Squash leaf curl virus (leaf curl of squash)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Means of Movement and Dispersal
- Plant Trade
- Vectors and Intermediate Hosts
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Squash leaf curl virus
Preferred Common Name
- leaf curl of squash
Other Scientific Names
- Melon leaf curl virus
- Squash leaf curl
- Watermelon curly mottle virus
International Common Names
- English: curly mottle of watermelon; leaf curl of melon; necrotic mosaic of melon
- SLCV00 (Squash leaf curl begomovirus)
Taxonomic TreeTop of page
- Domain: Virus
- Unknown: "ssDNA viruses"
- Unknown: "DNA viruses"
- Family: Geminiviridae
- Genus: Begomovirus
- Species: Squash leaf curl virus
Notes on Taxonomy and NomenclatureTop of page
The original strain of the virus (SLCV-CA) was reported in California, USA, damaging crops of squash marrows or summer squashes (Cucurbita pepo) and winter squashes (C. moschata) (Flock and Mayhew, 1981). The Watermelon curly mottle virus strain (WCMoV) reported in Arizona, USA, has a wider host range, also attacking buffalo gourd (Cucurbita foetidissima), cucumber (Cucumis sativus), muskmelon and honeydew melon (Cucumis melo) and watermelon (Citrullus lanatus) (Brown and Nelson, 1986, 1989; Rosemeyer et al., 1986). Other hosts include pumpkin (C. maxima), Phaseolus vulgaris and several wild species of Cucurbita. SLCV is nearly restricted to Cucurbitaceae, which are the only hosts on which it is of economic importance. Certain species have been reported to be resistant in glasshouse and field tests including Benincasa, Lagenaria, Luffa (McCreight, 1984; McCreight and Kishaba, 1991). P. vulgaris and Nicotiana benthamiana are the only reported non-cucurbitaceous hosts of WCMoV (Brown, 1986, 1989) and SLCV-E (Lazarowitz and Lazdins, 1991) and both species are only known as experimental hosts.
Melon leaf curl virus (Duffus et al., 1985) is probably a synonym of WCMoV in that their host ranges and symptomatology are indistinguishable, and the MLCV isolate was described in melon in California, USA, after WCMoV was reported in nearby melon fields in Arizona, USA.
The complete genome sequence for the SLCV-Extended host range isolate (SLCV-E) has been published but the corresponding biological properties are still unresolved, as are the relationships between SLCV-CA, SLCV-E, SLCV-R and WCMoV (syn. MLCV). Both SLCV-E and SLCV-R (partial sequence published) occurred in a mixture in the squash plant from which they were cloned (Lazarowitz, 1991). WCMoV (SLCV-E like?) and SLCV (SLCV-R-like?) are known to exist individually and in mixtures of three or four viral components in the same host species (Polston et al., 1989; Brown et al., 1995). Currently, at least two distinct begomovirus species are thought to be involved in this disease complex in the region, SLCV (R-like) and WCMoV (like SLCV-E).
DescriptionTop of page SLCV has geminate particles, 22 x 38 nm (Cohen et al., 1983). The virus is associated with maturing phloem sieve tube elements (Hoefert, 1987). Virions contain one circular, single-stranded DNA molecule. Bipartite viruses such as SLCV require two different chromosomes, packaged separately, while monopartite begomoviruses have a single chromosome. All New World begomoviruses have a bipartite arrangement, while Old World viruses can have either.
DistributionTop of page
WCMoV (syn? SLCV-E) and SLCV-CA (syn? SLCV-R) occur in agroecosystems in the south-western USA and northern Mexico including Arizona, California and Texas (USA) and Sonora (Mexico).
In 1999-2000, a newly described begomovirus, provisionally called Cucurbit leaf curl virus (CLCuV), was discovered in several cucurbit species from Arizona, California and Texas (USA) and Coahuilla (Mexico). During these growing seasons in Arizona and Texas, CLCuV was the most common begomovirus identified in field-infected melon, pumpkin and squash fields (Brown et al., 2000). SLCV-E-like isolates were still predominant in Hermosillo and Mayo Valley in Sonora, Mexico (JK Brown et al., unpublished data). CLCuV was most recently identified in field-infected cucumber from Coahuilla, Mexico and in watermelon from Arizona, USA.
