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. 2026 Sep 14;27(9):e70345. doi: 10.1111/mpp.70345

Cucurbit Leaf Crumple Virus: An Important Pathogen of Cucurbit and Snap Bean Crops

Dalvir Kaur Dhadly 1, Saritha Raman Kavalappara 1, Sudeep Bag 1,✉, Theodore McAvoy 2, Paul M Severns 3, Rajagopalbabu Srinivasan 4, Alvin M Simmons 5
PMCID: PMC13575508  PMID: 42740504

ABSTRACT

Taxonomy

Cucurbit leaf crumple virus (CuLCrV); Begomovirus cucurbitae; Geminiviridae; Geplafuvirales.

Geographical Distribution

The presence of CuLCrV is exclusively limited to North America, mainly Mexico and the United States.

Physical Properties

CuLCrV is a bipartite begomovirus comprising two circular single‐stranded DNA molecules (DNA‐A and DNA‐B), encapsidated within geminate icosahedral particles.

Genome and Organization

CuLCrV possesses a bipartite genome of DNA‐A (2632 nucleotides) and DNA‐B (2600 nucleotides). DNA‐A contains five open reading frames (ORFs): AV1 (coat protein), AC1 (replication‐associated protein), AC2 (transcriptional activator protein), AC3 (replication enhancer protein) and AC4. DNA‐B contains two ORFs: BV1 (nuclear shuttle protein) and BC1 (movement protein).

Transmission

CuLCrV is transmitted by the sweetpotato whitefly, Bemisia tabaci , in a persistent, circulative and non‐propagative manner.

Hosts

CuLCrV primarily infects crop members of the Cucurbitaceae and snap bean ( Phaseolus vulgaris , Fabaceae). Multiple weed species belonging to Brassicaceae, Convolvulaceae, Cucurbitaceae and Verbenaceae act as persistent virus reservoir hosts.

Symptoms

Symptom expression varies with host and infection timing. In cucurbits, infection induces leaf crumpling, thickening and downward curling of leaves, with green streaks and distortion of fruits. In snap bean, symptoms include leaf distortion, chlorosis and malformed pods.

Control

No commercial cultivars with resistance to CuLCrV are available for cucurbit crops, although some resistance has been reported in snap bean cultivars. Therefore, management relies primarily on integrated disease management.

Keywords: Begomovirus, cucurbits, mixed infection, weed reservoirs, whitefly

Cucurbit leaf crumple virus (CuLCrV) is an emerging pathogen in the New World. High whitefly abundance and transmission efficiency, mixed infections, weed reservoirs and limited host resistance challenge cucurbit and snap bean production.

graphic file with name MPP-27-e70345-g001.webp

1. Introduction

Begomoviruses, members of the genus Begomovirus in the family Geminiviridae, are among the most damaging plant viruses globally, causing significant yield losses in tropical and subtropical agriculture (Fondong 2025; Navas‐Castillo et al. 2011). With over 464 species currently recognized by the International Committee on Taxonomy of Viruses, Begomovirus is the largest genus in the virosphere (ICTV 2025). These viruses induce a wide range of symptoms in host plants, including mosaic and mottle patterns, chlorosis, leaf curling and deformation, stunted plant growth and fruit loss (Fiallo‐Olivé and Navas‐Castillo 2023). Based on their genome organization, begomoviruses are divided into two groups: monopartite viruses, which contain a single circular single‐stranded DNA (ssDNA) molecule; and bipartite viruses, which have two genomic components, DNA‐A and DNA‐B. Most begomoviruses that originated in the New World (North, Central and South America) are bipartite, whereas most of those originating in the Old World (Africa, Europe and Asia) may be either monopartite or bipartite (Briddon et al. 2010; Brown and Czosnek 2002; Rojas et al. 2005).

Cucurbit leaf crumple virus (CuLCrV; Begomovirus cucurbitae) is one of the most economically important bipartite begomoviruses in the southeastern United States, particularly in Georgia and Florida, where squash ( Cucurbita pepo ) and snap bean ( Phaseolus vulgaris ) may experience up to 100% infection during severe epidemics (Adkins et al. 2011; Agarwal et al. 2021; Brown et al. 2000; Dutta et al. 2024; Gautam 2019; Guzman et al. 2000; Hendricks and Roberts 2023; Kavalappara et al. 2021; LaTora et al. 2022). By comparison, this virus is of limited economic importance in the southwestern United States (Hagen, Rojas, Sudarshana, et al. 2008; Mondal et al. 2023). These contrasting regional patterns indicate that the epidemiological significance of CuLCrV varies across the cucurbit production regions in the United States.

Transmitted by the sweetpotato whitefly ( Bemisia tabaci ), CuLCrV frequently occurs as a mixed infection with other whitefly‐transmitted viruses (WTVs). In cucurbits, these include cucurbit chlorotic yellows virus (CCYV), cucurbit yellow stunting disorder virus (CYSDV; Crinivirus cucurbitae) and squash vein yellowing virus (SqVYV; Ipomovirus cucurbitavenaflavi) (Hendricks and Roberts 2023; Kavalappara et al. 2021; Kousik and Adkins 2020); whereas in snap bean CuLCrV commonly occurs with sida golden vein mosaic virus (SiGMV; Begomovirus sidaureidis) (Gautam et al. 2023). These co‐infections can complicate diagnosis because symptom expression often overlaps among associated viruses (Hendricks and Roberts 2023; Kavalappara et al. 2021). Particularly in Georgia, mixed infections involving CuLCrV, CCYV and CYSDV are highly prevalent and frequently exceed the incidence of single‐virus infections, resulting in greater symptom severity and reduced marketable yield (Acharya, McAvoy, et al. 2025; Acharya et al. 2026; Adeleke et al. 2022; Kavalappara et al. 2021). Similarly in Florida, CuLCrV occurs in complex virus associations involving CCYV, CYSDV, SqVYV, and watermelon crinkle leaf‐associated viruses (WCLaV‐1 and WCLaV‐2) (Hendricks and Roberts 2023). Co‐occurrence of CuLCrV with WCLaV‐1 and WCLaV‐2 has also been documented in Georgia (Kavalappara, Acharya, et al. 2024).

CuLCrV was initially identified in the autumn of 1998 from volunteer watermelon plants displaying symptoms of leaf crumpling and yellowing in the Imperial Valley of California (Guzman et al. 2000). During the same period, another begomovirus provisionally designated as cucurbit leaf curl virus was found infecting pumpkin ( C. pepo ), honeydew melon ( Cucumis melo ) and muskmelon ( C. melo ) in Arizona, Texas and Mexico (Brown et al. 2000). However, later research identified and designated this virus as CuLCrV (Hagen, Rojas, Sudarshana, et al. 2008). Subsequently, CuLCrV was reported in the southeastern United States on yellow straight‐neck squash and zucchini squash in Florida in 2006 (Akad et al. 2008). Shortly thereafter, it was detected from snap bean in Georgia (Larsen and Kmiecik 2010) and muskmelon, summer squash ( C. pepo ), and watermelon ( Citrullus lanatus ) in South Carolina (Keinath et al. 2018). Severe incidence of CuLCrV in association with CCYV and CYSDV on various autumn‐grown cucurbits, such as cantaloupe ( Cucumis melo var. cantalupensis), cucumber ( Cucumis sativus ), yellow squash and zucchini, was observed in field surveys conducted in Georgia during 2019–2020 (Gautam et al. 2020; Kavalappara et al. 2021).

The tremendous population increase of B. tabaci in Florida and Georgia in recent years, coupled with the heavy incidence of CuLCrV, has resulted in extensive losses in yield, fruit quality and marketability of cucurbits and snap bean crops (Candian et al. 2021; Dawson 2016; Dutta 2019; Gadhave et al. 2020; Martini et al. 2016; McAvoy 2017). The increasing impact of CuLCrV can be attributed to the high transmission efficiency (Ghosh and Ghanim 2021) and broad host range (Adkins et al. 2011; Sparks et al. 2018) of its whitefly vector, which allows large whitefly populations to develop. In the southeastern United States, year‐round vegetable production, mild winters and the prevalence of weed hosts sustain whitefly populations throughout the year (Barman et al. 2022; Sparks et al. 2018). CuLCrV has been detected in cucurbitaceous weeds such as balsam apple ( Momordica charantia ) and smellmelon ( Cucumis melo var. dudaim) (Adkins et al. 2008; Adkins, Webster, et al. 2009). Additionally, weeds such as wild radish ( Raphanus raphanistrum; Brassicaceae), Lantana camara (Verbenaceae) and cypressvine morning glory ( Ipomoea quamoclit; Convolvulaceae) and tall morning glory ( Ipomoea purpurea ; Convolvulaceae) have been demonstrated to act as virus reservoirs in Georgia (Dhadly et al. 2025a). The recent discovery of agroinoculation‐mediated seed transmission of CuLCrV (Dhadly et al. 2025b) in yellow squash suggests a potential transmission route that warrants further investigation under natural field conditions. With limited host resistance against the virus complex and its whitefly vector, current management practices rely on integrated approaches, including the use of reflective mulches, row covers and chemical insecticides. However, their effectiveness and adoption may be limited by increased production costs (LaTora et al. 2022). This underscores the need to deepen our understanding of CuLCrV biology, epidemiology and the development of sustainable management options.

2. CuLCrV Parameters

2.1. Geographical Distribution

Unlike its vector, CuLCrV has only been reported from North America (Figure 1A). Following its initial detection in California in 1998 (Guzman et al. 2000), the virus was reported in Arizona and Texas in the United States and Coahuila in north Mexico, between 1998 and 1999 (Brown et al. 2000). It later emerged in Florida in 2006 (Akad et al. 2008), was detected in Georgia by 2009 (Larsen and Kmiecik 2010) and subsequently in South Carolina in 2017 (Keinath et al. 2018) (Figure 1B). More recently, CuLCrV was identified in Campeche in southeast Mexico in 2019 (Rodríguez‐Negrete et al. 2021). In addition to cultivated cucurbits, CuLCrV has also been detected in weed hosts in California (Hagen, Rojas, Sudarshana, et al. 2008), Florida (Adkins et al. 2008; Adkins, Webster, et al. 2009; Hendricks and Roberts 2023) and Georgia (Dhadly et al. 2025a). This distribution highlights the virus's capacity for regional spread, likely driven by efficient whitefly transmission, the presence of reservoir hosts and favourable environmental conditions.

FIGURE 1.

FIGURE 1

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(A) Distribution of cucurbit leaf crumple virus (CuLCrV) in North America. (B) Distribution in the United States showing year of first detection by state. (C) Transmission electron micrograph (TEM) of virus particles in symptomatic yellow squash leaf from a field in Georgia (scale bar = 200 nm); the flexuous rod‐shaped particle is consistent with a crinivirus, namely, cucurbit chlorotic yellows virus or cucurbit yellow stunting disorder virus. (D) TEM of a CuLCrV geminate particle from the same sample (scale bar = 50 nm). (E) Schematic of the DNA‐A and DNA‐B components with predicted open reading frames (shown by arrows) and the conserved common region (shown by red T‐loop). Photo credits (C–D): Dr. Tian Tongyan, California Department of Food and Agriculture. Panel (A‐B and E) was created with BioRender.

