceaeMain
Cichorium endivia (endives)AsteraceaeMain
Cichorium intybus (chicory)AsteraceaeUnknown
Cirsium (thistle)AsteraceaeUnknown
Cirsium arvense (creeping thistle)AsteraceaeWild host
Citrullus lanatus (watermelon)CucurbitaceaeMain
Clematis flammulaRanunculaceaeUnknown
Clematis vitalba (old man's beard)RanunculaceaeUnknown
Cleome viscosa (Asian spiderflower)CapparaceaeUnknown
ColeusLamiaceaeMain
ColumneaGesneriaceaeMain
Columnea hirtaGesneriaceaeMain
Commelina communis (common dayflower)CommelinaceaeWild host
Conium maculatum (poison hemlock)ApiaceaeUnknown
Convolvulus arvensis (bindweed)ConvolvulaceaeWild host
Conyza canadensis (Canadian fleabane)AsteraceaeWild host
Coprosma repensRubiaceaeOther
Coronopus squamatusBrassicaceaeUnknown
CrepisAsteraceaeWild host
Crotalaria juncea (sunn hemp)FabaceaeMain
Crotalaria spectabilis (showy rattlepod)FabaceaeUnknown
Cucumis sativus (cucumber)CucurbitaceaeMain
Cucurbita moschata (pumpkin)CucurbitaceaeOther
Cucurbita pepo (marrow)CucurbitaceaeMain
Cuscuta (dodder)CuscutaceaeUnknown
CyclamenPrimulaceaeMain
Cynara cardunculus var. scolymus (globe artichoke)AsteraceaeMain
Cyphomandra betacea (tree tomato)SolanaceaeMain
DahliaAsteraceaeMain
Datura stramonium (jimsonweed)SolanaceaeWild host
Gera et al. (2000); Chatzivassiliou et al. (2001)
Daucus carota (carrot)ApiaceaeMain
Dianthus chinensis (china pink)CaryophyllaceaeWild host
Dieffenbachia (dumbcanes)AraceaeMain
Diplotaxis erucoidesBrassicaceaeOther
Ecballium elateriumCucurbitaceaeUnknown
Echinops ritro (small globe-thistle)AsteraceaeUnknown
Eclipta prostrata (eclipta)AsteraceaeWild host
Emilia sonchifolia (red tasselflower)AsteraceaeWild host
ErodiumUnknown
Erodium ciconiumGeraniaceaeUnknown
Erodium moschatumGeraniaceaeWild host
Eucharis × grandifloraEucharitidaeOther
Eupatorium capillifolium (Dog fennel)AsteraceaeUnknown
Eustoma grandiflorum (Lisianthus (cut flower crop))GentianaceaeMain
Ficus elastica (rubber plant)MoraceaeMain
Ficus pumila (creeping fig)MoraceaeMain
Forsythia viridissimaOleaceaeWild host
Fritillaria thunbergiiLiliaceaeUnknown
Fumaria officinalis (common fumitory)PapaveraceaeOther
Galinsoga parviflora (gallant soldier)AsteraceaeMain
Galium aparine (cleavers)RubiaceaeUnknown
Galium spuriumRubiaceaeWild host
Geranium carolinianum (Carolina geranium)GeraniaceaeUnknown
Gerbera (Barbeton daisy)AsteraceaeMain
Gerbera jamesonii (African daisy)AsteraceaeMain
Glycine max (soyabean)FabaceaeMain
Yoon et al. (2018); Golnaraghi et al. (2001); Golnaraghi et al. (2002); Nischwitz et al. (2006); Golnaraghi et al. (2004); Sikora et al. (2011)
Gnaphalium purpureumAsteraceaeUnknown
Gomphrena globosa (globe amaranth)AmaranthaceaeUnknown
Gera et al. (2000)
Gossypium (cotton)MalvaceaeMain
Gypsophila elegans (baby's breath)CaryophyllaceaeOther
Helianthus annuus (sunflower)AsteraceaeMain
Heliotropium europaeum (common heliotrope)BoraginaceaeUnknown
Helminthotheca echioides (bristly oxtongue)AsteraceaeUnknown
Hibiscus trionum (Venice mallow)MalvaceaeOther
HostaLiliaceaeOther
Momol et al. (2018); Momol et al. (2003)
Hoya carnosa (Wax plant)AsclepiadaceaeMain
Humulus scandens (Japanese hop)CannabaceaeWild host
Yoon et al. (2018); Yoon et al. (2018)
Iberis semperflorensBrassicaceaeOther
Parrella et al. (2013)
Impatiens (balsam)BalsaminaceaeMain
Impatiens walleriana (busy lizzy)BalsaminaceaeMain
Ipomoea hederaceaConvolvulaceaeUnknown
Ipomoea purpurea (tall morning glory)ConvolvulaceaeUnknown
Iris domestica (blackberry lily)IridaceaeMain
Jacquemontia tamnifolia (Smallflower morningglory)Main
KalanchoeCrassulaceaeMain
Lactuca sativa (lettuce)AsteraceaeMain
Lactuca serriola (prickly lettuce)AsteraceaeUnknown
Lamium amplexicaule (henbit deadnettle)LamiaceaeWild host
Lamium purpureum (purple deadnettel)LamiaceaeUnknown
Lathyrus sativus (grass pea)FabaceaeMain
Lens culinaris subsp. culinaris (lentil)FabaceaeMain
Lepidium didymum (lesser swine-cress)BrassicaceaeUnknown
Lepidium virginicum (Virginian peppercress)BrassicaceaeWild host
Leuzea carthamoidesAsteraceaeOther
Linaria canadensisScrophulariaceaeUnknown
Lolium perenne (perennial ryegrass)PoaceaeUnknown
Lupinus (lupins)FabaceaeMain
Lycium chinense (chinese wolfberry)SolanaceaeMain
LycopersiconSolanaceaeOther
Lycopus europaeus (European water horehound)LamiaceaeUnknown
Malva neglecta (common mallow)MalvaceaeUnknown
Malva sylvestrisMalvaceaeWild host
Malva verticillataMalvaceaeMain
Medicago polymorpha (bur clover)FabaceaeWild host
Melilotus officinalis (yellow sweet clover)FabaceaeWild host
Mentha piperita (Peppermint)LamiaceaeMain
Mentha suaveolensLamiaceaeUnknown
Mirabilis jalapa (four o'clock flower)NyctaginaceaeMain
Mollugo verticillataUnknown
Morus alba (mora)MoraceaeMain
Myosoton aquaticumCaryophyllaceaeWild host
Nicandra physalodes (apple of Peru)SolanaceaeMain
Nicotiana benthamianaSolanaceaeUnknown
Gera et al. (2000)
Nicotiana glutinosaUnknown
Gera et al. (2000)
Nicotiana rustica (wild tobacco)SolanaceaeMain
Gera et al. (2000)
Nicotiana tabacum (tobacco)SolanaceaeMain
OcimumLamiaceaeMain
Ocimum basilicum (basil)LamiaceaeMain
Oenanthe javanicaApiaceaeMain
Oncidium (dancing-lady orchid)OrchidaceaeMain
OrnithogalumLiliaceaeOther
