This datasheet covers the four bacterial species and pathovars that are associated with the bacterial spot of tomato and pepper: Xanthomonas euvesicatoria pv. euvesicatoria, Xanthomonas euvesicatoria pv. perforans, Xanthomonas hortorum pv. gardneri, Xanthomonas vesicatoria.

Preferred name: Xanthomonas vesicatoria
Authority: (Doidge) Vauterin, Hoste, Kersters & Swings
Taxonomic position: Bacteria: Proteobacteria: Gammaproteobacteria: Lysobacterales: Lysobacteraceae
Other scientific names: Pseudomonas exitiosa Gardner & Kendrick, Pseudomonas vesicatoria (Doidge) Stevens
Common names in English: bacterial leaf spot of pepper, bacterial leaf spot of tomato, bacterial scab of tomato, bacterial spot of pepper, black spot of tomato, leaf spot of tomato, stem canker of tomato
view more common names online...
Notes on taxonomy and nomenclature EPPO Categorization: A2 list
EU Categorization: RNQP (Annex IV)
view more categorizations online...
EPPO Code: XANTVE

The main hosts of bacterial spot xanthomonads are tomato (Solanum lycopersicum) and pepper (Capsicum spp.). X. euvesicatoria pv. euvesicatoria and X. hortorum pv. gardneri are reported as pathogens for both tomato and pepper. Meanwhile, X. vesicatoria primarily infects tomato and, until recently, X. euvesicatoria pv. perforans strains had only been isolated from tomato (Timilsina et al., 2015). However, over in recent years X. euvesicatoria pv. perforans has been isolated from pepper fields in Florida and Alabama (USA) (Potnis et al., 2015; Newberry et al., 2019).

Three pathotypes of strains have been distinguished among the bacterial spot causative agents: those exclusively infecting tomato (T races), those exclusively affecting pepper (P races), and those infecting both tomato and pepper. Several races have been identified based on the hypersensitive reaction (HR) triggered by effector proteins in Xanthomonas strains delivered via the type III secretion system into host cells and recognition by specific resistance proteins in tomato or pepper (Stall et al., 2009). Currently 11 pepper races and five tomato races have been documented (Bouzar et al., 1994; Stall et al., 2009; Adhikari et al., 2020; Jibrin et al., 2022). Pathogenic races are determined by the presence or absence of HR in the susceptible C. annuum cultivar Early Calwonder (ECW), its near-isogenic lines and Capsicum pubescens PI235047, or in different S. lycopersicum genetic lines. Nevertheless, future studies may reveal unknown races and potential novel dynamics in pathogen-host interaction.

A range of solanaceous and non-solanaceous plants, mainly weeds, have also been recorded as incidental hosts (Osdaghi et al., 2021).

Host list: Capsicum annuum, Capsicum baccatum, Capsicum chinense, Capsicum pubescens, Solanum lycopersicum, Tubocapsicum anomalum

Bacterial spot causative agents occur widely in tomato and pepper-growing areas, especially in tropical and subtropical regions with moderate or higher rainfall. The disease has primarily been observed in field crops but can also occur in greenhouses. The environmental conditions in Southern Europe are particularly favorable for disease expression in the field, given that the optimal growth temperature for xanthomonads is between 25 and 30°C (Holt, 1994).

The classification of the four lineages of bacterial spot xanthomonads has experienced several revisions due to the intricate taxonomic relationships within this group (see Notes on taxonomy and nomenclature). This has led to uncertainties regarding the specific species present in various geographical areas. As reported by Timilsina et al. (2015), shifts in the species composition of bacterial spot pathogens populations have occurred due to the global spread of dominant genotypes, and recombination between species has generated genetic diversity in these populations. Therefore, the global distribution and genetic diversity of bacterial spot xanthomonads are poorly understood. Strains belonging to X. euvesicatoria pv. euvesicatoria and X. vesicatoria were historically considered the dominant bacterial lineages with a worldwide distribution (Jones et al., 2004b). However, more recently, X. euvesicatoria pv. perforans and X. hortorum pv. gardneri were increasingly isolated in North and South America, the Middle East, East Africa, and regions bordering the Indian Ocean (Vancheva et al., 2021). Dramatic changes in the dominant lineages and population structure of bacterial spot in different local areas have been documented over the past few decades. For example, prior to 1991, X. euvesicatoria pv. euvesicatoria was the only species isolated from tomato in Florida (USA), while this taxon has been entirely replaced by X. euvesicatoria pv. perforans on this host since then (Klein-Gordon et al., 2021). Similar changes are also reported in Taiwan (Burlakoti et al., 2018).