2.2. Physical Properties

No studies have specifically been reported on the characterization of the physical properties of CuLCrV. Transmission electron microscopy (TEM) of the field‐infected squash plants indicated that CuLCrV has geminate (twinned) particles typical of members of the family Geminiviridae (Figure 1C,D).

2.3. Genome Organization

CuLCrV is a bipartite begomovirus consisting of two circular single‐stranded (ss) DNA molecules. DNA‐A is 2632 nucleotides (nt; NC_002984) and DNA‐B is 2600 nt (NC_002985). In many bipartite begomoviruses, the DNA‐A component encodes two proteins (AV1 and AV2) on the virion‐sense strand and four proteins (AC1‐4) on the complementary‐sense strand, whereas the DNA‐B component encodes one protein on each sense (BV1) and complementary sense strand (BC1, Hull 2014). However, in New World begomoviruses including CuLCrV, the AV2 gene (pre‐coat protein) is absent (Briddon et al. 2010; Fiallo‐Olivé and Navas‐Castillo 2023). Analysis of CuLCrV sequence for open reading frames (ORFs) shows that DNA‐A contains five ORFs, namely, AV1 (coat protein, CP; 217–966) on the virion‐sense strand and AC1 (replication‐associated protein, Rep; 1433–2563), AC2 (transcriptional activator protein, TrAP; 1102–1497), AC3 (replication enhancer protein, Ren; 963–1361) and AC4 (2044–2409) on the complementary‐sense strand (Hagen, Rojas, Sudarshana, et al. 2008). DNA‐B contains two ORFs: BV1 (nuclear shuttle protein, NSP; 521–1291) on the virion‐sense strand and BC1 (movement protein, MP; 1479–2408) on the complementary‐sense strand (Figure 1E; Hagen, Rojas, Sudarshana, et al. 2008). The genes are transcribed bidirectionally, with a noncoding intergenic region also known as the common region (CR) between the virion‐sense and complementary‐sense strands. This region contains a stem‐loop structure with the conserved nonanucleotide sequence TAATATTAC, which serves as the origin of replication (V‐ori). The CR between DNA‐A and DNA‐B is 194 nt long and 98% identical. The replication‐associated protein binds to high‐affinity pentanucleotide sites, GGTGTCCTGGTGT (Brown et al. 2002; Hagen, Rojas, Sudarshana, et al. 2008). Begomoviruses typically replicate via a rolling‐circle replication mechanism, but also via a recombination‐dependent replication mechanism (Jeske et al. 2001). DNA‐A of CuLCrV, being the most important genome component, which is homologous to monopartite begomoviruses, encodes proteins required for replication, regulation of gene expression, host defence resistance, encapsidation and insect transmission. In contrast, DNA‐B encodes two proteins with functions in intra‐ and intercellular movement in host plants (Briddon et al. 2010; Fiallo‐Olivé et al. 2021; Hagen, Rojas, Sudarshana, et al. 2008).

2.4. Phylogenetic Analyses

Sequence comparisons and phylogenetic analyses revealed that CuLCrV was most closely related to squash leaf curl virus (SLCuV) (Brown et al. 2002; Guzman et al. 2000) and this was further supported by the capacity of a CuLCrV isolate from Arizona (originally referred to as cucurbit leaf curl virus) to form infectious pseudorecombinants with SLCuV (Brown et al. 2002). The evolutionary relationships among CuLCrV and related Begomovirus isolates from cucurbit and snap bean in North America (Table S1) were analysed in this study using the maximum‐likelihood method based on the Tamura–Nei model in MEGA X with default parameters (Kumar et al. 2018; Tamura and Nei 1993). DNA‐A of CuLCrV Georgia isolate (PP617367.1) clustered tightly with CuLCrV isolates from Campeche, Mexico (MW273384.1), Baja California Sur, Mexico (OQ466347.1), Arizona (AF256200.4) and California (NC_002984.1), forming a distinct, well‐supported CuLCrV clade that is clearly separated from other bipartite begomoviruses infecting cucurbits and from snap bean‐associated begomoviruses (Figure 2A). A similar phylogenetic topology was observed when analysing the AV1 gene sequence (Figure 2B). Likewise, DNA‐B of CuLCrV Georgia isolate (PP617368.1) grouped closely with DNA‐B sequences from CuLCrV isolates from Campeche, Mexico (MW273385.1), Baja California Sur, Mexico (OQ466346.1), Arizona (AF327559.1) and California (NC_002985.1), forming a tight CuLCrV DNA‐B subgroup that was distinct from other cucurbit‐infecting and snap bean begomoviruses in the region (Figure 2C). This pattern was also reflected in the BC1 gene phylogeny (Figure 2D).

FIGURE 2.

FIGURE 2

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Maximum‐likelihood phylogenetic trees of cucurbit leaf crumple virus and related begomoviruses infecting cucurbit and snap bean ( Phaseolus vulgaris ) hosts in North America, based on (A) DNA‐A, (B) the AV1 gene, (C) DNA‐B and (D) the BC1 gene. Bootstrap support values greater than 50% are shown at the corresponding nodes. Coloured branches indicate CuLCrV isolates (pink), other cucurbit‐infecting begomoviruses (green) and snap bean‐infecting begomoviruses (blue). Tip labels include the GenBank/NCBI accession number, virus acronym and isolate location.

Interestingly, DNA‐A of Old World squash leaf curl China virus (SLCCNV) isolate (AM260205.1) clustered with watermelon chlorotic stunt virus (WmCSV) isolates from Mexico (PP622784.1, KY124280.1) and Arizona (PQ399661.1). A similar association was observed for the DNA‐B SLCCNV isolate (AM260207.1), which grouped with WmCSV DNA‐B sequences from Mexico (PP622788.1) and Arizona (PQ399662.1), suggesting possible shared evolutionary origins among these New World viruses. Recombination analysis was conducted using the Recombination Detection Program (RDP5; Martin et al. 2021) with default settings and detected no statistically supported recombination events in the DNA‐A or DNA‐B components of CuLCrV isolates analysed in the present study. Additionally, no evidence of interspecific recombination was identified between CuLCrV and other closely related cucurbit‐infecting begomoviruses included in the analysis, indicating limited recombination and overall genomic stability within the CuLCrV population.

2.5. Host Range

CuLCrV infects a wide range of cucurbitaceous crops including cantaloupe (Brown et al. 2000; Hagen, Rojas, Sudarshana, et al. 2008; Kavalappara et al. 2021), Charentais melon ( C. melo ‘Charentais’) (Hagen, Rojas, Sudarshana, et al. 2008), cucumber (Brown et al. 2002; Hagen, Rojas, Sudarshana, et al. 2008; Kavalappara et al. 2021), honeydew melon ( C. melo ) (Brown et al. 2000, 2002), muskmelon ( C. melo ) (Brown et al. 2002; Keinath et al. 2018), pumpkin ( C. pepo ) (Brown et al. 2000, 2002; Hagen, Rojas, Sudarshana, et al. 2008), summer squash (e.g., yellow squash and zucchini) (Akad et al. 2008; Brown et al. 2002; Hagen, Rojas, Sudarshana, et al. 2008; Kavalappara et al. 2021) and watermelon (Adeleke et al. 2022; Akad et al. 2008; Brown et al. 2002; Guzman et al. 2000; Hagen, Rojas, Sudarshana, et al. 2008; Keinath et al. 2018; Rodríguez‐Negrete et al. 2021). Among winter squash, CuLCrV has been reported to infect Turk's Turban ( Cucurbita maxima ) and Blue Hubbard squash ( C. maxima ) and acorn squash ( C. pepo ), Jack‐o’‐lantern pumpkins ( C. pepo ) and birdhouse gourd ( Lagenaria siceraria ) (Hagen, Rojas, Sudarshana, et al. 2008). Within the Fabaceae, the only confirmed natural host is snap bean (Adkins, Polston, and Turechek 2009; Hagen, Rojas, Sudarshana, et al. 2008; Larsen and Kmiecik 2010). Experimentally, CuLCrV also infects Nicotiana benthamiana (Solanaceae) (Brown et al. 2002), a widely used model host for infectious clone‐based studies of plant viruses.

In addition to crop hosts, CuLCrV has been detected in smellmelon (Adkins, Webster, et al. 2009) and balsam apple (Adkins et al. 2008). Wright's groundcherry (Physalis sp., Solanaceae) was also reported to harbour CuLCrV; however, further research is needed to confirm its host status (Hendricks and Roberts 2023). CuLCrV was also detected in weeds collected from vegetable fields such as cypress vine morning glory, tall morning glory, pigweed (Amaranthus sp., Amaranthaceae), wild radish and lantana ( Lantana camara ) (Dhadly et al. 2025a; see Section 2.9 for additional details) and perennial coyote melon ( Cucurbita palmata ) (Hagen, Rojas, Sudarshana, et al. 2008).

2.6. Symptomatology

CuLCrV infection produces diverse symptoms depending on the host species and the plant growth stage at the time of infection. In general, symptoms first appear on newly emerging leaves and include chlorotic spots, leaf crumpling, thickening and downward curling (Hagen, Rojas, Sudarshana, et al. 2008). In yellow squash inoculated with viruliferous whiteflies at the two‐true‐leaf stage, chlorotic spots typically develop by 5 days post‐inoculation (dpi; Figure 3A), progressing to pronounced leaf crumpling and downward curling by 30 dpi under controlled greenhouse conditions (Figure 3B; 28°C ± 3°C, 50% ± 20% relative humidity and a 16 ± 8 h light:dark cycle; October 2024). In the field, the symptoms observed in the autumn are mainly due to mixed infection of WTVs and potyviruses (Acharya, Kumar, et al. 2025; Devendran et al. 2023). Early‐season field infections frequently cause severe deformation of young leaves and pronounced plant stunting (Figure 3C,D). Infected plants often produce fruits with characteristic green streaks and a significant size reduction. In severe cases, fruit formation may be greatly reduced or completely absent and any fruits that do develop frequently fail to mature properly (Figure 3E). In contrast, late‐season infections are generally milder, with symptoms largely restricted to crumpling and curling of younger leaves, while older foliage (Figure 3F) and fruit quality remain unaffected. In zucchini squash, CuLCrV infection induces crumpling and downward curling of young leaves (Figure 3G). Among winter squash, butternut squash and calabaza squash develop less severe symptoms, with no symptoms observed on the fruits (Acharya, McAvoy, et al. 2025). In pumpkins, field‐infected plants show severe deformation of new growth (Figure 3H). Cucumber plants typically do not exhibit the characteristic CuLCrV symptoms on new growth; instead, interveinal chlorosis and yellowing of the lower leaves are more prominent due to mixed infection with CCYV and/or CYSDV (Figure 3I; Acharya, McAvoy, et al. 2025). In melons, infected cantaloupe plants exhibit leaf crumpling and curling of new growth, particularly along runners (Figure 3J). Infected watermelon plants display bright yellow chlorosis, marginal leaf necrosis and crumpling of young leaves (Figure 3K). Severe infections can result in complete fruit set failure or prevent fruits from reaching maturity, similar to observations in yellow squash. CuLCrV infection in watermelons was reported to reduce plant vigour and yield by 15%–20% in South Carolina, primarily due to reduced fruit size and failure of fruit to reach marketable maturity (Keinath et al. 2018; Keinath and Ling 2023; Rodríguez‐Negrete et al. 2021). In contrast, casaba and honeydew melons generally lack visible foliar symptoms (Hagen, Rojas, Sudarshana, et al. 2008).