Begomovirus-like symptoms have been observed in cucurbits (squashes and melons) in Costa Rica, Dominican Republic, Guatemala, Honduras and Nicaragua (JK Brown, pers comm) but only recently has the identity of one of the isolates been determined. In spring 2000, a newly described begomovirus species was identified in melon from Guatemala. The virus which is under more detailed investigation, has been provisionally named Melon chlorotic leaf curl virus (MCLCV) (Brown et al., 2001).
In the Philippines, Benigno (1977) and Dolores et al. (1988) refer to a Bemisia-transmitted leaf curl of squash, said to be seedborne and associated with Bemisia tabaci infestations. This reference predates the description of SLCV in the USA and is unlikely to refer to the same virus. A begomovirus of squash was also reported from China, SLCV-China (SLCV-CH) (Genbank Accession # AB027465) (Qinyan et al., 1999), and appears to be of Old World origin based on DNA sequence analysis; consequently it is a begomovirus species distinct from SLCV-like isolates from the USA and Mexico.
The distantly related, Old World Watermelon chlorotic stunt virus (WmCSV) has been described in watermelon in Yemen (Bedford et al., 1994) and Iran (Kheyr-Pour et al., 2000; Genbank Accessions # AJ245652 and AJ245653) and Sudan (Kheyr-Pour et al., 2000; Genbank Accessions # AJ245650 and AJ245651).
Distribution TableTop of page
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Israel||Present||CABI/EPPO, 2014; EPPO, 2014|
|Jordan||Present||Al-Musa et al., 2008; CABI/EPPO, 2014; EPPO, 2014|
|Lebanon||Present||EPPO, 2012; Sobh et al., 2012; CABI/EPPO, 2014; EPPO, 2014|
|Saudi Arabia||Present||CABI/EPPO, 2014; EPPO, 2014|
|Taiwan||Present||Tsai et al., 2007|
|Egypt||Present||CABI/EPPO, 2014; EPPO, 2014|
|Mexico||Restricted distribution||Montes Garcia et al., 1998; Ramirez-Arredondo et al., 1995; CABI/EPPO, 2014; EPPO, 2014|
|USA||Restricted distribution||CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014|
|-Arizona||Present||Brown and Nelson, 1986; Rosemeyer et al., 1986; Brown and Nelson, 1989; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014|
|-California||Present||Flock and Mayhew, 1981; Cohen et al., 1983; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014|
|-Texas||Present||1993||Isakeit et al., 1994; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014|
Central America and Caribbean
|Costa Rica||Present||CABI/EPPO and, 1997; EPPO, 2014|
|Dominican Republic||Present||CABI/EPPO and, 1997; EPPO, 2014|
|Guatemala||Present||CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014|
|Honduras||Present||CABI/EPPO and, 1997; EPPO, 2014|
|Nicaragua||Present||CABI/EPPO and, 1997; EPPO, 2014|
|Belgium||Absent, no pest record||EPPO, 2014|
Risk of IntroductionTop of page
SLCV is on the EPPO A2 list, but has not been classified as a quarantine pest by any other regional plant protection organisation. This is no doubt partly due to the fact that its appearance as a significant pest is very recent. If introduced outside of its native habitat, it presents a severe threat to the cultivation of cucurbits (especially courgettes, melons and watermelons) in the open in southern Europe or under glass in northern Europe, wherever B. tabaci and especially the B biotype, occurs.
Host plants of B. tabaci from areas where SLCV occurs should come from a production nursery free from SLCV and B. tabaci (or treated against B. tabaci) during the last growing season. This applies especially to the ornamental Euphorbia pulcherrima or other ornamental species that are well known to carry B. tabaci inconspicuously.
Host Plants and Other Plants AffectedTop of page
|Citrullus lanatus (watermelon)||Cucurbitaceae||Main|
|Cucumis melo (melon)||Cucurbitaceae||Main|
|Cucumis sativus (cucumber)||Cucurbitaceae||Main|
|Cucurbita maxima (giant pumpkin)||Cucurbitaceae||Main|
|Cucurbita moschata (pumpkin)||Cucurbitaceae||Main|
|Cucurbita pepo (marrow)||Cucurbitaceae||Main|
|Phaseolus vulgaris (common bean)||Fabaceae||Main|
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page
SLCV causes severe systemic stunting and leaf curling in most cucurbits. In pumpkin, squash and watermelon, a severe, chlorotic foliar mosaic or mottle accompanies leaf curling and stunting symptoms. Symptoms in Cucumis species caused by WCMoV are confined to leaf curling at the growing tips and often disappear as the plants mature and begin to flower and set fruit. Symptoms on fruit are not particularly notable, perhaps because very little, if any, fruit is set on most cultivars after infection occurs in susceptible cultivars.