Osteospermum ecklonisAsteraceaeOther
Oxalis acetosellaOxalidaceaeOther
Paederia foetida (skunkvine)RubiaceaeWild host
Panicum repens (torpedo grass)PoaceaeUnknown
Papaver rhoeas (common poppy)PapaveraceaeUnknown
Pelargonium (pelargoniums)GeraniaceaeMain
Peperomia obtusifolia (pepper-face)PiperaceaeOther
Pericallis cruenta (common cineraria)AsteraceaeMain
Persicaria pensylvanicaPolygonaceaeUnknown
PetuniaSolanaceaeMain
Petunia hybridaSolanaceaeMain
Gera et al. (2000)
PhalaenopsisOrchidaceaeMain
Phaseolus (beans)FabaceaeMain
Phaseolus vulgaris (common bean)FabaceaeMain
Phragmites australis (common reed)PoaceaeUnknown
Physalis ixocarpaSolanaceaeUnknown
Physalis peruviana (Cape gooseberry)SolanaceaeMain
Physalis philadelphicaSolanaceaeOther
Phytolacca americana (pokeweed)PhytolaccaceaeWild host
Pinus elliottii (slash pine)PinaceaeUnknown
Pinus palustris (longleaf pine)PinaceaeUnknown
Pinus taeda (loblolly pine)PinaceaeUnknown
Pisum sativum (pea)FabaceaeMain
Pittosporum tobira (Japanese pittosporum)PittosporaceaeMain
Liu et al. (2016); Gera et al. (2000)
Plantago lanceolata (ribwort plantain)PlantaginaceaeUnknown
Plantago major (broad-leaved plantain)PlantaginaceaeUnknown
Platycodon grandiflorus (Balloonflower)CampanulaceaeOther
Poa annua (annual meadowgrass)PoaceaeWild host
Polygonum aviculare (prostrate knotweed)PolygonaceaeUnknown
Portulaca oleracea (purslane)PortulacaceaeWild host
Potentilla reptans (sulfur cinquefoil)RosaceaeUnknown
Ranunculus (Buttercup)RanunculaceaeWild host
Ranunculus asiaticus (garden crowfoot)RanunculaceaeWild host
Ranunculus bulbosus (bulbous buttercup)RanunculaceaeUnknown
Ranunculus sardousRanunculaceaeUnknown
Raphanus raphanistrum (wild radish)BrassicaceaeUnknown
Raphanus sativus (radish)BrassicaceaeMain
Rhaponticum carthamoidesAsteraceaeUnknown
Robinia pseudoacacia (black locust)FabaceaeWild host
Rorippa indica (Indian marshcress)BrassicaceaeWild host
Rumex (Dock)PolygonaceaeWild host
Rumex crispus (curled dock)PolygonaceaeUnknown
Saintpaulia ionantha (African violet)GesneriaceaeMain
Salvia officinalis (common sage)LamiaceaeMain
Sanguisorba minorRosaceaeUnknown
Saponaria officinalis (soapwort)CaryophyllaceaeUnknown
Scabiosa (Scabious)DipsacaceaeUnknown
Sechium edule (chayote)CucurbitaceaeMain
Sedum sarmentosumCrassulaceaeWild host
Senecio vulgarisAsteraceaeWild host
Senna obtusifolia (sicklepod)FabaceaeUnknown
Sesamum indicum (sesame)PedaliaceaeMain
SileneUnknown
Silene latifolia subsp. alba (white campion)CaryophyllaceaeUnknown
Silybum marianum (variegated thistle)AsteraceaeUnknown
Sinapis (mustard)BrassicaceaeOther
Sinapis arvensis (wild mustard)BrassicaceaeUnknown
SinningiaGesneriaceaeMain
Sinningia speciosa (gloxinia)GesneriaceaeMain
SolanaceaeSolanaceaeMain
Solanum carolinense (horsenettle)SolanaceaeUnknown
Solanum elaeagnifolium (silverleaf nightshade)SolanaceaeUnknown
Solanum lycopersicum (tomato)SolanaceaeMain
Solanum melongena (aubergine)SolanaceaeMain
Solanum nigrum (black nightshade)SolanaceaeWild host
Solanum pimpinellifolium (currant tomato)SolanaceaeUnknown
Solanum tuberosum (potato)SolanaceaeMain
Solidago (Goldenrod)AsteraceaeUnknown
Sonchus (Sowthistle)AsteraceaeWild host
Sonchus arvensis (perennial sowthistle)AsteraceaeWild host
Sonchus asper (spiny sow-thistle)AsteraceaeWild host
Sonchus oleraceus (common sowthistle)AsteraceaeWild host
Sorghum halepense (Johnson grass)PoaceaeUnknown
SpathiphyllumAraceaeUnknown
Spinacia oleracea (spinach)ChenopodiaceaeOther
Stellaria media (common chickweed)CaryophyllaceaeWild host
Stephanotis floribunda (madagascar stephanotis)AsclepiadaceaeMain
Stevia rebaudianaAsteraceaeOther
Suaeda fruticosa (Shrubby seablite)ChenopodiaceaeUnknown
Tagetes (marigold)AsteraceaeMain
Tanacetum cinerariifolium (Pyrethrum)Wild host
Taraxacum officinale complex (dandelion)AsteraceaeWild host
Tephrosia purpurea (purple tephrosia)FabaceaeMain
Tragopogon dubiusAsteraceaeOther
Tragopogon mirusAsteraceaeOther
Tragopogon porrifolius (oysterplant)AsteraceaeOther
Tragopogon pratensisAsteraceaeOther
Tribulus terrestris (puncture vine)ZygophyllaceaeUnknown
Trichosanthes kirilowiiCucurbitaceaeWild host
Trifolium (clovers)FabaceaeUnknown
Trifolium repens (white clover)FabaceaeWild host
Tropaeolum majus (common nasturtium)TropaeolaceaeWild host
Tulbaghia violaceaWild host
Valeriana officinalis (common valerian)ValerianaceaeMain
Valerianella locusta (common cornsalad)ValerianaceaeMain
Verbena officinalis (vervain)VerbenaceaeUnknown
Veronica chamaedrysScrophulariaceaeUnknown
Veronica officinalisScrophulariaceaeUnknown
Veronica persica (creeping speedwell)ScrophulariaceaeOther
Vicia (vetch)FabaceaeUnknown
Vicia amoenaFabaceaeWild host
Vicia faba (faba bean)FabaceaeMain
Vicia hirsuta (hairy tare (UK))FabaceaeWild host
Vigna mungo (black gram)FabaceaeMain
Vigna radiata (mung bean)FabaceaeMain
Vigna unguiculata (cowpea)FabaceaeMain
Xanthium spinosum (bathurst burr)AsteraceaeUnknown
Xanthium strumarium (common cocklebur)AsteraceaeUnknown
Youngia japonica (oriental false hawksbeard)AsteraceaeWild host
Zantedeschia aethiopica (calla lily)AraceaeMain
ZinniaAsteraceaeMain
Zinnia elegans (zinnia)AsteraceaeMain