FIGURE 3.

FIGURE 3

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Symptoms associated with cucurbit leaf crumple virus infection. In yellow squash, greenhouse symptoms include (A) chlorotic spots and (B) leaf crumpling with downward curling. Field symptoms under mixed infection conditions in yellow squash include (C) downward curling and mottling; (D) plant stunting with thickened, deformed young leaves and green streaking on fruits during early infection; (E) malformed, undersized fruits compared with a fully developed fruit from an asymptomatic plant; and (F) mild crumpling, downward curling and chlorosis of young leaves during late infection. In other crop hosts, young leaves show (G) crumpling, mottling and downward curling in zucchini squash; (H) curling and deformation in pumpkin; (I) no symptoms on new growth but yellowing of lower leaves in cucumber; (J) crumpling and downward curling in cantaloupe; (K) chlorosis and crumpling in watermelon; and (L) interveinal chlorosis and vein greening in snap bean. In non‐crop hosts, symptoms include (M) leaf curling in cypress vine morning glory, a normal leaf is shown in a white circle for comparison; (N) interveinal chlorosis and leaf puckering in tall morning glory; (O) chlorosis in pigweed; (P) chlorosis with leaf blistering in wild radish; and (Q) rolling of upper leaves in lantana. White circles indicate enlarged and cropped sections corresponding to panels I–K to better illustrate symptom details.

In snap bean, symptoms observed under field conditions are associated with co‐infection by CuLCrV and SiGMV. These infections result in interveinal chlorosis, vein greening, blistering and rugosity (Figure 3L), as well as malformed pods that are often curled, misshapen and unmarketable (Agarwal et al. 2021; Larsen and Kmiecik 2010). Among non‐crop hosts, CuLCrV infection in association with CCYV produces a range of symptoms, including leaf curling in cypressvine morning glory (Figure 3M), interveinal chlorosis and leaf puckering in tall morning glory (Figure 3N), chlorosis in pigweed (Figure 3O) and chlorosis with leaf blistering in wild radish (Figure 3P). Leaf rolling was observed in the perennial weed lantana infected with CuLCrV (Figure 3Q).

Notably, CuLCrV produces the most severe symptoms in yellow squash and pumpkin, whereas a recovery phenotype is observed in watermelon and cantaloupe, in which severe symptoms observed during early stages of infection gradually attenuate as the disease progresses. This phenomenon has been documented under both experimental and field conditions (Hagen, Rojas, Kon, and Gilbertson 2008). Recovered plants can produce commercially acceptable yields despite early infection. This recovery is associated with reduced viral DNA titers through activation of host RNA‐silencing defences. Recovered plants also exhibit resistance to reinfection; however, this response can be suppressed by co‐infection with viruses encoding strong silencing suppressors, such as cucumber mosaic virus. Recovery is most efficient in indeterminate cucurbits with continuous vine growth and new meristematic tissue like cantaloupe and watermelon. This recovery is further associated with enhanced viral genome methylation in strongly recovering hosts, suggesting epigenetic suppression of viral replication or gene expression (Hagen, Rojas, Kon, and Gilbertson 2008).

2.7. Diagnosis

The most important step in the management of any virus in plants is through precise and timely detection with accuracy, specificity and inexpensive methods. In the case of CuLCrV, symptoms alone are not sufficient to diagnose with confidence owing to the high frequency of mixed infections with more than one virus. There are no serological tests available for CuLCrV detection; however, PCR and quantitative PCR (qPCR) are commonly used as confirmatory laboratory assays (Agarwal et al. 2021; Brown et al. 2000; Guzman et al. 2000; Hagen, Rojas, Sudarshana, et al. 2008). Multiplex reverse transcriptase‐based PCRs (RT‐PCRs) have also been developed for the simultaneous detection of RNA and DNA viruses in cucurbits, namely, CuLCrV, CYSDV, SqVYV and CCYV (Jailani et al. 2021). In Georgia, high‐throughput sequencing was used to identify CuLCrV in cantaloupe, cucumber and yellow squash in association with CCYV and CYSDV (Kavalappara et al. 2021). More recently, a multiplex TaqMan assay was developed that allowed simultaneous detection of WTVs from cucurbit leaf tissues and whitefly (Jailani and Paret 2024).

Field‐deployable, rapid isothermal amplification assays have also been developed for CuLCrV detection. Specifically, Kalischuk et al. (2022) developed recombinase polymerase amplification (RPA) and exonuclease RPA (exo‐RPA) assays providing results within 30 min, high sensitivity and no cross‐reactivity with non‐target viruses. A multiplex RT‐RPA assay is also available for the simultaneous detection of CuLCrV and criniviruses from crude plant extracts, enabling the rapid and cost‐effective identification of mixed infections (Jailani and Paret 2023). There is also a real‐time loop‐mediated isothermal amplification (LAMP) assay that provides results within 60 min and has greater sensitivity than endpoint PCR (Waliullah et al. 2020). An AmplifyRP XRT test kit (Agdia Inc.) is also commercially available for both field‐based and laboratory assays. More recently, an optimized RPA–lateral flow test (RPA‐LFT) assay was developed for on‐site CuLCrV detection directly from plant tissues and whitefly vectors, enabling instrument‐free diagnostics with high sensitivity (Jailani and Paret 2025). Despite the availability of these assays, conventional/endpoint PCR and qPCR remain the preferred confirmatory diagnostics due to their robustness and reproducibility across laboratories.

2.8. Transmission

CuLCrV is transmitted by the sweetpotato whitefly, B. tabaci , a species complex of more than 40 cryptic species (Brown et al. 2023). Among these, the MEAM1 and Mediterranean (MED) species are the most invasive and economically damaging members worldwide (Brown et al. 1995, 2023; De Barro 2011; Perring 2001; Wan et al. 2009). Molecular surveys indicate that MEAM1 predominates in vegetable and row‐crop farmscapes in Georgia, with no significant genetic differentiation among populations collected from crops, weeds or different geographic locations (Gautam et al. 2020). Although both MEAM1 and MED can acquire CuLCrV, successful transmission to healthy plants has been observed only for MEAM1 (Gautam et al. 2022).

Bemisia tabaci transmits CuLCrV in a persistent, circulative and non‐propagative manner, such that once acquired, whiteflies can retain and transmit CuLCrV for life (Boykin et al. 2007; Boykin 2014; Czosnek et al. 2002; Dinsdale et al. 2010; Gautam et al. 2022; Hagen, Rojas, Sudarshana, et al. 2008). An adult whitefly requires a minimum acquisition access period (AAP) of ~30 min on an infected plant to acquire CuLCrV, after which the virus circulates within the insect and becomes transmissible following a 6–8‐h latent period. Transmission to a healthy plant then requires an inoculation feeding period of 15–30 min (Keinath and Ling 2023). In experimental assays, successful whitefly‐mediated CuLCrV transmission was observed using a 48‐h AAP and inoculation access period (Gautam et al. 2022). Very low frequency of vertical (transovarial) and horizontal (mating) transmission of CuLCrV by B. tabaci has been detected experimentally. However, because neither route resulted in subsequent plant infection, these transmission pathways are unlikely to maintain CuLCrV inoculum between cropping seasons (Gadhave et al. 2020).

CuLCrV is phloem‐limited and is not known to be mechanically transmitted through infected plant sap. The use of infectious clones to study specific host–virus interactions is indispensable because they enable reproduction of CuLCrV infection under controlled conditions without the need for viruliferous whiteflies or naturally infected source plants. These constructs typically contain tandem repeats of the viral genome, including duplicated viral origins of replication (V‐ori), which facilitate efficient viral replication and allow the virus to complete its life cycle in plants. Delivery of these clones is typically achieved via Agrobacterium‐mediated inoculation or particle bombardment (Hagen, Rojas, Sudarshana, et al. 2008; Kavalappara, Devendran, et al. 2024). However, efficient agroinoculation of yellow squash is hindered by its densely packed mesophyll tissue. To overcome this limitation, Kavalappara, Devendran, et al. (2024) developed a microneedle‐assisted agroinoculation technique for delivering CuLCrV infectious clones into yellow squash (Kavalappara, Devendran, et al. 2024). In a follow‐up study using this system, CuLCrV accumulated at high levels in flower and fruit tissues and was also detected in the seed embryo. In grow‐out tests, weak symptoms were observed in progeny seedlings. Full‐length CuLCrV DNA‐A and DNA‐B were amplified in various seed and seedling tissues, including true leaves of a progeny plant (Dhadly et al. 2025b). These findings demonstrate the potential for transmission of CuLCrV via seeds under experimental agroinoculation conditions. However, whether natural CuLCrV infections result in seed transmission and the epidemiological significance of this pathway under field conditions remain to be determined (Srinivasan 2025).

2.9. Epidemiology

CuLCrV was first detected in the southwestern United States, where cantaloupe and honeydew melon dominate cucurbit production; it has become more problematic in the Southeast, where squash and other cucurbits are major crops. In the Southwest, limited reservoir hosts, recovery responses in infected plants and extended crop‐free period from November to February likely restrict interseasonal persistence (Gilbertson 2017). CuLCrV incidence is typically lower in spring‐planted cucurbits and higher in autumn crops in the southwestern United States, reflecting seasonal increases in whitefly populations (Hagen, Rojas, Sudarshana, et al. 2008). A similar trend occurs in Georgia, where whitefly populations are low in spring cucurbits but increase during summer. Under hot, dry summer conditions, the sweetpotato whitefly can complete a generation in as little as 14 days, promoting rapid population buildup on summer crops such as cotton (Barman et al. 2023; Sparks et al. 2018). Cotton serves as an important propagative host for whiteflies, enabling large whitefly populations to disperse into nearby vegetable fields at the onset of autumn. This leads to severe outbreaks of WTVs, including CuLCrV in cucurbit and snap bean crops (Adeleke et al. 2022; Gautam 2019; Kavalappara et al. 2021).

CuLCrV persistence is further facilitated by alternative host plants. The availability of multiple host species increases opportunities for primary spread and vector acquisition (Dawson 2016; Dhadly et al. 2025a; Martini et al. 2016; Moury et al. 2017). CuLCrV was detected in weeds growing around cucurbit fields during crop‐free periods, indicating that CuLCrV can persist in these weed hosts between cropping seasons. Greenhouse transmission assays were further conducted on these weed hosts, namely, cypress vine morning glory, lantana, tall morning glory and wild radish. Whiteflies successfully transmitted CuLCrV from infected squash to weeds and from infected weeds back to squash, supporting the role of these weed species as virus reservoirs (Dhadly et al. 2025a). Volunteer watermelon, smellmelon and balsam apple plants bridging the spring and autumn cropping seasons serve as over‐summering hosts for CuLCrV, whereas crop plants serve as primary overwintering hosts because the major production season occurs during winter months (Adkins et al. 2008, 2011; Adkins, Webster, et al. 2009).