The so-called silverleaf symptom seen on cucurbit plants infested by Bemisia tabaci (B biotype) (Costa and Brown, 1991) is not caused by SLCV.
List of Symptoms/SignsTop of page
|Growing point / distortion|
|Leaves / abnormal colours|
|Leaves / abnormal forms|
|Whole plant / distortion; rosetting|
|Whole plant / dwarfing|
Biology and EcologyTop of page
The SLCV complex is transmitted in a persistent manner by the whitefly, Bemisia tabaci. Both B. tabaci and B. tabaci (B biotype) are effective vectors in the USA and Mexico. Virus can be acquired in ca 30 minutes to several hours. A latent period of ca 6-10 hours is required before transmission can occur. Whiteflies can continue to transmit the virus for 7-10 days or perhaps for the life of the vector. There is no evidence for transovarial transmission of SLCV (Cohen et al., 1983; Brown et al., 1986; Rosell et al., 1999).
SLCV can be present in spring, summer, autumn and winter crops if B. tabaci population levels are sufficiently high for dispersal of the virus. B. tabaci is typically abundant in the late spring, summer and autumn in the southern river valleys of Arizona and California, and in summer and fall in the Rio Grande Valley, Texas, USA (watermelon, melons, squash) and northern Coahuilla, Mexico (watermelon, cucumber, squash, melons). The vector is most abundant in squash, melons and cucumbers during the autumn, winter and spring in Sonora, Mexico. Susceptible cultivars develop symptoms within 6-12 days post-infection and can serve as virus reservoirs from which secondary spread can readily occur. Spread between crops is the most important means by which virus causes epidemics or routine crop loss.
Several wild cucurbit species are known to be experimentally susceptible to SLCV infection but only one wild cucurbit species (in Texas) has been found naturally infected with the virus (T Isakeit and JK Brown, unpublished data). No natural reservoirs in which SLCV overseasons have been identified in the USA or Mexico.
Means of Movement and DispersalTop of page
SLCV is transmitted by Bemisia tabaci and B. tabaci (B biotype), and probably other biotypes that can feed on cucurbits; all in a persistent, circulative manner.
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Flowers/Inflorescences/Cones/Calyx||Yes||Pest or symptoms usually invisible|
|Fruits (inc. pods)||Yes||Pest or symptoms usually invisible|
|Leaves||Yes||Pest or symptoms usually visible to the naked eye|
|Stems (above ground)/Shoots/Trunks/Branches||Yes||Pest or symptoms usually visible to the naked eye|
|Plant parts not known to carry the pest in trade/transport|
|Growing medium accompanying plants|
|True seeds (inc. grain)|
Vectors and Intermediate HostsTop of page
ImpactTop of page
SLCV causes severe losses of all squash and pumpkin cultivars, and minor losses in cucumbers and melons in Arizona, California (Dodds et al., 1985; Brown and Nelson, 1986) and Texas, USA (Isakeit et al., 1994) and in Coahuilla and Sonora, Mexico (JK Brown, unpublished data; Ramirez-Arredondo et al., 1995). Although recorded to infect Phaseolus vulgaris, there is no indication that it has any importance on that host naturally (Brown and Nelson, 1986, 1989). The leaf curl disease was first observed in 1977/78 (Flock and Mayhew, 1981) and on cultivated buffalo gourd in Arizona in 1976/77 (Rosemeyer et al., 1986) and is caused by one (or more) of a group of viruses which were transmitted by B. tabaci (native to the deserts of the south-western USA and Sonora, Mexico) and has become even more important since the introduction and spread of B. tabaci (B biotype) in about 1987/88 (Costa and Brown, 1991).
DiagnosisTop of page
SLCV is not readily mechanically transmissible but can be transmitted by sap inoculation with difficulty. Severe mottle/mosaic and leaf curl develop on leaves that develop shortly after infection, and fruit is usually without symptoms (Brown and Nelson, 1986, 1989). Recommended indicator plant species are Cucurbita maxima, C. moschata or C. pepo (symptoms as above), and Cucumis melo, the latter to separate SLCV-CA from WCMoV. SLCV can be detected by PCR using universal primers for the core region of the coat protein gene (core CP) (Wyatt and Brown, 1996) followed by nucleotide (nt) sequencing of the cloned amplicon, and comparison with sequences of well-studied begomoviruses of cucurbits. If necessary, virus-specific PCR primers can be designed by taking into account the nt sequence of different SLCV strains (or species) in relation to each other and other closely related cucurbit-infecting begomoviruses. SLCV can also be detected in B. tabaci by DNA spot hybridization assay (Polston et al., 1990) or by PCR (Rosell et al., 1999) but unless component specific probes are employed or the nt sequence of the PCR amplicon is determined, respectively, accurate identification is not possible because several other closely related viruses may also be detected using these approaches.