Growth Stages

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Fruiting stage

Symptoms

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Symptoms are illustrated by reference to a selection of economically important vegetable, ornamental and industrial crop species. TSWV can induce a wide variety of symptoms that may vary on the same host species with cultivar, age and nutritional and environmental conditions. Isolates of TSWV usually do not differ in biological properties and differ only slightly in molecular and serological properties. A study of the diversity of eight TSWV isolates collected from north-eastern Spain showed a slight biological variability among the isolates when compared in 10 host species; no differences in transmission efficiencies were found among them (Roca et al., 1997).

Symptoms evoked by other orthotospovirus species do not differ principally from those caused by TSWV. The greatest care should be used to distinguish the orthotospovirus species by symptoms alone.

Inoculation of solanaceous species may result in the generation of defective interfering RNAs. These RNAs are formed by deletions in the L RNA. Isolates containing these DI RNAs usually have attenuated symptoms (Resende et al., 1991b; Inoue-Nagata et al., 1997) and are often poorly transmitted by thrips (Nagata et al., 1999).

On tomatoes, plants show bronzing, curling, necrotic streaks and spots on the leaves. Dark-brown streaks also appear on leaf petioles, stems and growing tips. The plants are small and stunted. The carotene, chlorophyll and xanthophyll levels decreased. The ripe fruit shows paler red or yellow areas on the skin. Sometimes affected plants are killed by severe necrosis. Symptoms are occasionally only found on the fruits (Pavan et al., 1996). Some fruits of TSWV-resistant plants can show peculiar ringspot symptoms caused as consequence of an insufficient hypersensitive response by the feeding of viruliferous thrips on the fruit during the early stages of development (de Haan et al., 1996; Aramburu et al., 2000).

On Capsicum, symptoms are mainly stunting and yellowing of the whole plant. Leaves may show chlorotic line patterns or mosaic with necrotic spots. Necrotic streaks appear on stems extending to the terminal shoots. On ripe fruits, yellow spots with concentric rings or necrotic streaks have been observed. On lettuces, infection starts in leaves on one side of the plant, which becomes chlorotic with brown patches. The discoloration extends to the heart leaves and cessation of growth on one side of the plant produces characteristic distortion.

On chrysanthemums, there is a wide variation among cultivars. Usually black stem streaks and wilt are observed. On gloxinias, infected leaves show yellow or brown leaf spotting, or brown oak-leaf patterns. On Impatiens, some cultivars of New Guinea hybrids infected with INSV and TSWV develop stunting, black discoloration at the base of the leaf, or brown leaf spots. On groundnuts, symptoms of the disease now attributed to a distinct orthotospovirus, groundnut bud necrosis are bud necrosis, chlorosis of foliage, limb collapse and plant death.

List of Symptoms/Signs

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SignLife StagesType
Fruit / abnormal shape
Fruit / discoloration
Leaves / abnormal colours
Leaves / abnormal forms
Leaves / necrotic areas
Whole plant / dwarfing

Biology and Ecology

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All orthotospoviruses are transmitted and spread in nature by insects of the family Thripidae (Thysanoptera), belonging to the genera Frankliniella and Thrips. They include Frankliniella bispinosa (Webb et al., 1998), F. intonsa (Wijkamp et al., 1995), F. fusca, F. occidentalis (EPPO/CABI, 1992), F. schultzei, Thrips palmi, T. setosus and T. tabaci. T. flavus (Singh and Krishnareddy, 1996), F. tenuicornis (Kormelink, 1994) and Scirtothrips dorsalis (Amin et al., 1981) have been reported as vectors, but their status as such has yet to be confirmed.

The type of transmission is persistent, circulative and propagative. The virus is acquired only by the larvae and transmitted by late second instars and adults. The former will transmit when the virus is acquired by early first instars. Virus is acquired by more intensive feeding, and inoculation can be achieved by shallow feeding on leaf epidermal cells. The shortest reported acquisition and inoculation periods vary between 10 and 15 minutes for F. occidentalis and T. tabaci. The efficiency of transmission for F. occidentalis increases with feeding time, being 4% for 15 min., 33% for 1 hour and 77% for 4 days (Roselló et al., 1996) and is higher for males than females. F. occidentalis needs mean access periods of 1 hour to acquire and 1-2 hours to transmit TSWV (Wijkamp et al., 1996b). The ability of F. occidentalis populations to acquire orthotospovirus decreases with the age of the larvae. Larvae that acquire the virus early can transmit the virus (60-80%) when they are in the late second instars. The latent (incubation) period of TSWV and Impatiens necrotic spot virus (INSV) in F. occidentalis does not significantly differ and depends on temperature. The mean latent period is about 10 days, in which the larvae pupates and emerges as an adult (Roselló et al., 1996). Consequently, the virus is retained when the vector moults and is transmitted by the adults; although, virus transmission is intermittent (Wijkamp et al., 1993). There is no evidence of transovarial transmission (Wijkamp et al., 1996a). These observations show that TSWV and other orthotospoviruses are persistent in their vectors. TSWV and INSV have been serologically detected in viruliferous thrips (Cho et al., 1988; Wijkamp et al., 1993; Aramburu et al., 1996). The virus can also be detected in adults, which migrate from healthy to infected plants and feed on the latter.

In the western part of Europe and the Mediterranean region, TSWV, thought to be transmitted by T. tabaci, has been virtually absent since the Second World War. This disappearance is not understood. It is possible that this vector was present in relatively low numbers. The absence has also been explained by the almost complete disappearance of tobacco culture in Western Europe, a replacement of transmitting by non-transmitting T. tabaci populations, and the enhanced application of insecticides. F. occidentalis, indigenous to North America, began to spread internationally around 1980 and is now widely reported in Europe and the Mediterranean (OEPP/EPPO, 1989). The recent epidemic spread of TSWV in protected and outdoor crops in the European and Mediterranean region is closely associated with the establishment and rapid infestation by this efficient vector. INSV became important with the expansion of this vector. INSV and TSWV are efficiently transmitted by F. occidentalis. Groundnut ringspot orthotospovirus (GRSV) and Tomato chlorotic spot orthotospovirus (TCSV) are also, but less efficiently, transmitted by this species (Wijkamp et al., 1996b). Frankliniella schultzei is reported to be an important vector of GRSV in tomato in Brazil (Pavan et al., 1996).