3. Virus Management

Host plant resistance represents a cornerstone of sustainable virus management; however, management of CuLCrV is particularly challenging due to the limited availability of resistant cultivars. Current control strategies primarily rely on cultural and chemical methods (LaTora et al. 2022; Nyoike et al. 2008). However, heavy whitefly abundance, especially in the southeastern United States during the autumn, together with the development of insecticide resistance and changes in insecticide susceptibility, can reduce the effectiveness of chemical control (Dimase, De Marchi, et al. 2024; Dimase, Rossitto De Marchi, et al. 2024; Perier, Cremonez, Parkins, et al. 2024; Perier, Cremonez, Smith, et al. 2024; Smith et al. 2016). Additionally, excessive insecticide application poses risks to applicators, non‐target organisms and the environment while disrupting natural pest control by beneficial insects.

3.1. Host Plant Resistance

Host plant resistance to whiteflies and/or CuLCrV represents a sustainable strategy for long‐term disease management. However, frequent mixed infections complicate resistance breeding, as durable control would require resistance to multiple whitefly‐transmitted viruses. Leaf trichomes are widely documented to influence B. tabaci abundance (Amini et al. 2021; Oliveira et al. 2021; Oriani and Vendramim 2010; Shi et al. 2023; Snyder et al. 1998; Wang et al. 2017). Nonglandular trichomes often favour whiteflies, whereas glandular trichomes may deter adults through repellent or toxic exudates (Do Prado et al. 2016; Oriani and Vendramim 2010). The glabrous leaf trait in melon has been identified as a source of resistance to whiteflies (Riley et al. 2001). In contrast, diploid watermelon ( Citrullus colocynthis ) accessions PI 386015, PI 386018 and PI 386024 were among the most resistant to B. tabaci under controlled conditions despite higher trichome densities than cultivated watermelon (Simmons and Levi 2001).

Resistance and tolerance to whiteflies and WTVs vary among cucurbit hosts. Cucumber is resistant to squash silverleaf (SSL) disorder and to curling/crumpling symptoms caused by CuLCrV. Zucchini is more tolerant to leaf silvering and WTV symptom severity than yellow squash, whereas butternut winter squash shows comparatively greater tolerance among long‐season cucurbits (Acharya, McAvoy, et al. 2025). Additionally, several breeding lines and wild germplasm accessions have been evaluated as potential sources of resistance to CuLCrV. In melon, breeding line MR‐1 and plant introductions PI 124111, PI 124112, PI 179901, PI 234607, PI 313970 and PI 414723 show partial resistance to CuLCrV under field and greenhouse conditions (McCreight et al. 2008). PI 236355 exhibited complete resistance in greenhouse assays. Genetic analysis identified a single recessive gene, culcrv, conferring resistance in PI 313970 (McCreight et al. 2008). Despite the identification of the culcrv gene in melon, its application in large‐scale breeding programmes remains limited. Germplasm screening in Georgia and Florida identified Cucurbita moschata , C. ecuadorensis and C. okeechobeensis as potential resistance sources to CuLCrV, CYSDV and whiteflies (Luckew et al. 2022). In watermelon, screening in 2022–2023 identified C. mucosospermus accessions (PI 494528 and PI 595203) and C. ecirrhosus accession (Grif 16,444) with reduced CuLCrV‐like symptom severity and virus loads (Luckew et al. 2025). Reduced virus load in Grif 16,444 may be associated with reported whitefly non‐preference, as observed in another C. ecirrhosus accession (PI 673135; Simmons et al. 2019).

In snap bean, commercially available cultivars and other genotypes have also been evaluated for resistance to CuLCrV and sida golden mosaic Florida virus (SiGMFV). Twenty Phaseolus genotypes showed moderate to high resistance to CuLCrV and SiGMFV under field conditions (Agarwal et al. 2021) and commercial cultivars Sybaris, Tema, PV 857, Momentum and Prevail exhibited moderate resistance (Dutta 2022). Collectively, these resistance sources provide valuable resources for breeding cultivars with improved tolerance to whiteflies and/or CuLCrV.

3.2. Vector Management Strategies

Management of vectors is a widely used strategy for controlling vector‐transmitted viruses. Chemical insecticides remain one of the primary tools used by vegetable growers to manage whiteflies and WTVs, including CuLCrV in open field conditions. Although several products can reduce whitefly populations and SSL intensity (Castle et al. 2009; Nyoike and Liburd 2008, 2010), current insecticide modes of action recommended for cucurbits and snap beans in the southeastern United States are outlined in Table S2 and updated regularly through regional extension sources (Georgia Pest Management Handbook 2024; Southeast U.S. Vegetable Crop Handbook 2026). Regularly monitoring crops for whiteflies and timely providing control measures can result in the best protection for the crops (Sparks et al. 2018). Recent field trials indicate that cyantraniliprole and flupyradifurone provide the strongest protection against adult whiteflies, whereas afidopyropen and spirotetramat plus pyriproxyfen are most effective against nymphs (LaTora et al. 2022). Cyantraniliprole and flupyradifurone also reduce virus symptom severity, including CuLCrV symptoms in yellow squash, likely because they rapidly suppress vector feeding and thereby limit virus inoculation and secondary spread (Caballero et al. 2015; Castle et al. 2017; LaTora et al. 2022; Nauen et al. 2015). However, insecticides can reduce virus disease severity and not incidence because a single viruliferous whitefly is sufficient to transmit this virus (Rahman et al. 2017). Moreover, reduced susceptibility and variable responses of B. tabaci populations to several insecticide classes have been reported in the southeastern United States, highlighting the importance of resistance monitoring and rotation among insecticide modes of action (Caballero et al. 2013; Dimase, De Marchi, et al. 2024; Dimase, Rossitto De Marchi, et al. 2024; Perier, Cremonez, Parkins, et al. 2024; Perier, Cremonez, Smith, et al. 2024; Rossitto De Marchi et al. 2021; Schuster et al. 2008; Smith et al. 2016). Field observations from the southeastern United States further suggest that the residual activity of some insecticides has declined over time, necessitating more frequent applications to maintain whitefly control (Whittaker 2019; Sparks, personal communication). Nevertheless, insecticide applications, when used judiciously within an integrated pest management framework, remain an important part of whitefly and WTVs management.

Beyond synthetic insecticides, additional tools may complement CuLCrV management. Acibenzolar‐S‐methyl, a salicylic acid analogue, does not directly affect whiteflies but induces systemic acquired resistance by activating the salicylic acid pathway and may be integrated into CuLCrV management programs to enhance plant defences (LaTora et al. 2022). Extracts of Chenopodium ambrosioides have also been reported to reduce adult whitefly numbers on yellow squash seedlings, although their effects on CuLCrV incidence were limited (LaTora et al. 2022). Kaolin clay, a mineral particle film, decreases whitefly adult settling and oviposition when applied to cucurbit foliage (Díaz‐Pérez et al. 2023; Liang and Liu 2009). Kaolin clay may be used alone or in rotation with conventional insecticides as a whitefly repellent, and its efficacy for whitefly and WTV management may be enhanced when combined with limonene, a citrus essential‐oil derived botanical repellent (Johnston et al. 2022; Martini et al. 2022). Similarly, neem oil and soap reduced whitefly indices in zucchini squash fields (Díaz‐Pérez et al. 2023).

3.3. Cultural Practices for Mitigating Whiteflies and the WTVs, Including CuLCrV

Preventing CuLCrV infections in cucurbits begins with the use of virus‐free seedlings. Protecting seedlings during their early growth stages is critical, as they are most vulnerable to virus transmission. The use of insect exclusion netting in greenhouses in conjunction with insecticides has been shown to significantly reduce whitefly numbers and virus incidence (LaTora et al. 2022).

UV‐reflective mulches applied during field‐bed preparation serve as an effective CuLCrV management tool in open‐field cucurbit production (LaTora et al. 2022; Martini et al. 2022). Insect‐proof row covers installed immediately after planting to protect seedlings during the most vulnerable stage have been shown to be as effective as, or more effective than, insecticides in reducing whitefly populations, SSL intensity, virus severity and boosting marketable squash yield (LaTora et al. 2022). Live mulches are another promising strategy, offering comparable whitefly control, virus suppression and yields to reflective mulch and insecticide treatments (Hilje and Stansly 2008; Nyoike et al. 2008), while enhancing the abundance of natural enemies that contribute to whitefly suppression and reducing virus transmission (Frank and Liburd 2005; Razze et al. 2016). For example, intercropping buckwheat ( Fagopyrum esculentum ) with yellow squash, in combination with white plastic mulch on beds, reduced adult whitefly abundance (LaTora et al. 2022). The timing of these practices should be adjusted according to regional cropping seasons and local whitefly activity.

Field sanitation is another important component of CuLCrV cultural management. Destroying plant material immediately after harvesting eliminates both the virus and the whiteflies surviving on crop debris (Sparks et al. 2018). Likewise, removing weed hosts in and around cucurbit fields during the off‐season can reduce virus reservoirs and limit early‐season spread (Adkins et al. 2011; Dhadly et al. 2025a).

4. Outlook and Practical Implications

The knowledge base surrounding CuLCrV has expanded considerably since its initial detection in the New World; however, critical gaps remain. A shift toward integrated, technology‐driven and sustainable strategies is needed to improve management of CuLCrV. Advances in molecular breeding and gene editing offer promising avenues for durable resistance. Marker‐assisted selection and genomic selection provide powerful tools for pyramiding virus resistance genes and whitefly resistance QTLs (Jaganathan et al. 2026). CRISPR (Clustered Regularly Interspaced Short Palindromic Repeats)/Cas systems can further complement modern breeding by enabling precise targeting of virus genome or host susceptibility genes to reduce virus accumulation, with inducible systems enhancing specificity (Ghorbani, Faal, et al. 2020; Pramanik et al. 2021). Gene editing also offers a means to overcome linkage drag from wild introgressions by uncoupling beneficial resistance loci from undesirable traits (Jaganathan et al. 2026). Transgenic strategies, including coat protein‐mediated resistance previously demonstrated in squash against aphid‐transmitted viruses (USDA 1994, 1996), provide a conceptual framework for developing CuLCrV‐resistant cultivars.