The map includes only records where the Xanthomonas species has been identified. Articles mentioning 'Xanthomonas spp.’ or 'bacterial spot of tomato’ without mentioning the species are not included in the database. A map showing the records of Xanthomonas axonopodis pv. vesicatoria is available as an archive in EPPO Global Database.

EPPO Region: Austria, Azerbaijan, Belarus, Bulgaria, Czech Republic, France (mainland), Greece (mainland), Hungary, Israel, Italy (mainland, Sardegna, Sicilia), Kazakhstan, Morocco, Poland, Romania, Russia (Central Russia, Southern Russia, Western Siberia), Serbia, Slovakia, Slovenia, Spain (mainland), Tunisia, Türkiye
Africa: Egypt, Ethiopia, Ghana, Kenya, Madagascar, Malawi, Morocco, Mozambique, Niger, Nigeria, Reunion, Senegal, Seychelles, South Africa, Tanzania, Togo, Tunisia, Zambia, Zimbabwe
Asia: India (Karnataka), Israel, Kazakhstan, Nepal, Pakistan, Philippines, Taiwan
North America: Canada (Ontario), Mexico, United States of America (Arizona, California, Georgia, Hawaii, Iowa, Michigan, New Mexico, Ohio, Oklahoma)
Central America and Caribbean: Antigua and Barbuda, Barbados, Bermuda, Costa Rica, Cuba, Dominica, Dominican Republic, El Salvador, Grenada, Guadeloupe, Guatemala, Honduras, Jamaica, Martinique, Puerto Rico, Saint Lucia, St Kitts-Nevis, St Vincent and the Grenadines, Trinidad and Tobago, Virgin Islands (US)
South America: Argentina, Brazil (Bahia, Goias, Minas Gerais, Pernambuco, Rio de Janeiro, Santa Catarina, Sao Paulo), Chile, Colombia, Paraguay, Suriname, Uruguay, Venezuela
Oceania: Australia (New South Wales, Queensland), Fiji, Micronesia, New Caledonia, New Zealand, Palau, Tonga

The primary source of inoculum for bacterial spot xanthomonads are infected seeds and planting material (Potnis et al., 2015). Bacteria can survive from one season to another, mainly on tomato and pepper seeds, for at least 10 years (Bashan et al., 1982b). A positive correlation was observed between the inoculum concentration of the bacterium on pepper blossoms and the percentage of infested seeds (Dutta et al., 2013). These pathogens can also survive in infected debris and the soil to some extent, possibly in the rhizosphere of non-host plants (Bashan et al., 1982a). Diverse solanaceous and non-solanaceous weeds may be alternative hosts (Osdaghi et al., 2021).

Once introduced in an area, the spread of bacteria is primarily by rain-splash or overhead irrigation, with the handling of young plants also playing an important role (Goode & Sasser, 1980). Viable bacteria have been detected in aerosols over commercial fields, indicating a putative aerial dispersal (McInnes et al., 1988; Bernal & Berger, 1996 ). Leaves are infected through stomata, while fruits are infected through small wounds, such as abrasions, and insect punctures. Young leaves and fruits are more sensitive to infection and the bacteria can multiply epiphytically on young plants in the absence of symptoms. Thinning of directly seeded tomato seedlings (practice of selectively removing weaker seedlings to improve the health and productivity of the plants) is reported to promote spread of the disease, and it is recommended to thin in the afternoon, when plants are dry, and to use prophylactic hand washes (Pohronezny et al., 1990). The disease is favored by heavy rainfall, high humidity (Diab et al., 1982) and temperatures between 25°C and 30°C (EFSA, 2014).

Symptoms

All bacterial spot xanthomonads can induce a wide variety of symptoms on their host plants, including angular lesions that later become brown and necrotic on the leaves, fruits, petioles and stems. Some symptoms may be mistaken for those caused by other organisms (see below). Pathogen aggressiveness and the development of symptoms often depend on the host-pathogen combination, with X. euvesicatoria pv. euvesicatoria appearing to be significantly more aggressive on bell pepper than on tomato (Ignjatov et al., 2010).

Fruits of tomato show superficial corky spots or scabs, with water-soaked margins, oval or irregular in shape and with a diameter of around 2-10 mm. The ‘flecks’ caused by Pseudomonas syringae pv. tomato are distinctly smaller (diameter <1 mm), black, circular and elevated. Differences are also observed with symptoms of Clavibacter michiganensis on fruits, consisting of brown spots, slightly raised, and surrounded by a white halo (a distinctive 'bird's eye' appearance). Scabbing of fruit (but without water-soaking) may also be a symptom of the phytotoxicity of plant protection products. Bacterial spot xanthomonads fruit lesions in pepper are scab-like, raised and rapidly necrotising. Lesions on tomato or pepper leaves appear as irregular water-soaked areas, initially green and later becoming brown and necrotic. Speck lesions caused by P. syringae pv. tomato look similar in a first stage, but are surrounded by a more distinct yellow halo; lesions are often in streaks and the yellow haloes run together to give large chlorotic areas (Goode & Sasser, 1980). Severe infections in pepper cause defoliation, favoring sunscald of the fruits on hot and sunny days.