 

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
B - Dry (arid and semi-arid) Preferred < 860mm precipitation annually
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C

Means of Movement and Dispersal

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The main pathways of TSWV dispersal are through infected plants for planting, pot plants, and through insects of the family Thripidae (Thysanoptera), belonging to the genera Frankliniella and Thrips, and associated with plants materials. Since the virus is vector-transmitted, the viruliferous vectors constitute the main pathway of local and natural dispersal. International introduction is likely through traded infected pot plants and plant material for planting.

Vector transmission

TSWV is liable to spread naturally with its vectors (OEPP/EPPO, 1989). All orthotospoviruses are transmitted and spread in nature by insects of the family Thripidae (Thysanoptera), belonging to the genera Frankliniella and Thrips. They include Frankliniella bispinosa (Webb et al., 1998), F. intonsa (Wijkamp et al., 1995), F. fuscaF. occidentalis (EPPO/CABI, 1992), F. schultzeiThrips palmiT. setosus and T. tabaciT. flavus (Singh and Krishnareddy, 1996), F. tenuicornis (Kormelink, 1994) and Scirtothrips dorsalis (Amin et al., 1981) have been reported as vectors, but their status as such has yet to be confirmed. The type of transmission is persistent, circulative and propagative. 

In international trade, TSWV may be carried by susceptible host plants, whether pot plants or plants for planting, and will be especially liable to spread if these plants also carry vectors.

Seed transmission has not been demonstrated.

Seedborne Aspects

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Seed Treatment

Seed treatment with thiamethoxam and imidacloprid reduced TSWV prevalence in seed potato by 75% in the early season and 66% in the late season (Pourrahim et al., 2008).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Breeding and propagation Yes Yes
Crop production Yes Yes
Cut flower trade Yes Yes
Horticulture Yes Yes
Nursery trade Yes Yes
Ornamental purposes Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Host and vector organisms Yes Yes
Plants or parts of plantsMultiplying virions that can be acquired by Thrips larvae (vector) Yes Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Flowers/Inflorescences/Cones/Calyx
Seedlings/Micropropagated plants

Impact Summary

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CategoryImpact
Economic/livelihood Negative

Impact

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TSWV and other orthotospoviruses have become an increasingly important factor contributing to economic losses in many food and ornamental crops throughout the world; losses may be as high as 100% (Berling et al., 1990; Rodriguez, 1990; Roselló et al., 1996). TSWV incidence in Brazil, Hawaii, USA, and South Africa can be so high that farmers are forced out of production. Destructive outbreaks of TSWV have occurred in France and Spain in protected and field crops of tomatoes, Capsicum and Anemone, associated with the establishment and rapid spread of the vector Frankliniella occidentalis (OEPP/EPPO, 1989). In Liguria, Italy, Capsicum production can be severely affected while adjacent tomato and lettuce crops remain healthy. Devastating epidemics can occur in tobacco in Bulgaria, Greece and other south-eastern European countries. In some areas of Argentina, Brazil, Italy and South Africa, TSWV has become one of the most important diseases in tomato. In general, economic loss data are limited, but the following examples of economic impact have been obtained from the literature.

In groundnuts, TSWV has been shown to reduce yield in direct proportion to the intensity of infection. Healthy plants produced 50% more kernels than plants with maximum infection by TSWV (Saharan et al., 1983). The virus reduces height, root length and yield depending on the plant growth stage at the time of infection (Rao et al., 1979). In plants showing symptoms within 45 days of sowing, 100% losses were observed. Losses decreased with increasing age of plants at infection. Gopal and Uphadhyaya (1991) reported yield losses of 50% in Raichur, India. Field trials indicated that early infection with the virus caused a heavy yield loss compared with late infection. Siddaramaiah et al. (1980) reported a drastic reduction in dried pod weight, fresh fodder weight and shelling percent in plants affected at an early stage. Narendrappa and Siddaramaiah (1986) also reported that infection of plants up to 65 days old caused significant reductions in yield while no losses were recorded in plants infected after 95 days. The incidence of bud necrosis (caused by TSWV) has been shown to be lower in close plant spacings than in wide ones (Anon., 1981). Culbreath et al. (1992) reported that the number of seed produced, average weight per seed and total seed yield were lower for TSWV symptomatic plants than for healthy plants in Georgia, USA, in 1988, 1989 and 1990. TSWV has also been reported to reduce the oil content in groundnuts; earlier infection causing greater losses. Plants infected 15 days before harvest showed a 13.2% oil reduction (Ali and Rao, 1982).

Fiederow and Kralowska (1995) reported decreases in yields of tomatoes and Capsicum of 38.7 and 92.2%, respectively. In tomatoes, earlier infection has been shown to cause greater yield losses (Kumar and Irulappen, 1991). A lower number of fruits, fruits weight, and yield were recorded in tomato plants infected at an early stage than those infected at more mature stages; however, the quality of production was altered even in late-infected plants due to abnormal ripening and no difference in marketable fruit yield was obtained (Moriones et al., 1998).

The biochemical changes caused in tomato following infection with TSWV have also been studied. Chlorophyll, xanthophylls and carotene levels decreased in infected plants (Sutha et al., 1998).

In peas in India, TSWV-affected plants were pale-green and stunted with reduced leaf petioles, stipules and tendrils. Only a few pods were produced and these were necrotic. Disease incidence ranged from 10 to 25%, causing serious yield losses (Singh and Gupta, 1994). In greenhouse tests in Canada during 1989-1990 and 1990-1991, TSWV affected Lathyrus sativus and Pisum sativum var. arvense. In L. sativus, symptoms varied from loss of chlorophyll, wilting and drying-up of the foliage to bleaching and drying-up of stem segments at the nodes. In P. sativum var. arvense, purplish-brown streaks were prominent on the stems and petioles. Flower and pod abortion occurred in severely affected plants (Zimmer et al., 1992).

In cucurbit and solanaceous vegetables in Okinawa, Japan, severe losses have been caused by TSWV. Cucumber plants inoculated at the cotyledon stage were shorter, lateral shoots were fewer and shorter and yields were 40% lower than in non-inoculated plants (Hokama and Tokahashi, 1987).

Dahlias in the Netherlands in 1992, 1993 and 1994 had infection levels of 15, 5 and 2%, respectively. Yield of cuttings/tuber was reduced by up to 20% (Asjes et al., 1997).

In Hawaii, USA, TWSV has destroyed 50-90% of lettuce crops (Cho et al., 1987).

Potato crops have been affected by TSWV in India (Khurana et al., 1997), Portugal, Brazil and Argentina (Granval de Millan et al., 1998). Doubt exists as to whether infected tubers will produce healthy plants when they sprout.