Emerging technologies have not yet been validated for CuLCrV, but they offer promising opportunities for future disease monitoring and control. Advances in diagnostics such as digital droplet PCR and CRISPR‐based platforms such as DETECTR and SHERLOCK enable highly sensitive detection, while portable sequencing tools such as the Oxford Nanopore MinION support rapid virus monitoring (Combala et al. 2025). Emerging in planta sensor technologies may further enable non‐destructive, real‐time pathogen detection (Singh et al. 2025). Artificial intelligence (AI) and machine learning are increasingly integrated into disease monitoring and decision‐support systems. For example, image‐based platforms can automate whitefly detection from field traps (Ciampi et al. 2023), while tools such as SquashRX (Malik and Dutta 2025) incorporate epidemiological and weather data to generate region‐specific WTV risk forecasts in South Georgia. AI can also support future CuLCrV research through precision agriculture, resistance screening and genome editing using tools such as AlphaFold and large language model‐assisted CRISPR systems (Jumper et al. 2021; Fang et al. 2025; Ruffolo et al. 2025). Although exogenous dsRNA has shown limited success against DNA viruses (Delgado‐Martín et al. 2022; Rego‐Machado et al. 2020), artificial microRNA strategies have shown promise (Vu et al. 2013), suggesting that RNA interference‐based approaches warrant further exploration for CuLCrV. Collectively, these emerging technologies may expand the toolbox available for CuLCrV management (Figure 4).

FIGURE 4.

FIGURE 4

Open in a new tab

Current and prospective management strategies for cucurbit leaf crumple virus under a systems approach. Early detection includes rapid virus detection and whitefly monitoring tools. Viral genomic surveillance focuses on monitoring virus diversity and evolution. Cultural and chemical control includes practises to reduce virus sources and whitefly populations. Breeding for virus resistance involves conventional and advanced approaches for developing resistant cultivars. Emerging technologies associated with each module are illustrated in the surrounding panels. (Created with BioRender).

Author Contributions

Dalvir Kaur Dhadly: writing – original draft, writing – review and editing, data curation, investigation, formal analysis, validation, visualization. Saritha Raman Kavalappara: writing – review and editing, resources, supervision, methodology, investigation, validation, visualization. Sudeep Bag: conceptualization, methodology, investigation, project administration, funding acquisition, writing – original draft, writing – review and editing, resources, supervision, data curation. Theodore McAvoy: writing – review and editing, resources. Paul M. Severns: writing – review and editing. Rajagopalbabu Srinivasan: writing – review and editing, resources. Alvin M. Simmons: writing – review and editing, funding acquisition, project administration.

Funding

This study was partially supported by the USDA‐HATCH grant awarded to S.B., number 1020319 and Agreement No. 59‐6080‐5‐001‐ Managing Whiteflies and Whitefly‐transmitted Viruses in Vegetable Crops in the Southeastern U.S., from the U.S. Department of Agriculture‐Agricultural Research Service. D.K.D., acknowledges the scholarship provided by the Department of Plant Pathology, University of Georgia, for her graduate program and UGA Graduate School Summer Research Grants 2024 for Doctoral Students. Any opinions, findings, conclusions or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the views of the U.S. Department of Agriculture and funding agency.

Conflicts of Interest

The authors declare no conflicts of interest.

Supporting information

Table S1: List of virus species used for phylogenetic analyses.

MPP-27-e70345-s002.docx (20.3KB, docx)

Table S2: List of modes of action recommended for whiteflies that are labelled for cucurbit and snap bean in the Southeastern USA.

MPP-27-e70345-s001.docx (16.3KB, docx)

Acknowledgements

S.B. wishes to thank Tian Tongyan for his invaluable support and assistance with electron microscopy.

Data Availability Statement

Data sharing not applicable to this article as no datasets were generated or analysed during the current study.