Bacterial spot xanthomonads can cause canker-like splits in stems, but their presence alone is not diagnostic as similar symptoms may also be caused by P. syringae pv. tomato, C. michiganensis and Alternaria solani. Pith necrosis has been associated with the presence of X. euvesicatoria pv. perforans, but this symptom could be confused with those of Pseudomonas mediterranea and Pseudomonas corrugata (Aiello et al., 2013).

Morphology

Xanthomonads causing bacterial spot of tomato and pepper are aerobic, mobile, Gram-negative rods, occurring singly or in pairs, 0.6 x 1.0-1.5 µm, with a single polar flagellum. Like other species of the genus, they produce characteristic yellow pigments (xanthomonadins). On general media, such as yeast-glucose-calcium carbonate agar (YGCA) or yeast peptone glucose agar (YPGA), colonies are mucoid-fluidal, convex, and yellow with entire edges. Unlike P. syringae pv. tomato, bacterial spot xanthomonads are non-fluorescent on King's B medium.

Detection and inspection methods

Symptoms of bacterial spot xanthomonads can be confused with those caused by other pathogens. Confirmation through diagnostic analysis is necessary when the presence of spot-causing bacteria is suspected. The EPPO Diagnostic protocol Standard PM 7/110 (2) (EPPO, 2023) offers comprehensive guidelines for the preliminary screening of plant material or seeds. The Standard includes isolation and molecular tests and, if required, confirmatory pathogenicity tests in susceptible cultivars of tomato and pepper plants. The EPPO Standard recommends multiplex real-time PCR (Strayer et al., 2016) and real-time PCR tests for specific identification of Xanthomonas species causing bacterial spot disease (Baldwin et al., 2023). Serological methods are not recommended in the Standard; few antibodies are commercially available for immunofluorescence and ELISA, and no validation data could be retrieved.

The International Seed Health Initiative (ISHI) advises using a minimum of 10 000 tomato or pepper seeds for seed detection purposes to ensure effective screening (ISF, 2017). This approach, with subsamples capped at 10 000 seeds, aims to identify contamination levels as low as 0.03% with a 95% confidence interval. In instances where there is a high likelihood of saprophytic bacteria overshadowing the presence of Xanthomonas spp., opting for smaller subsample sizes, such as five sets of 2 000 seeds each, is recommended.

A procedure by which consignments of tomato seeds should be subjected to phytosanitary import inspection, including sampling and identification, is provided in EPPO Standard PM 3/80 (2) (EPPO, 2021).

Trade of infected seeds and plants for planting (transplants) are associated with long-distance dissemination of Xanthomonas spp. which are responsible of causing bacterial spot. Bacteria may escape from infected plants as exudates, and short-distance dispersal is then facilitated by splashing water, such as irrigation or rain. This becomes particularly concerning during transplant in greenhouse production when several thousands of transplants are growing closely together, as well as in the field, especially in the case of sprinkler irrigation. Short-distance-spread of bacteria is also possible through contaminated tools (EFSA, 2014).

Economic impact

Bacterial spot xanthomonads are widespread and are considered significant pathogens of tomato and pepper in field-grown crops in warm-temperate and tropical countries, especially under overhead irrigation. They can also occur in greenhouses. Fruit yield losses are most substantial when infection occurs early, as observed in tomato in USA, and pepper in Israel (Dougherty, 1978; Bashan et al., 1985). Damage to the leaves tends to expose fruits to the sun, increasing the risk of sunscald. Although fruit lesions are often only superficial, they result in loss of marketability.

Bacterial spot has been recognized as a severe disease accompanied by significant damages in production in countries of the EPPO region, such as Serbia (Ignjatov et al., 2010; Vlajić et al., 2017), Bulgaria and the Republic of North Macedonia (Kizheva et al., 2011), or Türkiye (Aysan & Sahin, 2003). Although no recent data are available on economic losses caused by these pathogens in the European Union (EU), infections resulting in up to 30 % losses have been reported (EFSA, 2014). Yield losses of up to 66% have been reported in the USA (Pohronezny & Volin, 1983). Tomato yield loss estimated to be 7 413 USD per ha have been reported in Florida (USA) (Vallad et al., 2013). Furthermore, outbreaks in 2009-2010 in 2000 ha of processing tomatoes in Northwest Ohio and Southeast Michigan resulted in total losses of up to 7.8 USD million (Ma et al., 2011).