Risk and Impact Factors

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Invasiveness
  • Invasive in its native range
  • Has a broad native range
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Benefits from human association (i.e. it is a human commensal)
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Host damage
  • Increases vulnerability to invasions
  • Monoculture formation
  • Negatively impacts agriculture
Impact mechanisms
  • Parasitism (incl. parasitoid)
  • Pathogenic
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

Uses List

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General

  • Research model

Diagnosis

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Rapid immunofilter paper assay (RIPA) can detect the presence of TSWV in 30 minutes (López Lambertini et al., 2003). The use of ImmunoStrip tests to diagnose TSWV in tomato and lettuce samples is mentioned in Abou-Jawdah et al. (2006). For the diagnosis of TSWV in artichoke, a non-isotopic dot blot hybridisation protocol with DNA probes is detailed in Minutillo et al. (2012).

Both ELISA and RT-PCR are used for the diagnosis of TSWV in groundnut. There is no significant difference between the accuracy of these two assays (Dang et al., 2009).

Detection and Inspection

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TSWV is sap-transmissible to diagnostic species (Ie, 1970). Petunia hybrida is one of the most useful species because of the rapidity with which the brown local lesions develop (within 2-3 days under favourable conditions without systemic virus spread). This species is used as an indicator plant for the presence of viruliferous thrips in greenhouses (Allen and Matteoni, 1991). Leaf discs of this species are successfully used in transmission studies (Wijkamp and Peters, 1993). Nicotiana benthamiana, N. clevelandii, N. glutinosa, N. rustica and N. tabacum often show small to large local necrotic or ring-like lesions followed by systemic infection. The first symptom is then a vein chlorosis which develops into a mottling, mosaic or necrosis, or even death of the plant (N. benthamiana and N. clevelandii). N. glutinosa is usually selected to clone TSWV isolates by mechanical inoculation (de Avila et al., 1993). The number of lesions and their size, and the type and severity of the systemic symptoms depends on the virus isolate. N. benthamiana is perhaps the most susceptible species. Datura stramonium is sometimes preferred as a diagnostic host. Plant material co-infected with tobacco mosaic virus has to be tested on N. glutinosa and those with PVY on D. stramonium. Cucumis sativus shows chlorotic lesions with necrotic centres on cotyledons 4-5 days after inoculation. Some isolates give necrotic lesions on the true leaves.

Sap transmission of TSWV is usually efficient if young test plants and freshly prepared inocula are used. However, difficulties are sometimes encountered in transmitting the virus from older infected plants. Sap transmission efficiency can be greatly increased by using neutral phosphate buffers containing a reducing agent, for example, sodium sulfite. It is recommended to maintain the virus not in plants, but by storage of infected plant material at -70°C to prevent co-infection with other viruses, the generation of mutants (defective interference isolates) and loss of transmissibility by thrips.

Antisera to most species, often prepared to the nucleocapsid fraction, are now available. These antisera are often very specific and discriminate readily between the different species. Antisera with the desired quality to the complete virus are more difficult to prepare as the viruses are often difficult to purify.

ELISA is the most commonly used serological test for extracts from infected plants (Gonsalves and Trujillo, 1986; Resende et al., 1991a) and thrips (Cho et al., 1988; Wijkamp et al., 1993); however, the detection of infectious thrips by squash-blot assay provides a practical means of testing the large number of thrips needed for analysis in epidemiological studies (Aramburu et al., 1996) and direct tissue blotting was more sensitive than ELISA for detection of TSWV in leaf extracts (Hsu and Lawson, 1991). The double antibody sandwich (DAS) direct ELISA using polyclonal antibodies to the nucleocapsid protein is recommended to the use of antibodies to the other structural proteins for detecting the distinct species (Ávila et al.; 1990; Law and Moyer, 1990). Twenty TSWV isolates compared in ELISA using polyclonal and six monoclonal antibodies were differentiated into two serogroups and three serotypes (de Avila et al., 1990). When tested in different forms of ELISA, TSWV does not exhibit great variations in its response using one and the same antiserum (Adam et al., 1996). Adam et al. (1995) described an assay using antibodies to the G proteins of TSWV which will detect orthotospoviruses generally.

Groundnut ringspot orthotospovirus (RSV) and Tomato chlorotic spot orthotospovirus (TCSV) react faintly with antisera to TSWV (Ávila et al., 1993). The other orthotospovirus species do not react with antisera to TSWV, TCSV and GRSV. Watermelon silver mottle orthotospovirus (WSMV), Groundnut bud necrosis orthotospovirus (GBNV) and Watermelon bud necrosis orthotospovirus (WBNV) are serologically related, although the reaction between their N sera is also weak. Iris yellow spot orthotospovirus (IYSV) and the Brazilian IYSV isolate show almost identical reactions. The species Chrysanthemum stem necrosis orthotospovirus (CNSV), Impatiens necrotic spot orthotospovirus (INSV), Physalis severe mottle virus (PSMV), Peanut chlorotic fan-spot virus (PCFV), Peanut yellow spot virus (PYSV) and Zucchini lethal chlorosis orthotospovirus (ZLCV) do not mutually show any cross-reaction and also show no relationship with other species.

The use of cDNA probes (Ronco et al., 1989; Rice et al., 1990) and riboprobes (Huguenot et al., 1990) has been proposed but is not yet widely applied in the identification and diagnosis of TSWV. A PCR-based assay has been developed by Mumford et al. (1994). A method based on immunocapture and PCR amplification has been reported by Nolasco et al. (1993). Immunocapture and detection by nucleic acid hybridization as a diagnostic method for TSWV is mentioned in Martínez et al. (2005). The use of a polyprobe method for the detection of TSWV in tomato is detailed in Aparicio et al. (2009). RT-PCR using primers designed from conserved regions of the orthotospovirus genome (Chu et al., 2001) and dot blot hybridization with digoxigenin-labelled probes were applied for the universal detection of orthotospovirus species (Eiras et al., 2001). The detection of TSWV in individual thrips by real time fluorescent RT-PCR using TaqMan chemistry has shown to be a very sensitive technique suitable for large-scale testing (Boonham et al., 2002; Debreczeni et al., 2011).

Particles of TSWV and other orthotospovirus species can be detected by electron microscopy using leaf dip preparations but it is necessary to prevent distortion of particles by prior fixation. Although the preparation of thin sections from tissues of infected plants is time-consuming, this method is very reliable for the identification of orthotospoviruses, because of the distinct shape and size of the particles, accumulation of particles within the cisternae of the endoplasmic reticulum and presence of typical viroplasms. Louro (1995) has devised a tissue-print immunoassay for TSWV. Detection should not rely solely on serological techniques in dubious cases. The serological results have to be confirmed by EM, inoculation to an indicator host or hosts, or molecular techniques, when working with hosts in which infections with TSWV and other orthotospoviruses are not well studied or with which the experimenter has no or limited experience.