References

  1. Acharya, N. , Kumar M., Bag S., et al. 2025. “Prevalence of Aphid‐Transmitted Potyviruses in Pumpkin and Winter Squash in Georgia, USA.” Viruses 17, no. 2: 233. [DOI] [PMC free article] [PubMed] [Google Scholar]
  2. Acharya, N. , Kumar M., Bag S., et al. 2026. “Cultivars of Short‐Season Cucurbit Vegetable Crops (Yellow Squash and Zucchini) Exhibit Variable Tolerance to Whitefly‐Associated Silverleaf Disorder and Whitefly‐Transmitted Viruses in the Southeastern United States.” Plant Disease. 10.1094/PDIS-04-26-0730-RE. [DOI] [PubMed] [Google Scholar]
  3. Acharya, N. , McAvoy T., Bag S., and Riley D. G.. 2025. “Selecting Fall‐Season Cucurbit Crops for Whitefly‐Transmitted Disease Tolerance.” CAES Field Report C1350. https://fieldreport.caes.uga.edu/publications/C1350/selecting‐fall‐season‐cucurbit‐crops‐for‐whitefly‐transmitted‐disease‐tolerance.
  4. Adeleke, I. A. , Kavalappara S. R., McGregor C., Srinivasan R., and Bag S.. 2022. “Persistent and Asymptomatic Viral Infections and Whitefly‐Transmitted Viruses Impacting Cantaloupe and Watermelon in Georgia, USA.” Viruses 14, no. 6: 1310. [DOI] [PMC free article] [PubMed] [Google Scholar]
  5. Adkins, S. , Polston J. E., and Turechek W. W.. 2009. “Cucurbit Leaf Crumple Virus Identified in Common Bean in Florida.” Plant Disease 93, no. 3: 320. [DOI] [PubMed] [Google Scholar]
  6. Adkins, S. , Webb S. E., Baker C. A., and Kousik C. S.. 2008. “Squash Vein Yellowing Virus Detection Using Nested Polymerase Chain Reaction Demonstrates That the Cucurbit Weed Momordica charantia Is a Reservoir Host.” Plant Disease 92, no. 7: 1119–1123. [DOI] [PubMed] [Google Scholar]
  7. Adkins, S. , Webster C. G., Baker C. A., Weaver R., Rosskopf E. N., and Turechek W. W.. 2009. “Detection of Three Whitefly‐Transmitted Viruses Infecting the Cucurbit Weed Cucumis melo var. dudaim in Florida.” Plant Health Progress 10, no. 1: 39. [Google Scholar]
  8. Adkins, S. , Webster C. G., Kousik C. S., et al. 2011. “Ecology and Management of Whitefly‐Transmitted Viruses of Vegetable Crops in Florida.” Virus Research 159, no. 2: 110–114. [DOI] [PubMed] [Google Scholar]
  9. Agarwal, G. , Kavalappara S. R., Gautam S., et al. 2021. “Field Screen and Genotyping of Phaseolus vulgaris Against Two Begomoviruses in Georgia, USA.” Insects 12, no. 1: 49. [DOI] [PMC free article] [PubMed] [Google Scholar]
  10. Akad, F. , Webb S., Nyoike T. W., et al. 2008. “Detection of Cucurbit Leaf Crumple Virus in Florida Cucurbits.” Plant Disease 92: 648. [DOI] [PubMed] [Google Scholar]
  11. Amini, S. N. , Golizadeh A., Tafaghodinia B., et al. 2021. “Interaction Between Host Plant Morphological Characteristics With Life History of the Greenhouse Whitefly Trialeurodes vaporariorum .” Arthropod–Plant Interactions 15: 875–885. [Google Scholar]
  12. Barman, A. K. , Roberts P. M., Prostko E. P., and Toews M. D.. 2022. “Seasonal Occurrence and Reproductive Suitability of Weed Hosts for Sweetpotato Whitefly Bemisia tabaci (Hemiptera: Aleyrodidae) in South Georgia.” Journal of Entomological Science 57, no. 1: 1–11. [Google Scholar]
  13. Barman, A. K. , Toews M. D., and Roberts P. M.. 2023. “Sampling and Managing Whiteflies in Georgia Cotton.” UGA Cooperative Extension Circular 1184. University of Georgia Cooperative Extension. https://fieldreport.caes.uga.edu/publications/C1184/sampling‐and‐managing‐whiteflies‐in‐georgia‐cotton/.
  14. Boykin, L. M. 2014. “ Bemisia tabaci Nomenclature: Lessons Learned.” Pest Management Science 70, no. 10: 1454–1459. [DOI] [PubMed] [Google Scholar]
  15. Boykin, L. M. , Shatters R. G. Jr., Rosell R. C., et al. 2007. “Global Relationships of Bemisia tabaci (Hemiptera: Aleyrodidae) Revealed Using Bayesian Analysis of Mitochondrial COI DNA Sequences.” Molecular Phylogenetics and Evolution 44: 1306–1319. [DOI] [PubMed] [Google Scholar]
  16. Briddon, R. W. , Patil B. L., Bagewadi B., Nawaz‐ul‐Rehman M. S., and Fauquet C. M.. 2010. “Distinct Evolutionary Histories of the DNA‐A and DNA‐B Components of Bipartite Begomoviruses.” BMC Evolutionary Biology 10, no. 1: 97. [DOI] [PMC free article] [PubMed] [Google Scholar]
  17. Brown, J. K. , Coats S. A., Bedford I. D., Markham P. G., Bird J., and Frohlich D. R.. 1995. “Characterization and Distribution of Esterase Electromorphs in the Whitefly Bemisia tabaci (Genn.) (Homoptera: Aleyrodidae).” Biochemical Genetics 33, no. 7: 205–214. [DOI] [PubMed] [Google Scholar]
  18. Brown, J. K. , and Czosnek H.. 2002. “Whitefly Transmission of Plant Viruses.” In Advances in Botanical Research, vol. 36, 65–100. Press. [Google Scholar]
  19. Brown, J. K. , Idris A. M., Alteri C., and Stenger D. C.. 2002. “Emergence of a New Cucurbit‐Infecting Begomovirus Species Capable of Forming Viable Reassortants With Related Viruses in the Squash Leaf Curl Virus Cluster.” Phytopathology 92, no. 7: 734–742. [DOI] [PubMed] [Google Scholar]
  20. Brown, J. K. , Idris A. M., Olsen M. W., Miller M. E., Isakeit T., and Anciso J.. 2000. “Cucurbit Leaf Curl Virus, a New Whitefly Transmitted Geminivirus in Arizona, Texas, and Mexico.” Plant Disease 84: 809. [DOI] [PubMed] [Google Scholar]
  21. Brown, J. K. , Paredes‐Montero J. R., and Stocks I. C.. 2023. “The Bemisia tabaci Cryptic (Sibling) Species Group – Imperative for a Taxonomic Reassessment.” Current Opinion in Insect Science 57: 101032. [DOI] [PubMed] [Google Scholar]
  22. Caballero, R. , Cyman S., and Schuster D. J.. 2013. “Monitoring Insecticide Resistance in Biotype B of Bemisia tabaci (Hemiptera: Aleyrodidae) in Florida.” Florida Entomologist 96, no. 4: 1243–1256. [Google Scholar]
  23. Caballero, R. , Schuster D. J., Peres N. A., et al. 2015. “Effectiveness of Cyantraniliprole for Managing Bemisia tabaci (Hemiptera: Aleyrodidae) and Interfering With Transmission of Tomato Yellow Leaf Curl Virus on Tomato.” Journal of Economic Entomology 108, no. 3: 894–903. [DOI] [PubMed] [Google Scholar]
  24. Candian, J. S. , Coolong T., Dutta B., et al. 2021. “Yellow Squash and Zucchini Cultivar Selection for Resistance to Cucurbit Leaf Crumple Virus in the Southeastern United States.” HortTechnology 31, no. 4: 504–513. [Google Scholar]
  25. Castle, S. , Palumbo J., Merten P., Cowden C., and Prabhaker N.. 2017. “Effects of Foliar and Systemic Insecticides on Whitefly Transmission and Incidence of Cucurbit Yellow Stunting Disorder Virus.” Pest Management Science 73, no. 7: 1462–1472. [DOI] [PubMed] [Google Scholar]
  26. Castle, S. J. , Prabhaker N., Henneberry T. J., and Toscano N. C.. 2009. “Host Plant Influence on Susceptibility of Bemisia tabaci (Hemiptera: Aleyrodidae) to Insecticides.” Bulletin of Entomological Research 99, no. 3: 263–273. [DOI] [PubMed] [Google Scholar]
  27. Ciampi, L. , Zeni V., Incrocci L., et al. 2023. “A Deep Learning‐Based Pipeline for Whitefly Pest Abundance Estimation on Chromotropic Sticky Traps.” Ecological Informatics 78: 102384. [Google Scholar]
  28. Combala, M. , Tibiri E. B., Pita J. S., et al. 2025. “Improving the Diagnosis of Cassava Mosaic Begomoviruses Using Oxford Nanopore Technology Sequencing.” Scientific Reports 15: 41432. [DOI] [PMC free article] [PubMed] [Google Scholar]
  29. Czosnek, H. , Ghanim M., and Ghanim M.. 2002. “The Circulative Pathway of Begomoviruses in the Whitefly Vector Bemisia tabaci —Insights From Studies With Tomato Yellow Leaf Curl Virus.” Annals of Applied Biology 140, no. 3: 215–231. [Google Scholar]
  30. Dawson, K. 2016. Whitefly Populations Troubling Georgia Gegetable Growers. University of Georgia. https://vegetablegrowersnews.com/news/whitefly‐populations‐troubling‐georgia‐vegetable‐growers/. [Google Scholar]
  31. De Barro, P. J. 2011. “Chapter 9– Bemisia tabaci , the Capacity to Invade.” In The Whitefly, Bemisia tabaci (Homoptera: Aleyrodidae) Interaction With Geminivirus‐Infected Host Plants: Bemisia tabaci , Host Plants and Geminiviruses, edited by Thompson W., 181–204. Springer Netherlands. [Google Scholar]
  32. Delgado‐Martín, J. , Ruiz L., Janssen D., and Velasco L.. 2022. “Exogenous Application of dsRNA for the Control of Viruses in Cucurbits.” Frontiers in Plant Science 13: 895953. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Devendran, R. , Kavalappara S. R., Simmons A. M., and Bag S.. 2023. “Whitefly‐Transmitted Viruses of Cucurbits in the Southern United States.” Viruses 15, no. 11: 2278. [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Dhadly, D. K. , Kavalappara S. R., McAvoy T., et al. 2025a. “Weed Hosts: Hidden Reservoirs for Whitefly‐Transmitted Viruses Threatening Georgia Cucurbits.” Plant Disease 109, no. 11: 2352–2361. [DOI] [PubMed] [Google Scholar]
  35. Dhadly, D. K. , Kavalappara S. R., McAvoy T., et al. 2025b. “Cucurbit Leaf Crumple Virus Is Seed Transmitted in Yellow Squash ( Cucurbita pepo ).” Plant Disease 109, no. 1: 63–72. [DOI] [PubMed] [Google Scholar]
  36. Díaz‐Pérez, J. C. , McAvoy T., Luo X., and Bag S.. 2023. “Alternative Methods to Manage Silverleaf Whitefly [ Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae)] in Zucchini ( Cucurbita pepo L.).” Journal of Horticultural Research 31, no. 1: 61–70. [Google Scholar]
  37. Dimase, M. , De Marchi B. R., Lahiri S., et al. 2024. “First Report of Bemisia tabaci MEAM1 (Hemiptera: Aleyrodidae) Resistance to Spirotetramat in Florida.” Crop Protection 180: 106661. [Google Scholar]
  38. Dimase, M. , Rossitto De Marchi B. R., Barreto da Silva F., et al. 2024. “Monitoring the Susceptibility of Bemisia tabaci Middle East–Asia Minor 1 (Hemiptera: Aleyrodidae) to Afidopyropen, Cyantraniliprole, Dinotefuran, and Flupyradifurone in South Florida Vegetable Fields.” Journal of Economic Entomology 117, no. 4: 1606–1615. [DOI] [PMC free article] [PubMed] [Google Scholar]
  39. Dinsdale, A. , Cook L., Riginos C., Buckley Y. M., and Barro P. D.. 2010. “Refined Global Analysis of Bemisia tabaci (Hemiptera: Sternorrhyncha: Aleyrodoidea: Aleyrodidae) Mitochondrial Cytochrome Oxidase 1 to Identify Species‐Level Genetic Boundaries.” Annals of the Entomological Society of America 103, no. 2: 196–208. [Google Scholar]
  40. Do Prado, J. C. , Peñaflor M. F. G. V., Cia E., et al. 2016. “Resistance of Cotton Genotypes With Different Leaf Colour and Trichome Density to Bemisia tabaci Biotype B.” Journal of Applied Entomology 140: 405–413. [Google Scholar]
  41. Dutta, B. 2019. “Sida Golden Mosaic and Cucurbit Leaf Crumple Virus on Snap Beans.” University of Georgia Vegetable Blog 15: 357. https://site.caes.uga.edu/vegpath/2019/10/sida‐golden‐mosaic‐and‐cucurbit‐leaf‐crumple‐virus‐on‐snap‐beans/. [Google Scholar]
  42. Dutta, B. 2022. “Consideration for Variety Selection in Snap Bean Against Whitefly‐Transmitted Viruses.” University of Georgia Vegetable Blog 1567. https://site.caes.uga.edu/vegpath/2022/08/consideration‐for‐variety‐selection‐in‐snap‐bean‐against‐whitefly‐transmitted‐viruses/.
