Squash leaf curl virus
(leaf curl of squash)

Pictures

Picture	Title	Caption	Copyright
Title Symptoms Caption Squash leaf curl virus (leaf curl of squash); symptoms of SLCV CA in winter squash. Copyright ©Judith K. Brown	Symptoms	Squash leaf curl virus (leaf curl of squash); symptoms of SLCV CA in winter squash.	©Judith K. Brown
Title Symptoms Caption Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in watermelon. Copyright ©Judith K. Brown	Symptoms	Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in watermelon.	©Judith K. Brown
Title Symptoms Caption Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in zucchini squash. Copyright ©Judith K. Brown	Symptoms	Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in zucchini squash.	©Judith K. Brown
Title Symptoms Caption Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in melon. Copyright ©Judith K. Brown	Symptoms	Squash leaf curl virus (leaf curl of squash); symptoms of WCMoV in melon.	©Judith K. Brown

Identity

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

English acronym

• SLCV

EPPO code

• SLCV00 (Squash leaf curl begomovirus)

Taxonomic Tree

• Domain: Virus
•     Group: "ssDNA viruses"
•         Group: "DNA viruses"
•             Family: Geminiviridae
•                 Genus: Begomovirus
•                     Species: Squash leaf curl virus

Notes on Taxonomy and Nomenclature

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).

Description

Distribution

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 Table

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.

Risk of Introduction

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 Affected

Growth Stages

Symptoms

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/Signs

Biology and Ecology

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 Dispersal

Vector transmission

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 Trade

Vectors and Intermediate Hosts

Impact

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).
 

Diagnosis

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 Inspection

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/Conditions

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 Control

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.

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).

References

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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

Benigno DRA, 1977. Note: leaf curl disease of squash. Philippine Agriculturist, 61(7/8):304-305

Bharathan N, Narayanan KR, McMillan RT Jr, 1992. Characteristics of sweetpotato whitefly-mediated silverleaf syndrome and associated double-stranded RNA in squash. Phytopathology, 82(2):136-141

Brown JK, 1991. An update on the whitefly-transmitted geminiviruses in the Americas and the Caribbean Basin. FAO plant Protection Bulletin, 39(1):5-23

Brown JK, 1994. Current status of Bemisia tabaci as a plant pest and virus vector in agroecosystems worldwide. FAO Plant Protection Bulletin, 42(1/2):3-32

Brown JK, Bird J, 1992. Whitefly-transmitted geminiviruses and associated disorders in the Americas and the Caribbean Basin. Plant Disease, 76(3):220-225

Brown JK, Idris AM, Olsen MW, Miller ME, Isakeit T, Anciso J, 2000. Cucurbit leaf curl virus, a new whitefly transmitted geminivirus in Arizona, Texas, and Mexico. Plant Disease, 84(7):809; 3 ref

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, Nelson MR, 1986. Whitefly-borne viruses of melons and lettuce in Arizona. Phytopathology, 76(2):236-239

Brown JK, Nelson MR, 1989. Characterisation of watermelon curly mottle virus, a geminivirus distinct from squash leaf curl virus. Annals of Applied Biology, 115(2):243-252

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

CABI/EPPO, 2014. Squash leaf curl virus. [Distribution map]. Distribution Maps of Plant Diseases, No.October. Wallingford, UK: CABI, Map 748 (Edition 2)

Cohen S, Duffus JE, Larsen RC, Liu HY, Flock RA, 1983. Purification, serology, and vector relationships of squash leaf curl virus, a whitefly-transmitted geminivirus. Phytopathology, 73(12):1669-1673

Cohen S, Duffus JE, Liu HY, 1989. Acquisition, interference, and retention of cucurbit leaf curl viruses in whiteflies. Phytopathology, 79(1):109-113

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

Dodds JA, Lee JG, Nameth ST, Lpmmlen FF, 1984. Aphid- and whitefly-transmitted cucurbit viruses in Imperial County, California. Phytopathology, 74(2):221-225