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.

As there is no direct means of controlling the virus, the method of control must either be aimed at the thrips vectors or involve the application of sanitation measures. Seedling beds should be isolated from ornamental plants and susceptible crops and the surrounding areas kept free from weeds. The inside and outside of glasshouses should be kept free of weeds, thus reducing all possible sources of infection and reducing thrips populations. Fine-mesh netting may possibly be useful to exclude thrips (Lacasa et al., 1994). Susceptible decorative plants should preferably not be grown in the vicinity of the glasshouse. The glasshouse should be regularly and frequently inspected after planting. The presence of thrips in the crops should be monitored using yellow sticky cards. If the disease appears in a crop, infected plants should be rogued and destroyed immediately and the house treated with insecticide against thrips.

Similar precautions should be taken with field crops. Although chemical control is possible (Bournier, 1990), F. occidentalis has been found to develop resistant populations if certain insecticides are used repeatedly (OEPP/EPPO, 1989). It is, therefore, important to rotate insecticides with different active ingredients. For ornamental hosts (chrysanthemums, pelargoniums) for which virus-free certification schemes are applied, TSWV is now one of the most important viruses to be tested for (OEPP/EPPO, 1992). Promising results with biological and integrated control measures against thrips in glasshouses have been achieved in several countries (Gillespie, 1989; Ramakers et al., 1989; Trottin-Caudal and Grasselly, 1989; Sanchez et al., 2000). Frankliniella occidentalis, infesting plants and present in plastic houses, could be drastically reduced in number using UV-absorbing plastic sheets as cover (Antignus et al., 1996).

Reports about the virus-vector relationship and reduction in TSWV epidemics by chemical treatment are very limited. However, higher levels of control of TSWV were observed in dahlia with weekly sprays of mineral oil, polydimethylsiloxane and deltamethrin (Asjes and Blom-Barnhoorn, 2001). Host-plant resistance, intensive insecticide treatment and the use of reflective mulch significantly reduced the incidence of thrips and TSWV (Riley and Pappu, 2000). In Louisiana, USA, aluminium-surfaced mulch reduced the numbers of trapped thrips by 33-68% and the incidence of TSWV by 60-78% in tomatoes and Capsicum (Greenough et al., 1990). These results support the general view that primary transmission during adult thrips dispersal and host seeking accounted for most observed incidences of TSWV in tomato and groundnut (Camann et al., 1995) and probably also in most crops.

Yudin et al. (1990) devised disease-prediction and economic models that enable growers with lettuce fields affected by TSWV to make management decisions early in the planting cycle. Early disease incidence was a better predictor of disease incidence at harvest than thrips abundance because the proportion of infectious insects is essential to analyse the epidemics. The incidences of TSWV in early transplanted tomato crops with the highest thrips population was comparable to those found in late transplanted tomatoes with very low population densities (Aramburu et al., 1997).

The effect of the groundnut variety, planting date, plant density, insecticides used and disease history on the incidence and severity of TSWV was assessed in groundnut fields in Georgia, USA. These effects can be evaluated for each field in a risk assessment index. This index can vary between 25 and 125 points for each field. Fields with an index of 25-55 were considered to have a low risk, those between 60-80 a moderate risk, and those with more than 85 a high risk (Brown et al., 1998). Previous crops, adjoining crops, tillage practices, row patterns and weather also have effects, but were not indexed as these factors are insufficiently defined.

Screening vegetative dahlia propagation material for TSWV infections resulted in an almost healthy dahlia crop and seed tubers in the Netherlands (Schadewijk, 1996).

The growth regulators gibberellic acid, naphthalene acetic acid and chlormequat sprayed once on tomato plants either before or after TSWV inoculation inhibited the infectivity of the virus (Sapatnekar and Sawant, 2001).

Several sources of resistance to TSWV have been found in species of Solanum (Kumar et al., 1995; Cho et al., 1996). Two dominant and three recessive genes were responsible for resistance in S. pimpinellifolium and two tomato cultivars (Finlay, 1953). Introduction of these genes in tomato lines did not result in field resistance (Watterson et al., 1989). Lack of success in introducing this resistance into commercial tomato cultivars may be due to the existence of different TSWV strains or pathotypes. The tomato cv. Stevens, obtained from a cross between S. lycopersicum and S. peruvianum, has broad resistance to different TSWV isolates (van Zijl et al., 1986) and has been preferred by breeders for incorporating resistance into cultivated tomatoes. This resistance, introgressed in cv. Stevens, is conferred by a single dominant gene denoted Sw-5 with a 98.7% penetrance (Stevens et al., 1992) and also provided a high level of resistance to other members of the genus orthotospovirus, including Groundnut ringspot orthotospovirus (GRSV), Tomato chlorotic spot orthotospovirus (TCSV) and Groundnut bud necrosis orthotospovirus (GBNV). TSWV isolates, breaking this resistance, have been found in field crops in South Africa, Australia, USA (California and Hawaii) and Spain (Thompson and van Zijl, 1996; Latham and Jones, 1998; Aramburu and Marti, 2003; Batuman et al., 2017). New and selected accessions from Solanum species showed high resistance to TSWV and other viruses and seem to be of interest for enhancing the durability of the resistance to TSWV in commercial varieties (Roselló et al., 1999; Picó et al., 2002).

In lettuces, two cultivars (Tinto and Ancora) are reported to be resistant to TSWV in Hawaii, USA (O'Malley and Hartmann, 1989). This resistance was not confirmed in later studies. In groundnuts, breeding lines with a lower incidence of spotted wilt and lower disease severity ratings have been, or will be, released (Culbreath et al., 1996). Some field tolerance to GBNV occurs in Indian cultivars (Nigam et al., 1990) and could be explained by mature and tissue resistance (Buiel and Parlevliet, 1996). In tobacco, Nicotiana sanderae was immune and N. alata and N. langsdorffii were highly resistant to TSWV (Palakarcheva and Yancheva, 1989).

A gene designated Tsw, which prevents systemic spread of TSWV by a hypersensitive response has been identified in several C. chinense accessions (Black et al., 1993; Boiteux, 1995). This resistance proved to be less stable when young plants became infected and were kept at high temperatures (Roggero et al., 1996; Moury et al., 1998; Soler et al., 1998); by contrast, INSV infection was restricted to the inoculated leaves in Capsicum annuum and C. chinense under high temperatures (Roggero et al., 1999). The Tsw gene has phenotypic and genetic similarities of resistance in pepper with tomato plants carrying the Sw-5 gene; however, distinct viral gene products control the outcome of TSWV infection (Jahn et al., 2000); so, TSWV isolates that overcome tomato resistance gene Sw-5 failed to overcome hypersensitive resistance to TSWV in C. chinense PI 152225 and PI 159236 (Latham and Jones, 1998). Line 159236 was not resistant to GRSV (Boiteux and Nagata, 1993).