  43. Dutta, B. , Myers B., Coolong T., Srinivasan B., Sparks A. N., and Bag S.. 2024. “Whitefly‐Transmitted Plant Viruses in South Georgia.” University of Georgia Cooperative Extension Publication No. B 1507. https://extension.uga.edu/publications/detail.html?number=B1507.
  44. Fang, Z. , Ran H., Zhang Y., et al. 2025. “AlphaFold 3: An Unprecedented Opportunity for Fundamental Research and Drug Development.” Precision Clinical Medicine 8, no. 3: pbaf015. [DOI] [PMC free article] [PubMed] [Google Scholar]
  45. Fiallo‐Olivé, E. , Lett J. M., Martin D. P., et al. 2021. “ICTV Virus Taxonomy Profile: Geminiviridae 2021.” Journal of General Virology 102, no. 12: e001696. [DOI] [PMC free article] [PubMed] [Google Scholar]
  46. Fiallo‐Olivé, E. , and Navas‐Castillo J.. 2023. “Begomoviruses: What Is the Secret(s) of Their Success?” Trends in Plant Science 28, no. 6: 715–727. [DOI] [PubMed] [Google Scholar]
  47. Fondong, V. N. 2025. “DNA Satellites Impact Begomovirus Diseases in a Virus‐Specific Manner.” International Journal of Molecular Sciences 26, no. 12: 5814. [DOI] [PMC free article] [PubMed] [Google Scholar]
  48. Frank, D. L. , and Liburd O. E.. 2005. “Effects of Living and Synthetic Mulch on the Population Dynamics of Whiteflies and Aphids, Their Associated Natural Enemies, and Insect‐Transmitted Plant Diseases in Zucchini.” Environmental Entomology 34: 857–865. [Google Scholar]
  49. Gadhave, K. R. , Gautam S., Dutta B., Coolong T., Adkins S., and Srinivasan R.. 2020. “Low Frequency of Horizontal and Vertical Transmission of Cucurbit Leaf Crumple Virus in Whitefly Bemisia tabaci Gennadius.” Phytopathology 110, no. 6: 1235–1241. [DOI] [PubMed] [Google Scholar]
  50. Gautam, S. 2019. “The Role of Bemisia tabaci in the Transmission of Vegetable Viruses in the Farmscape of Georgia.” Doctoral dissertation.
  51. Gautam, S. , Buck J. W., Dutta B., et al. 2023. “Sida Golden Mosaic Virus, an Emerging Pathogen of Snap Bean ( Phaseolus vulgaris L.) in the Southeastern United States.” Viruses 15: 357. [DOI] [PMC free article] [PubMed] [Google Scholar]
  52. Gautam, S. , Crossley M. S., Dutta B., et al. 2020. “Low Genetic Variability in Bemisia tabaci MEAM1 Populations Within Farmscapes of Georgia, USA.” Insects 11, no. 12: 834. [DOI] [PMC free article] [PubMed] [Google Scholar]
  53. Gautam, S. , Mugerwa H., Buck J. W., et al. 2022. “Differential Transmission of Old and New World Begomoviruses by Middle East–Asia Minor 1 (MEAM1) and Mediterranean (MED) Cryptic Species of Bemisia tabaci .” Viruses 14, no. 5: 1104. [DOI] [PMC free article] [PubMed] [Google Scholar]
  54. Georgia Pest Management Handbook: Commercial Edition – Volume 1 . 2024. “University of Georgia Cooperative Extension.” https://extension.uga.edu/programs‐services/integrated‐pest‐management/publications.html.
  55. Ghorbani, Faal, P. , Farsi M., Seifi A., et al. 2020. “Virus‐Induced CRISPR‐Cas9 System Improved Resistance Against Tomato Yellow Leaf Curl Virus.” Molecular Biology Reports 47: 3369–3376. [DOI] [PubMed] [Google Scholar]
  56. Ghosh, S. , and Ghanim M.. 2021. “Factors Determining Transmission of Persistent Viruses by Bemisia tabaci and Emergence of New Virus–Vector Relationships.” Viruses 13, no. 9: 1808. [DOI] [PMC free article] [PubMed] [Google Scholar]
  57. Gilbertson, R. 2017. “Cucurbit Leaf Crumple.” In Compendium of Cucurbit Diseases and Pests, edited by Keinath A., Wintermantel W., and Zitter T., Second ed., 111–113. APS Press. [Google Scholar]
  58. Guzman, P. , Sudarshana M. R., Seo Y.‐S., et al. 2000. “A New Bipartite Geminivirus (Begomovirus) Causing Leaf Curl and Crumpling in Cucurbits in the Imperial Valley of California.” Plant Disease 84: 488. [DOI] [PubMed] [Google Scholar]
  59. Hagen, C. , Rojas M. R., Kon T., and Gilbertson R. L.. 2008. “Recovery From Cucurbit Leaf Crumple Virus (Family Geminiviridae, Genus Begomovirus) Infection Is an Adaptive Antiviral Response Associated With Changes in Viral Small RNAs.” Phytopathology 98, no. 9: 1029–1037. [DOI] [PubMed] [Google Scholar]
  60. Hagen, C. , Rojas M. R., Sudarshana M. R., et al. 2008. “Biology and Molecular Characterization of Cucurbit Leaf Crumple Virus, an Emergent Cucurbit‐Infecting Begomovirus in the Imperial Valley of California.” Plant Disease 92: 781–793. [DOI] [PubMed] [Google Scholar]
  61. Hendricks, K. E. , and Roberts P. D.. 2023. “Distribution and Incidence of Watermelon Crinkle Leaf‐Associated Virus 1, Watermelon Crinkle Leaf‐Associated Virus 2, and Other Viruses in Cucurbits in South Florida.” Plant Health Progress 24, no. 4: 488–497. [Google Scholar]
  62. Hilje, L. , and Stansly P. A.. 2008. “Living Ground Covers for Management of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae) and Tomato Yellow Mottle Virus (ToYMoV) in Costa Rica.” Crop Protection 27: 10–16. [Google Scholar]
  63. Hull, R. 2014. “Chapter 6—Genome Composition and Expression.” In Plant Virology, 5th ed., 319–320. Academic Press. [Google Scholar]
  64. ICTV . 2025. “Genus: Begomovirus. ICTV Report.” https://ictv.global/report/chapter/geminiviridae/geminiviridae/begomovirus.
  65. Jaganathan, D. , Dutta B., Basu S., et al. 2026. “Breeding Vegetables for Whitefly Resistance: Past, Present, and Future in the AI Era.” Frontiers in Plant Science 16: 1724403. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Jailani, A. A. K. , Hendricks K., Roberts P. D., and Paret M. L.. 2021. “Development of a Simple One‐Step Multiplex RT‐PCR System for Simultaneous Detection of DNA and RNA Viruses of Cucurbit Leaf Crumple Virus, Cucurbit Yellow Stunting Disorder Virus, Squash Vein Yellowing Virus, and Cucurbit Chlorotic Yellows Virus.” Physiological and Molecular Plant Pathology 116: 101734. [Google Scholar]
  67. Jailani, A. A. K. , and Paret M. L.. 2023. “Development of a Multiplex RT‐RPA Assay for Simultaneous Detection of Three Viruses in Cucurbits.” Molecular Plant Pathology 24, no. 11: 1443–1450. [DOI] [PMC free article] [PubMed] [Google Scholar]
  68. Jailani, A. A. K. , and Paret M. L.. 2024. “A Multiplex RT‐qPCR Assay for Simultaneous Detection of Cucurbit Viruses From Individual Whitefly and Plant Samples.” Plant Disease 108, no. 12: 3508–3517. [DOI] [PubMed] [Google Scholar]
  69. Jailani, A. A. K. , and Paret M. L.. 2025. “First Report of On‐Site Detection of Cucurbit Leaf Crumple Virus by an Optimized RPA–Lateral Flow Assay With an Alternative Endonuclease.” International Journal of Molecular Sciences 26, no. 21: 10611. [DOI] [PMC free article] [PubMed] [Google Scholar]
  70. Jeske, H. , Lütgemeier M., and Preiß W.. 2001. “DNA Forms Indicate Rolling Circle and Recombination‐Dependent Replication of Abutilon Mosaic Virus.” EMBO Journal 20: 6158–6167. [DOI] [PMC free article] [PubMed] [Google Scholar]
  71. Johnston, N. , Paris T., Paret M., Freeman J., and Martini X.. 2022. “Repelling Whitefly ( Bemisia tabaci ) Using Limonene‐Scented Kaolin: A Novel Pest Management Strategy.” Crop Protection 155: 105905. [Google Scholar]
  72. Jumper, J. , Evans R., Pritzel A., et al. 2021. “Highly Accurate Protein Structure Prediction With AlphaFold.” Nature 596, no. 7873: 583–589. [DOI] [PMC free article] [PubMed] [Google Scholar]
  73. Kalischuk, M. , Hendricks K., Hochmuth R., Freeman J., Roberts P. D., and Paret M.. 2022. “A Portable Recombinase Polymerase Amplification Assay for the Rapid Detection of Cucurbit Leaf Crumple Virus in Watermelon Leaves and Fruits.” Journal of Plant Pathology 104, no. 1: 215–224. [Google Scholar]
  74. Kavalappara, S. R. , Acharya N., Suarez E., McAvoy T., and Bag S.. 2024. “First Report of Watermelon Crinkle Leaf‐Associated Virus‐2 (WCLaV‐2) in Watermelon ( Citrullus lanatus ) in Georgia, U.S.A.” Plant Disease 108: 2246. [Google Scholar]
  75. Kavalappara, S. R. , Devendran R., Simmons A. M., and Bag S.. 2024. “Microneedle Assisted Delivery of the Cloned Components of Cucurbit Leaf Crumple Virus in Yellow Squash ( Cucurbita pepo ).” Journal of Virological Methods 329: 114992. [DOI] [PubMed] [Google Scholar]
  76. Kavalappara, S. R. , Milner H., Konakalla N. C., et al. 2021. “High Throughput Sequencing‐Aided Survey Reveals Widespread Mixed Infections of Whitefly‐Transmitted Viruses in Cucurbits in Georgia, USA.” Viruses 13, no. 6: 988. [DOI] [PMC free article] [PubMed] [Google Scholar]
  77. Keinath, A. P. , and Ling K.‐S.. 2023. “Cucurbit Leaf Crumple Virus (CuLCrV).” Emerging Viruses in Cucurbits Working Group Publication No 2301. https://ecucurbitviruses.org/wp‐content/uploads/2023/05/CuLCrV‐April‐2023.pdf.
  78. Keinath, A. P. , Ling K.‐S., Adkins S., et al. 2018. “First Report of Cucurbit Leaf Crumple Virus Infecting Three Cucurbit Crops in South Carolina.” Plant Health Progress 19: 322–323. [Google Scholar]
  79. Kousik, C. S. , and Adkins S.. 2020. “Detection of Cucurbit Yellow Stunting Disorder Virus Infecting Watermelon in South Carolina.” Plant Health Progress 21, no. 2: 133–134. [Google Scholar]
  80. Kumar, S. , Stecher G., Li M., Knyaz C., and Tamura K.. 2018. “MEGA X: Molecular Evolutionary Genetics Analysis Across Computing Platforms.” Molecular Biology and Evolution 35: 1547–1549. [DOI] [PMC free article] [PubMed] [Google Scholar]
  81. Larsen, R. , and Kmiecik K.. 2010. “First Report of Cucurbit Leaf Crumple Virus in Snap Bean in Georgia.” Phytopathology 100: S68. [Google Scholar]
  82. LaTora, A. G. , Codod C. B., Legarrea S., et al. 2022. “Combining Cultural Tactics and Insecticides for the Management of the Sweetpotato Whitefly, Bemisia tabaci MEAM1, and Viruses in Yellow Squash.” Horticulturae 8, no. 4: 341. [Google Scholar]
  83. Liang, G. , and Liu T. X.. 2009. “Repellency of a Kaolin Particle Film, Surround, and a Mineral Oil, Sunspray Oil, to Silverleaf Whitefly (Homoptera: Aleyrodidae) on Melon in the Laboratory.” Journal of Economic Entomology 95: 317–324. [DOI] [PubMed] [Google Scholar]
  84. Luckew, A. , Meru G., Wang Y. Y., et al. 2022. “Field Evaluation of Cucurbita Germplasm for Resistance to Whiteflies and Whitefly‐Transmitted Viruses.” HortScience 57, no. 2: 337–344. [Google Scholar]
  85. Luckew, A. , Sari N., Pandey S., et al. 2025. “Watermelon Germplasm With Resistance to Whitefly‐Transmitted Viruses.” HortScience 60, no. 2: 245–253. [Google Scholar]
  86. Malik, A. A. , and Dutta B.. 2025. “Squashrx: A Decision‐Support Tool for Managing Whitefly‐Transmitted Viruses in Squash in Georgia.” Integrated Pest Management. https://ipm.uga.edu/2025/10/02/squashrx‐app/. [DOI] [PubMed]
  87. Martin, D. P. , Varsani A., Roumagnac P., et al. 2021. “RDP5: A Computer Program for Analyzing Recombination in, and Removing Signals of Recombination From, Nucleotide Sequence Datasets.” Virus Evolution 7, no. 1: veaa087. [DOI] [PMC free article] [PubMed] [Google Scholar]
  88. Martini, X. , Paret M., and Freeman J.. 2016. “Early Outbreaks of Sweetpotato Whiteflies.” Citrus Industry 11: 26–27. https://www.researchgate.net/publication/310749519_Early_outbreaks_of_sweetpotato_whiteflies. [Google Scholar]