Detection and InspectionTop of page
SLCV is not readily mechanically transmissible but can be so transmitted with difficulty. Symptoms are recognisable as severe mottle/mosaic and leaf curl on newer leaves and fruit is usually without symptoms. Often, adult or immature B. tabaci whiteflies are present on leaves of infected plants (Brown and Nelson, 1986, 1989). Whitefly-mediated transmission to recommended indicator plants (Cucurbita maxima or C. moschata or C. pepo) is needed to separate SLCV virus from non-whitefly transmitted viruses that may occur in mixed infection.
Similarities to Other Species/ConditionsTop of page
It may be stressed that the SLCV complex, like Lettuce infectious yellows virus (EPPO/CABI, 1996), has only been recorded in North America. Several other B. tabaci-transmitted viruses are now known to attack cucurbits in other parts of the world, including the little-known Cucumber vein yellowing virus in the Middle East (Yilmaz et al., 1989; Mansour and Al Musa, 1993), which are not begomoviruses and have entirely different characteristics. The newly described begomoviruses in the USA and Mexico, CLCuV and MCLCV, may be confused with SLCV in certain hosts but in general they cause distinctive symptoms in their hosts. CLCuV causes chlorotic sectoring and leaf curling on Cucurbita and Citrullus species and a chlorotic mosaic/mottle on Cucumis species in contrast to SLCV-infected Cucurbita species which exhibit severe leaf curling and a striking mottling and distortion as well as stunting. MCLCV causes interveinal chlorosis, leaf curling and stunting in melon, compared with SLCV which causes a mild curling on new growth which in most cultivars disappears as plants mature, even though the virus remains detectable in infected plants.
Prevention and ControlTop of page
SLCV is spread naturally only by its vector Bemisia tabaci which spreads it between fields (and presumably glasshouses) in infested areas. In international trade, it is unlikely to be carried by plants of its main cultivated hosts, since these are short-lived vegetable crops not normally transported except as seedlings. Young seedlings for transplanting might constitute a pathway, but would still be unlikely to be moved in intercontinental trade. The vegetables as such would not be likely to carry B. tabaci. So the main risk of movement is in B. tabaci on other host plants (for example, ornamentals), given the fact that the vector moves readily from one host to another and that the virus can persist in the vector for several weeks (to life) after virus acquisition (Cohen et al., 1983; Rosell et al., 1999).
In the field, disease control is mainly aimed at eliminating or excluding the whitefly vector, B. tabaci. Insecticides have been used in an attempt to reduce whitefly density. Protecting rows of seedlings with spun-bonded polyester as a floating cover has also been effective (Natwick and Durazo, 1985). Some cultivars of Cucurbita pepo have shown resistance to SLCV (McCreight, 1984, 1991).
ReferencesTop of page
Ali-Shtayeh MS; Jamous RM; Husein EY; Alkhader MY, 2010. First report of Squash leaf curl virus in squash (Cucurbita pepo), melon (Cucumis melo), and cucumber (Cucumis sativa) in the Northern West Bank of the Palestinian Authority. Plant Disease, 94(5):640. http://apsjournals.apsnet.org/loi/pdis
Al-Musa A; Anfoka G; Misbeh S; Abhary M; Ahmad FH, 2008. Detection and molecular characterization of Squash leaf curl virus (SLCV) in Jordan. Journal of Phytopathology, 156(5):311-316. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1439-0434.2007.01389.x
Bedford ID; Briddon RW; Brown JK; Rosell RC; Markham PG, 1994. Geminivirus transmission and biological characterisation of Bemisia tabaci (Gennadius) biotypes from different geographic regions. Annals of Applied Biology, 125(2):311-325.
Bedford ID; Briddon RW; Jones P; Alkaff N; Markham PG, 1994. Differentiation of three whitefly-transmitted geminiviruses from the Republic of Yemen. European Journal of Plant Pathology, 100(3-4):243-257.