High levels of resistance to TSWV has been obtained in inbred lines of tomato transformed with the nucleoprotein (N) gene (de Haan et al., 1996). Similar levels of resistance have also been found in Nicotiana tabacum and N. benthamiana (Vaira et al., 1995), and chrysanthemum (Sherman et al., 1998) containing the nucleoprotein gene of TSWV. This transgenic resistance to TSWV in N. tabacum is effective in reducing the incidence of the disease under field conditions (Herrero et al., 2000). Sense or antisense copies of the N or Nsm genes can confer resistance. Groundnut lines transgenic for the antisense nucleocapsid (N) gene showed a lower TSWV incidence in field assays (Magbanua et al., 2000). Other TSWV sequences, spanning 70% of the genome, appear not to be effective in inducing resistance in transgenic tobacco (Prins et al., 1996). A broad resistance to GRSV, TSWV and TCSV was found in tobacco plants expressing the N gene sequences of these viruses (Prins et al., 1995). Transgenic plants expressing the transgene with green fluorescent protein fused to segments of the nucleocapsid (N) gene of TSWV showed multiple virus resistance (Jan et al., 2000).

The ability of TSWV isolates to overcome the resistance conferred by Sw-5 gene in tomato and the resistance conferred by the nucleocapsid gene in transgenic tobacco has been associated with the M RNA segment (Hoffmann et al., 2001).

Phytosanitary Measures

Susceptible host plants in greenhouses should be regularly inspected for orthotospovirus infections and vectors. Removal or roguing of infected plants, especially when the incidence is low, is an option to control further spread. Application of this practise may depend on the crop and its age, and the question whether the infection will or will not spread in the crop. Vectors should be actively controlled at the place of production. In general, heavily infected crops should be destroyed. Where appropriate, healthy planting material should be used. All plant residues left after harvested crops in greenhouses and fields should be eliminated. The soil has to be disinfected after the removal or harvest of severely infected crops with a high infestation of thrips. The emerging of viruliferous adults from infected pupae may form a serious threat to the new crop.

References

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Parrella G, Troiano E, Cherchi C, Giordano P, 2020. Severe outbreaks of parietaria mottle virus in tomato in Sardinia, southern Italy. Journal of Plant Pathology. 102 (3), 915-915. DOI:10.1007/s42161-020-00492-8

Pasev G, Radeva-Ivanova V, Lyall R, Nankar A, Kostova D, Turina M, Turina M, Vallino M, 2020. First report of Tomato spotted wilt virus on lisianthus (Eustoma grandiflorum) in Bulgaria. Journal of Plant Pathology. 103 (1), 375-375. DOI:10.1007/s42161-020-00702-3

Pérez-Colmenares Y, Mejías A, Rodríguez-Román E, Avilán D, Gómez J C, Marys E, Olachea J E, Zambrano K, 2015. Identification of Tomato spotted wilt virus associated with fruit damage during a recent virus outbreak in pepper in Venezuela. Plant Disease. 99 (6), 896. http://apsjournals.apsnet.org/loi/pdis

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Polizzi G, Bellardi M G, 2007. First report of Tomato spotted wilt virus on Coprosma repens (mirror bush) in Italy. Plant Disease. 91 (10), 1362. DOI:10.1094/PDIS-91-10-1362C

Pourrahim R, Golnaraghi A R, Farzadfar S, 2012. Occurrence of Impatiens necrotic spot virus and Tomato spotted wilt virus on potatoes in Iran. Plant Disease. 96 (5), 771. DOI:10.1094/PDIS-01-12-0051-PDN

Rabiee S, Hosseini S, Hosseini A, 2015. Occurrence and distribution of some sunflower viruses from sunflower fields in Kerman and Isfahan provinces, Iran. Archives of Phytopathology and Plant Protection. 48 (3), 223-228. DOI:10.1080/03235408.2014.884827

Renukadevi P, Nagendran K, Nakkeeran S, Karthikeyan G, Jawaharlal M, Alice D, Malathi V G, Pappu H R, 2015. First report of Tomato spotted wilt virus infection of chrysanthemum in India. Plant Disease. 99 (8), 1190. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-01-15-0126-PDN

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Salamon P, Nemes K, Salánki K, Palkovics L, 2012. First report of natural infection of pea (Pisum sativum) by Tomato spotted wilt virus in Hungary. Plant Disease. 96 (2), 295. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-06-11-0508

Salem N M, Mansour A, Badwan H, 2012. Identification and partial characterization of tomato spotted wilt virus on lettuce in Jordan. Journal of Plant Pathology. 94 (2), 431-435. http://sipav.org/main/jpp/index.php/jpp/article/view/2572

Senthilraja C, Renukadevi P, Malathi V G, Nakkeeran S, Pappu H R, 2018. Occurrence of tomato spotted wilt virus infecting snapdragon (Antirrhinum majus) in India. Plant Disease. 102 (8), 1676. DOI:10.1094/pdis-02-18-0250-pdn

Shahraeen N, Farzadfar S, Lesemann D E, 2003. Incidence of viruses infecting winter oilseed rape (Brassica napus ssp. oleifera) in Iran. Journal of Phytopathology. 151 (11/12), 614-616. DOI:10.1046/j.0931-1785.2003.00774.x

Shahraeen N, Ghotbi T, Mehraban A H, 2002. Occurrence of Impatiens necrotic spot virus in ornamentals in Mahallat and Tehran provinces in Iran. Plant Disease. 86 (6), 694. DOI:10.1094/PDIS.2002.86.6.694A

Sialer M M F, Gallitelli D, 2000. The occurrence of Impatiens necrotic spot virus and Tomato spotted wilt virus in mixed infection in tomato. Journal of Plant Pathology. 82 (3), 244.