  89. Martini, X. , Paret M., Polston J. E., et al. 2022. “Recommendations for Management of Whiteflies, Whitefly‐Transmitted Viruses, and Insecticide Resistance for Production of Cucurbit Crops in Florida.” University of Florida IFAS Extension EDIS Publication IN871. https://edis.ifas.ufl.edu/publication/IN871.
  90. McAvoy, G. 2017. “Take Caution to Curb Cucurbit Leaf Crumple Virus.” https://www.growingproduce.com/crop‐protection/disease‐control/take‐caution‐to‐curb‐cucurbit‐leaf‐crumple‐virus/.
  91. McCreight, J. D. , Liu H., and Turini T. A.. 2008. “Genetic Resistance to Cucurbit Leaf Crumple Virus in Melon.” HortScience 43, no. 1: 122–126. [Google Scholar]
  92. Mondal, S. , Hladky L. J., and Wintermantel W. M.. 2023. “Differential Seasonal Prevalence of Yellowing Viruses Infecting Melon Crops in Southern California and Arizona Determined by Multiplex RT‐PCR and RT‐qPCR.” Plant Disease 107, no. 9: 2653–2664. [DOI] [PubMed] [Google Scholar]
  93. Moury, B. , Fabre F., Hébrard E., and Froissart R.. 2017. “Determinants of Host Species Range in Plant Viruses.” Journal of General Virology 98: 862–873. [DOI] [PubMed] [Google Scholar]
  94. Nauen, R. , Wölfel K., Lueke B., et al. 2015. “Development of a Lateral Flow Test to Detect Metabolic Resistance in Bemisia tabaci Mediated by CYP6CM1, a Cytochrome P450 With Broad Spectrum Catalytic Efficiency.” Pesticide Biochemistry and Physiology 121: 3–11. [DOI] [PubMed] [Google Scholar]
  95. Navas‐Castillo, J. , Fiallo‐Olivé E., and Sánchez‐Campos S.. 2011. “Emerging Virus Diseases Transmitted by Whiteflies.” Annual Review of Phytopathology 49: 219–248. [DOI] [PubMed] [Google Scholar]
  96. Nyoike, T. W. , and Liburd O. E.. 2008. “Application of Imidacloprid and Cultural Techniques for Management of Whiteflies in Cucurbits.” Proceedings of the Florida State Horticultural Society 121: 249–251. [Google Scholar]
  97. Nyoike, T. W. , and Liburd O. E.. 2010. “Effect of Living (Buckwheat) and UV Reflective Mulches With and Without Imidacloprid on Whiteflies, Aphids, and Marketable Yields of Zucchini Squash.” International Journal of Pest Management 56, no. 1: 31–39. [Google Scholar]
  98. Nyoike, T. W. , Liburd O. E., and Webb S. E.. 2008. “Suppression of Whiteflies, Bemisia tabaci (Hemiptera: Aleyrodidae), and Incidence of Cucurbit Leaf Crumple Virus, a Whitefly‐Transmitted Virus of Zucchini Squash New to Florida, With Mulches and Imidacloprid.” Florida Entomologist 91, no. 3: 460–465. [Google Scholar]
  99. Oliveira, C. , Hoffmann L., Toscano L., et al. 2021. “Resistance of Cotton Genotypes to Silverleaf Whitefly ( Bemisia tabaci [Gennadius] Biotype B).” International Journal of Tropical Insect Science 41: 1697–1707. [Google Scholar]
  100. Oriani, M. A. D. G. , and Vendramim J. D.. 2010. “Influence of Trichomes on Attractiveness and Ovipositional Preference of Bemisia tabaci (Gennadius) B Biotype (Hemiptera: Aleyrodidae) on Tomato Genotypes.” Neotropical Entomology 39: 1002–1007. [DOI] [PubMed] [Google Scholar]
  101. Perier, J. D. , Cremonez P. S., Parkins A. J., Kheirodin A., Simmons A. M., and Riley D. G.. 2024. “Modified Maximum Dose Bioassay for Assessing Insecticide Response in Field Populations of Bemisia tabaci (Hemiptera: Aleyrodidae).” Journal of Entomological Science 59, no. 4: 475–487. [Google Scholar]
  102. Perier, J. D. , Cremonez P. S., Smith H. A., Simmons A. M., and Riley D. G.. 2024. “Susceptibility of Bemisia tabaci (Hemiptera: Aleyrodidae) Adult Populations to Imidacloprid in Georgia, USA.” Journal of Entomological Science 59, no. 2: 182–192. [Google Scholar]
  103. Perring, T. M. 2001. “The Bemisia tabaci Species Complex.” Crop Protection 20, no. 9: 725–737. [Google Scholar]
  104. Pramanik, D. , Shelake R. M., Park J., et al. 2021. “CRISPR/Cas9‐Mediated Generation of Pathogen‐Resistant Tomato Against Tomato Yellow Leaf Curl Virus and Powdery Mildew.” International Journal of Molecular Sciences 22: 1878. [DOI] [PMC free article] [PubMed] [Google Scholar]
  105. Rahman, M. A. K. , Rahmat Z., Iqbal M. A., and Zafar Y.. 2017. “Genetics and Genomics of Cotton Leaf Curl Disease, Its Viral Causal Agents and Whitefly Vector: A Way Forward to Sustain Cotton Fiber Security.” Frontiers in Plant Science 8: 1157. [DOI] [PMC free article] [PubMed] [Google Scholar]
  106. Razze, J. M. , Liburd O. E., and McSorley R.. 2016. “Preference of Bemisia tabaci Biotype B on Zucchini Squash and Buckwheat and the Effect of Delphastus catalinae on Whitefly Populations.” Pest Management Science 72: 1335–1339. [DOI] [PubMed] [Google Scholar]
  107. Rego‐Machado, C. M. , Nakasu E., Silva J., Lucinda N., Nagata T., and Inoue‐Nagata A. K.. 2020. “siRNA Biogenesis and Advances in Topically Applied dsRNA for Controlling Virus Infections in Tomato Plants.” Scientific Reports 10: 22277. [DOI] [PMC free article] [PubMed] [Google Scholar]
  108. Riley, D. , Batal D., and Wolff D.. 2001. “Resistance in Glabrous‐Type Cucumis melo L. to Whiteflies (Homoptera: Aleyrodidae).” Journal of Entomological Science 36: 46–56. 10.18474/0749-8004-36.1.46. [DOI] [Google Scholar]
  109. Rodríguez‐Negrete, E. A. , Jordan‐Ramírez R., Leyva‐López N. E., and Méndez‐Lozano J.. 2021. “Cucurbit Leaf Crumple Virus Associated to Watermelon Disease in Campeche State, Mexico.” Plant Disease 105, no. 4: 1235. [Google Scholar]
  110. Rojas, M. R. , Hagen C., Lucas W. J., and Gilbertson R. L.. 2005. “Exploiting Chinks in the Plant's Armor: Evolution and Emergence of Geminiviruses.” Annual Review of Phytopathology 43, no. 1: 361–394. [DOI] [PubMed] [Google Scholar]
  111. Rossitto De Marchi, B. , Smith H., Turechek W., and Riley D.. 2021. “A Maximum Dose Bioassay to Assess Efficacy of Key Insecticides Against Bemisia tabaci MEAM1 (Hemiptera: Aleyrodidae).” Journal of Economic Entomology 11, no. 2: 914–921. [DOI] [PubMed] [Google Scholar]
  112. Ruffolo, J. A. , Nayfach S., Gallagher J., et al. 2025. “Design of Highly Functional Genome Editors by Modelling CRISPR–Cas Sequences.” Nature 645, no. 8080: 518–525. [DOI] [PMC free article] [PubMed] [Google Scholar]
  113. Schuster, D. , Spurgeon S., and Riley D.. 2008. “Imidacloprid Resistance Monitoring and Management in Whitefly.” In Proceedings of the 2008 Southeastern Regional Vegetable Conference, Savannah, GA, edited by Kelley W. T., 702. University of Georgia. [Google Scholar]
  114. Shi, P. , Liu J., Ye J., et al. 2023. “Population Changes of Bemisia tabaci (Hemiptera: Aleyrodidae) on Different Colored Poinsettia Leaves With Different Trichome Densities and Chemical Compositions.” Journal of Economic Entomology 116: 1276–1285. [DOI] [PubMed] [Google Scholar]
  115. Simmons, A. M. , Jarret R. L., Cantrell C. L., and Levi A.. 2019. “ Citrullus ecirrhosus: Wild Source of Resistance Against Bemisia tabaci (Hemiptera: Aleyrodidae) for Cultivated Watermelon.” Journal of Economic Entomology 112, no. 5: 2425–2432. [DOI] [PubMed] [Google Scholar]
  116. Simmons, A. M. , and Levi A.. 2001. “Sources of Whitefly (Homoptera: Aleyrodidae) Resistance in Citrullus for the Improvement of Cultivated Watermelon.” HortScience 37, no. 3: 581–584. [Google Scholar]
  117. Singh, N. , Zhang Q., Xu W., Whitham S. A., and Dong L.. 2025. “A Biohydrogel‐Enabled Microneedle Sensor for in Situ Monitoring of Reactive Oxygen Species in Plants.” ACS Sensors 10: 1797–1810. [DOI] [PMC free article] [PubMed] [Google Scholar]
  118. Smith, H. A. , Nagle C. A., MacVean C. A., and McKenzie C. L.. 2016. “Susceptibility of Bemisia tabaci MEAM1 (Hemiptera: Aleyrodidae) to Imidacloprid, Thiamethoxam, Dinotefuran, and Flupyradifurone in South Florida.” Insects 7, no. 4: 57. [DOI] [PMC free article] [PubMed] [Google Scholar]
  119. Snyder, J. , Simmons A., and Thacker R.. 1998. “Attractancy and Ovipositional Response of Adult Bemisia argentifolii (Homoptera: Aleyrodidae) to Type IV Trichome Density on Leaves of Lycopersicon hirsutum Grown in Three Day‐Length Regimes.” Journal of Entomological Science 33: 270–281. [Google Scholar]
  120. Southeastern Vegetable Extension Workers Group . 2026. Southeastern U.S. Vegetable Crop Handbook. 27th ed. University of Tennessee. https://vegetables.tennessee.edu/southeastern‐vegetable‐crop‐handbook/. [Google Scholar]
  121. Sparks, A. , Roberts P., Barman A., Riley D., and Toews M.. 2018. “Cross‐Commodity Management of Silverleaf Whitefly in Georgia.” University of Georgia Cooperative Extension Publication No. C 1141. https://extension.uga.edu/publications/detail.html?number=C1141.
  122. Srinivasan, R. 2025. “Begomovirus cucurbitae. CABI Compendium.” 10.1079/cabicompendium.41201948. [DOI]
  123. Tamura, K. , and Nei M.. 1993. “Estimation of the Number of Nucleotide Substitutions in the Control Region of Mitochondrial DNA in Humans and Chimpanzees.” Molecular Biology and Evolution 10: 512–526. [DOI] [PubMed] [Google Scholar]
  124. USDA . 1994. “Environmental Assessment for Upjohn Company/Asgrow Seed Company Petition for Determination of Non‐Regulated Status for ZW‐20 Squash.” U.S. Department of Agriculture.
  125. USDA . 1996. “Environmental Assessment for Upjohn Company/Asgrow Seed Company Petition for Determination of Non‐Regulated Status for CZW‐3 Squash.” U.S. Department of Agriculture.
  126. Vu, T. V. , Choudhury N. R., and Mukherjee S. K.. 2013. “Transgenic Tomato Plants Expressing Artificial microRNAs for Silencing the Pre‐Coat and Coat Proteins of a Begomovirus, Tomato Leaf Curl New Delhi Virus, Show Tolerance to Virus Infection.” Virus Research 172: 35–45. [DOI] [PubMed] [Google Scholar]
  127. Waliullah, S. , Ling K.‐S., Cieniewicz E. J., Oliver J. E., Ji P., and Ali M. E.. 2020. “Development of Loop‐Mediated Isothermal Amplification Assay for Rapid Detection of Cucurbit Leaf Crumple Virus.” International Journal of Molecular Sciences 21, no. 5: 1756. [DOI] [PMC free article] [PubMed] [Google Scholar]
  128. Wan, F. , Zhang G., Liu S., et al. 2009. “Invasive Mechanism and Management Strategy of Bemisia tabaci (Gennadius) Biotype B: Progress Report of 973 Program on Invasive Alien Species in China.” Science in China. Series C, Life Sciences 52, no. 1: 88–95. [DOI] [PubMed] [Google Scholar]
  129. Wang, X. , Li P., and Liu S.. 2017. “Whitefly Interactions With Plants.” Current Opinion in Insect Science 19: 70–75. [DOI] [PubMed] [Google Scholar]
  130. Whittaker, J. 2019. “Whiteflies Bugging Vegetable and Cotton Producers.” https://www.gfb.org/special‐blog/post/whiteflies‐bugging‐vegetable‐cotton‐producers.

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Table S1: List of virus species used for phylogenetic analyses.

MPP-27-e70345-s002.docx (20.3KB, docx)

Table S2: List of modes of action recommended for whiteflies that are labelled for cucurbit and snap bean in the Southeastern USA.

MPP-27-e70345-s001.docx (16.3KB, docx)

Data Availability Statement

Data sharing not applicable to this article as no datasets were generated or analysed during the current study.

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