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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

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Hoefert LL, 1987. Association of squash leaf curl virus with nuclei of squash vascular cells. Phytopathology, 77(11):1596-1600; 15 ref

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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

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McCreight JD, Kishaba AN, 1991. Reaction of cucurbit species to squash leaf curl virus and sweetpotato whitefly. Journal of the American Society for Horticultural Science, 116(1):137-141

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Natwick ET, Durazo A III, 1985. Polyester covers protect vegetables from whiteflies and virus diseases. California Agriculture, 39(7/8):21-22

Polston JE, Al-Musa A, Perring TM, Dodds JA, 1990. Association of the nucleic acid of squash leaf curl geminivirus with the whitefly Bemisia tabaci. Phytopathology, 80(9):850-856

Polston JE, Dodds JA, Perring TM, 1989. Nucleic acid probes for detection and strain discrimination of cucurbit geminiviruses. Phytopathology, 79:1123-1127

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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

Tsai WS, Shih L, Green SK, Jan FJ, 2007. Occurrence and molecular characterization of Squash leaf curl Philippines virus in Taiwan. Plant Disease, 91(7):907. HTTP://www.apsnet.org

Wyatt SD, Brown JK, 1996. Detection of subgroup III geminivirus isolates in leaf extracts by degenerate primers and polymerase chain reaction. Phytopathology, 86(12):1288-1293; 36 ref

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Distribution References

Al-Kuwaiti N A, 2017. Molecular characterization of a leaf curl disease infecting zucchini squash in Iraq. International Journal of Agriculture and Biology. 19 (4), 649-653.

Al-Musa A, Anfoka G, Misbeh S, Abhary M, Ahmad F H, 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 DOI:10.1111/j.1439-0434.2007.01389.x

Anon, 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. [ed. by Smith I M, McNamara D G, Scott P R, Holderness M]. Wallingford, UK: CAB INTERNATIONAL. vii + 1425 pp.

Brown J K, Nelson M R, 1986. Whitefly-borne viruses of melons and lettuce in Arizona. Phytopathology. 76 (2), 236-239. DOI:10.1094/Phyto-76-236

Brown J K, Nelson M R, 1989. Characterisation of watermelon curly mottle virus, a geminivirus distinct from squash leaf curl virus. Annals of Applied Biology. 115 (2), 243-252. DOI:10.1111/j.1744-7348.1989.tb03383.x

CABI, EPPO, 2014. Squash leaf curl virus. [Distribution map]. In: Distribution Maps of Plant Diseases, Wallingford, UK: CABI. Map 748 (Edition 2).

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Cohen S, Duffus J E, Larsen R C, Liu H Y, Flock R A, 1983. Purification, serology, and vector relationships of squash leaf curl virus, a whitefly-transmitted geminivirus. Phytopathology. 73 (12), 1669-1673. DOI:10.1094/Phyto-73-1669

EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO.

Flock R A, Mayhew D E, 1981. Squash leaf curl, a new disease of cucurbits in California. Plant Disease. 65 (1), 75-76. DOI:10.1094/PD-65-75

Isakeit T, Robertson N L, Brown J K, Gilbertson R L, 1994. First report of squash leaf curl virus on watermelon in Texas. Plant Disease. 78 (10), 1010. DOI:10.1094/PD-78-1010D

Rosemeyer M E, Brown J K, Nelson M R, 1986. Five viruses isolated from field-grown buffalo gourd (Cucurbita foetidissima), a potential crop for semiarid lands. Plant Disease. 70 (5), 405-409. DOI:10.1094/PD-70-405

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 DOI:10.1094/PDIS-04-12-0365-PDN

Tsai W S, Shih L, Green S K, Jan F J, 2007. Occurrence and molecular characterization of Squash leaf curl Philippines virus in Taiwan. Plant Disease. 91 (7), 907. HTTP://www.apsnet.org DOI:10.1094/PDIS-91-7-0907A

Distribution Maps