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Sivaprasad Y, Garrido P, Mendez K, Pachacama S, Garrido A, Ramos L, 2017. First report of Tomato spotted wilt virus infecting pepper in Ecuador. Journal of Plant Pathology. 99 (1), 304. http://www.sipav.org/main/jpp/index.php/jpp/article/view/3837/2479

Sivaprasad Y, Garrido P, Mendez K, Pachacama S, Garrido A, Ramos L, 2018. First report of Tomato spotted wilt virus infecting chrysanthemum in Ecuador. Journal of Plant Pathology. 100 (1), 113-113. DOI:10.1007/s42161-018-0010-5

Soler S, López C, Nuez F, 2005. Natural occurrence of viruses in Lycopersicon spp. in Ecuador. Plant Disease. 89 (11), 1244. HTTP://www.apsnet.org DOI:10.1094/PD-89-1244C

Soler S, Prohens J, López C, Aramburu J, Galipienso L, Nuez F, 2010. Viruses infecting tomato in València, Spain: occurrence, distribution and effect of seed origin. Journal of Phytopathology. 158 (11/12), 797-805. DOI:10.1111/j.1439-0434.2010.01706.x

Spanò R, Mascia T, Lucia B de, Torchetti E M, Rubino L, Gallitelli D, 2011a. First report of a resistancebreaking strain of Tomato spotted wilt virus from Gerbera jamesonii in Apulia, southern Italy. Journal of Plant Pathology. 93 (4, Supplement), S4.63. http://sipav.org/main/jpp/index.php/jpp/issue/view/118

Spanò R, Mascia T, Minutillo S A, Gallitelli D, 2011. First report of Tomato infectious chlorosis virus from tomato in Apulia, southern Italy. Journal of Plant Pathology. 93 (4, Supplement), S4.64. http://sipav.org/main/jpp/index.php/jpp/issue/view/118

Stanković I, Bulajić A, Vučurović A, Ristić D, Jović J, Krstić B, 2011. First report of Tomato spotted wilt virus on Gerbera hybrida in Serbia. Plant Disease. 95 (2), 226. DOI:10.1094/PDIS-10-10-0704

Stanković I, Bulajić A, Vučurović A, Ristić D, Milojević K, Nikolić D, Krstić B, 2012. First report of Tomato spotted wilt virus infecting onion and garlic in Serbia. Plant Disease. 96 (6), 918. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-02-12-0157-PDN

Stanković I, Bulajić A, Vučurović A, Ristić D, Milojević K, Nikolić D, Krstić B, 2013. First report of Tomato spotted wilt virus on chrysanthemum in Serbia. Plant Disease. 97 (1), 150-151. DOI:10.1094/PDIS-08-12-0778-PDN

Steyer S, Olivier T, Skelton A, Nixon T, Hobden E, 2010. Columnea latent viroid (CLVd): first report in tomato in France. Plant Pathology. 59 (4), 794. DOI:10.1111/j.1365-3059.2010.02261.x

Sui X, McGrath M T, Zhang S, Wu Z, Ling K S, 2018. First report of Tomato chlorotic spot virus infecting Tomato in New York. Plant Disease. 102 (2), 460. DOI:10.1094/PDIS-07-17-0991-PDN

Sun X H, Gao L L, Wang S L, Wang C L, Yang Y Y, Wang X Y, Zhu X P, 2016. First report of Tomato spotted wilt virus infecting pumpkin in China. Journal of Plant Pathology. 98 (3), 687. http://www.sipav.org/main/jpp/index.php/jpp/article/view/3748/2389

Sundaraj S, Srinivasan R, Webster C G, Adkins S, Perry K, Riley D, 2011. First report of Tomato chlorosis virus infecting tomato in Georgia. Plant Disease. 95 (7), 881. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-02-11-0122

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Thomas J E, Schwinghamer M W, Parry J N, Sharman M, Schilg M A, Dann E K, 2004. First report of Tomato spotted wilt virus in chickpea (Cicer arietinum) in Australia. Australasian Plant Pathology. 33 (4), 597-599. DOI:10.1071/AP04065

Trkulja V, Salapura J M, Ćurković B, Stanković I, Bulajić A, Vučurović A, Krstić B, 2013. First report of Tomato spotted wilt virus on gloxinia in Bosnia and Herzegovina. Plant Disease. 97 (3), 429. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-08-12-0777-PDN

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Vilchez M, Paulus A O, Moyer J W, Schrader W L, 2005. First report of tomato spotted wilt virus affecting globe artichoke in California, USA. Acta Horticulturae. 607-610. http://www.actahort.org

Wan Y R, Yin Y Q, Wu Q J, 2017. First report of tomato spotted wilt virus infecting balloon flower in China. Journal of Plant Pathology. 99 (3), 814. http://www.sipav.org/main/jpp/index.php/jpp/article/view/3978/2622

Wangai A W, Mandal B, Pappu H R, Kilonzo S, 2001. Outbreak of Tomato spotted wilt virus in tomato in Kenya. Plant Disease. 85 (10), 1123. DOI:10.1094/PDIS.2001.85.10.1123B

Webster C G, Turechek W W, Mellinger H C, Frantz G, Roe N, Yonce H, Vallad G E, Adkins S, 2011. Expansion of Groundnut ringspot virus host and geographic ranges in solanaceous vegetables in peninsular Florida. Plant Health Progress. PHP-2011-0725-01-BR. http://www.plantmanagementnetwork.org/php/elements/sum2.aspx?id=9490

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Xiao L, Li Y Y, Lan P X, Tan G L, Ding M, Li R H, Li F, 2016. First report of Tomato spotted wilt virus infecting cowpea in China. Plant Disease. 100 (1), 233. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-04-15-0495-PDN

Yeturu S, Viera W, Garrido P, Insuasti M, 2016. First report of Tomato spotted wilt virus infecting tree tomato (Solanum betaceum Cav.) in Ecuador. Journal of Plant Pathology. 98 (3), 691. http://www.sipav.org/main/jpp/index.php/jpp/article/view/3753/2394

Yoon J Y, Choi G S, Choi S K, 2017. First report of Tomato spotted wilt virus in Eustoma grandiflorum in Korea. Plant Disease. 101 (3), 515. DOI:10.1094/pdis-10-16-1514-pdn

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Yoon Y N, Jo Y, Cho W K, Choi H, Jang Y, Lee Y H, Bae J Y, Lee B C, 2018. First report of Tomato spotted wilt virus infecting soybean in Korea. Plant Disease. 102 (2), 461-462. DOI:10.1094/PDIS-07-17-1051-PDN

Yu M C, Yang C X, Wang J Z, Hou Q S, Zhang S, Cao M J, 2021. First report of Tomato spotted wilt virus isolated from Nasturtium (Tropaeolum majus) with a serious leaf mosaic disease in China. Plant Disease. 105 (3), 716-716. DOI:10.1094/PDIS-03-20-0688-PDN

Zarzyńska-Nowak A, Rymelska N, Borodynko N, Hasiów-Jaroszewska B, 2016. The occurrence of Tomato yellow ring virus on tomato in Poland. Plant Disease. 100 (1), 234. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-05-15-0521-PDN

Zheng Y X, Huang C H, Cheng Y H, Kuo F Y, Jan F J, 2010. First report of Tomato spotted wilt virus in sweet pepper in Taiwan. Plant Disease. 94 (7), 920. DOI:10.1094/PDIS-94-7-0920B

Zindović J, Bulajić A, Krstić B, Ciuffo M, Margaria P, Turina M, 2011. First report of Tomato spotted wilt virus on pepper in Montenegro. Plant Disease. 95 (7), 882. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-03-11-0167

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19/12/20 Updated by:

Dirk Janssen, Instituto de Investigación y Formación Agraria y Pesquera (IFAPA), Seville, Spain

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