Invasive Species Compendium

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Datasheet

Squash leaf curl virus
(leaf curl of squash)

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Datasheet

Squash leaf curl virus (leaf curl of squash)

Summary

  • Last modified
  • 14 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Squash leaf curl virus
  • Preferred Common Name
  • leaf curl of squash
  • Taxonomic Tree
  • Domain: Virus
  •   Unknown: "ssDNA viruses"
  •     Unknown: "DNA viruses"
  •       Family: Geminiviridae
  •         Genus: Begomovirus

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Pictures

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PictureTitleCaptionCopyright
Symptoms of SLCV CA in winter squash.
TitleSymptoms
CaptionSymptoms of SLCV CA in winter squash.
CopyrightJudith K. Brown
Symptoms of SLCV CA in winter squash.
SymptomsSymptoms of SLCV CA in winter squash.Judith K. Brown
Symptoms of WCMoV in watermelon.
TitleSymptoms
CaptionSymptoms of WCMoV in watermelon.
CopyrightJudith K. Brown
Symptoms of WCMoV in watermelon.
SymptomsSymptoms of WCMoV in watermelon.Judith K. Brown
Symptoms of WCMoV in zucchini squash.
TitleSymptoms
CaptionSymptoms of WCMoV in zucchini squash.
CopyrightJudith K. Brown
Symptoms of WCMoV in zucchini squash.
SymptomsSymptoms of WCMoV in zucchini squash.Judith K. Brown
Symptoms of WCMoV in melon.
TitleSymptoms
CaptionSymptoms of WCMoV in melon.
CopyrightJudith K. Brown
Symptoms of WCMoV in melon.
SymptomsSymptoms of WCMoV in melon.Judith K. Brown

Identity

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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

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

Notes on Taxonomy and Nomenclature

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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

Top 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.

Distribution

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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

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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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

IsraelPresentCABI/EPPO, 2014; EPPO, 2014
JordanPresentAl-Musa et al., 2008; CABI/EPPO, 2014; EPPO, 2014
LebanonPresentEPPO, 2012; Sobh et al., 2012; CABI/EPPO, 2014; EPPO, 2014
Saudi ArabiaPresentCABI/EPPO, 2014; EPPO, 2014
TaiwanPresentTsai et al., 2007

Africa

EgyptPresentCABI/EPPO, 2014; EPPO, 2014

North America

MexicoRestricted distributionMontes Garcia et al., 1998; Ramirez-Arredondo et al., 1995; CABI/EPPO, 2014; EPPO, 2014
USARestricted distributionCABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014
-ArizonaPresentBrown and Nelson, 1986; Rosemeyer et al., 1986; Brown and Nelson, 1989; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014
-CaliforniaPresentFlock and Mayhew, 1981; Cohen et al., 1983; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014
-TexasPresent1993Isakeit et al., 1994; CABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014

Central America and Caribbean

Costa RicaPresentCABI/EPPO and, 1997; EPPO, 2014
Dominican RepublicPresentCABI/EPPO and, 1997; EPPO, 2014
GuatemalaPresentCABI/EPPO and, 1997; CABI/EPPO, 2014; EPPO, 2014
HondurasPresentCABI/EPPO and, 1997; EPPO, 2014
NicaraguaPresentCABI/EPPO and, 1997; EPPO, 2014

Europe

BelgiumAbsent, no pest recordEPPO, 2014

Risk of Introduction

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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

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Plant nameFamilyContext
Citrullus lanatus (watermelon)CucurbitaceaeMain
Cucumis melo (melon)CucurbitaceaeMain
Cucumis sativus (cucumber)CucurbitaceaeMain
Cucurbita (pumpkin)CucurbitaceaeMain
Cucurbita maxima (giant pumpkin)CucurbitaceaeMain
Cucurbita moschata (pumpkin)CucurbitaceaeMain
Cucurbita pepo (marrow)CucurbitaceaeMain
Phaseolus vulgaris (common bean)FabaceaeMain

Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

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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

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SignLife StagesType
Growing point / distortion
Leaves / abnormal colours
Leaves / abnormal forms
Whole plant / distortion; rosetting
Whole plant / dwarfing

Biology and Ecology

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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

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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

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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 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
Bark
Growing medium accompanying plants
Roots
True seeds (inc. grain)
Wood

Vectors and Intermediate Hosts

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VectorSourceReferenceGroupDistribution
Bemisia tabaciEFSA Panel on Plant Health, 2013. Insect

Impact

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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

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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

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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

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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

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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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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.

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

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

Flock RA; Mayhew DE, 1981. Squash leaf curl, a new disease of cucurbits in California. Plant Disease, 65(1):75-76

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

Isakeit T; Robertson NL; Brown JK; Gilbertson RL, 1994. First report of squash leaf curl virus on watermelon in Texas. Plant Disease, 78(10):1010

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.

Mansour A; Al-Musa A, 1993. Cucumber vein yellowing virus: host range and virus vector relationships. Journal of Phytopathology, 137(1):73-78

McCreight JD, 1984. Tolerance of Cucurbita spp. to squash leaf curl. Report of Cucurbit Genetics Co-operative, USA, 7:71-72.

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

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.

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.

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.

Rosemeyer ME; Brown JK; Nelson MR, 1986. Five viruses isolated from field-grown buffalo gourd (Cucurbita foetidissima), a potential crop for semiarid lands. Plant Disease, 70(5):405-409

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

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.

Yilmaz MA; Ozaslan M; Ozaslan D, 1989. Cucumber vein yellowing virus in Cucurbitaceae in Turkey. Plant Disease, 73(7):610

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