Brown JK; Idris AM; Rogan D; Hussein MH; Palmieri M, 2001. Melon chlorotic leaf curl virus, a new whitefly-transmitted geminivirus in Guatemala. Plant Disease, 85: (submitted).
Brown JK; Ostrow KM; Idris AM; Stenger DC, 1999. Biotic, molecular, and phylogenetic characterization of bean calico mosaic virus, a distinct Begomovirus species with affiliation in the squash leaf curl virus cluster. Phytopathology, 89(4):273-280; 58 ref.
Brown JK; Wendt K; Wyatt SD, 1995. Genetic variability of Squash leaf curl virus isolates by DNA hybridization and component-specific polymerase chain reaction. In: Proceedings of Cucurbitaceae '94. Lester G, Dunlap J, et al., eds. USA: Gateway Printing, 5-11.
Costa HS; Brown JK, 1991. Variation in biological characteristics and esterase patterns among populations of Bemisia tabaci, and the association of one population with silverleaf symptom induction. Entomologia Experimentalis et Applicata, 61(3):211-219.
Dolores LM; Valdez RB; Roxas VP, 1988. Viral diseases of squash (Cucurbita spp.) in the Philippines and sources of resistance. Philippine Journal of Crop Science, 13(Supplement 1):S30; [in ^italic~4th scientific meeting of the Federation of Crop Science Societies of the Philippines held in Apo View Hotel, Davao City, April 27-30, 1988^roman~].
Duffus JE; Liu HY; Johnson MR, 1985. Melon leaf curl virus-a new geminivirus with host and serological variations from Squash leaf curl virus. Phytopathology, 75:1312.
EFSA Panel on Plant Health, 2013. Scientific Opinion on the risks to plant health posed by Bemisia tabaci species complex and viruses it transmits for the EU territory. EFSA Journal, 11(4). 3162. http://www.efsa.europa.eu/sites/default/files/scientific_output/files/main_documents/3162.pdf
EPPO, 2012. 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
Lazarowitz SG, 1987. The molecular characterization of geminiviruses. Plant Molecular Biology Reporter, 4(4):177-192.
Lazarowitz SG, 1991. Molecular characterization of two bipartite geminiviruses causing squash leaf curl disease: role of viral replication and movement functions in determining host range. Virology (New York), 180(1):70-80; 25 ref.
Lazarowitz SG; Lazdins IB, 1991. Infectivity and complete nucleotide sequence of the cloned genomic components of bipartite squash leaf curl geminivirus with a broad host range phenotype. Virology (New York), 180(1):58-69; 47 ref.
McCreight JD, 1984. Tolerance of Cucurbita spp. to squash leaf curl. Report of Cucurbit Genetics Co-operative, USA, 7:71-72.
Montes Garcia C; Garza Ortega S; Brown JK, 1998. Inheritance of the resistance to Squash leaf curl virus in Cucurbita pepo (L.). Proceedings of Cucurbitaceae 98. Evaluation and Enhancement of Cucurbit Germplasm, Asilomar Conference Center, Pacific Grove, CA, November 30-December 4, 1998, 328-330.
Polston JE; Dodds JA; Perring TM, 1989. Nucleic acid probes for detection and strain discrimination of cucurbit geminiviruses. Phytopathology, 79:1123-1127.
Ramfrez Arredondo JA; ArmentaCßrdens I; Delgadillo Sßnchez F; Rivera-Bustamante RF; Garz=n-Tiznado JA, 1995. Geminiviruses transmitted by whitefly (Bemisia tabaci Genn.) in crops of pepper and squash in Mayo Valley, Sonora, Mexico. Revista Mexicana de Fitopatologi^acute~a, 13(2):100-105; 18 ref.
Rosell RC; Torres-Jerez I; Brown JK, 1999. Tracing the geminivirus-whitefly transmission pathway by polymerase chain reaction in whitefly extracts, saliva, haemolymph, and honeydew. Phytopathology, 89(3):239-246; 46 ref.
Smith IM; McNamara DG; Scott PR; Holderness M, 1997. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization., Ed. 2:vii + 1425 pp.; many ref.
Sobh H; Samsatly J; Jawhari M; Najjar C; Haidar A; Abou-Jawdah Y, 2012. First report of Squash leaf curl virus in cucurbits in Lebanon. Plant Disease, 96(8):1231. http://apsjournals.apsnet.org/loi/pdis
Distribution MapsTop of page
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