Control

Due to the seedborne nature of bacterial spot xanthomonads, management of the disease has been a major challenge since its original description. As no effective methods or chemical control agents are available for infected crops, disease control requires the adoption of integrated management measures, primarily focused on prevention and exclusion. Pathogen-free seeds and transplants are crucial to avoid the introduction and spread of bacteria. Tomato seed extraction from fruits using appropriate fermentative or acid treatments to reduce xanthomonads population is required in the EU (Anonymous, 2019). For both tomato and pepper seeds, hot water soak, dry heat therapy or selected chemicals have been recommended (EFSA, 2014).

Once infection occurs, control of disease in the field is particularly difficult. Cultural practices, such as four-year crop rotations, are recommended. Avoiding the handling of wet plant material and minimizing free moisture on foliage helps prevent disease development and spread. The application of protective chemicals or biological treatments is advised to reduce the severity and spread during transplant production.

The most common approach for managing bacterial spot pathogens is the preventive application of copper-based bactericides, but their success is limited as bacterial resistance to such chemicals has appeared worldwide (Lamichhane et al., 2018). Additionally, in the EU a new legislation limits the use of copper compounds (Anonymous, 2018). Several biological control approaches have been studied, including bacteriophages (Jones et al., 2012; Balogh et al., 2018; Gašić et al., 2018; Ríos-Sandoval et al., 2020), plant growth-promoting rhizobacterium (Naue et al., 2014; Liu et al., 2018) and antagonistic bacteria. Integration of biological control agents, such as tailocins (phage-tail-like bacteriocins produced by Pseudomonas fluorescens SF4c (Príncipe et al., 2018) and Bacillus velezensis GF267 (de Paula Kuyat Mates et al., 2019), and SARS inducers (harpin and acibenzolar-S-methyl) enhances the effectiveness of bacterial spot management (Obradovic et al., 2005; Abo-Elyours & El-Hendawy, 2008). The application of bacteriophages alone or in combination with biocontrol agents or copper hydroxide reduces disease incidence.

Efforts have been made to develop tomato and pepper lines with resistance to bacterial spot xanthomonads, and sources of resistance have been identified and incorporated into breeding programs and varieties (Stall et al., 2009). However, the persistence of resistance can change rapidly due to the evolving geographical distribution of the pathogen and the rapid emergence of new pathogenic variants (Potnis et al., 2015).

Phytosanitary risk

Xanthomonas spp. are important bacterial pathogens that affect tomato and pepper production. Environmental conditions in Southern Europe are particularly favorable for bacterial spot expression in the field, as the optimal growth temperature for xanthomonads is between 25°C and 30°C, but the disease also occurs in greenhouses (EFSA, 2014).

Long-distance spread of Xanthomonas spp. causing bacterial spot of tomato and pepper, is commonly related to the movement of infected seeds and planting material (transplants). Once introduced into a production area, such as a cultivation plot or a greenhouse, the pathogens disperse easily, and short-distance dispersal is facilitated by splashing water (from irrigation and rain) or contaminated tools (EFSA, 2014).

As the main means of spread of bacterial spot xanthomonads are with seeds and plants for planting, the use of healthy seed and plantlets are a key phytosanitary measure (Picard et al., 2018).

Currently Xanthomonas euvesicatoria pv. euvesicatoria, Xanthomonas euvesicatoria pv. perforans, Xanthomonas hortorum pv. gardneri and Xanthomonas vesicatoria are classified as regulated non-quarantine pests (RNQP) in many EPPO countries, including those within the EU (Anonymous, 2019), and measures to prevent their presence on seeds and planting material are mandatory. The EU measures are as follows: the presence of these bacteria is not allowed in propagating material of ornamental plants or other plants for planting intended for ornamental purposes, vegetable seeds, and vegetable propagating and planting material other than seeds (threshold level 0%) (Anonymous, 2019 Annex IV, Parts D, F, and I). For tomato seeds, an appropriate extraction method for bacterial elimination is required (there is no indication whether or not pepper seeds should be treated). Tomato and pepper seeds should originate in areas known to be free from these pathogens where no symptoms of bacterial spot have been observed during the complete cycle of vegetation of the plants at site of production. Alternatively, seeds should have been subjected to official testing on a representative sample using appropriate methods and have been found free of the pathogens (Anonymous, 2019 Annex V, Parts C, E, and H).

Abo-Elyousr KA & El-Hendawy HH (2008) Integration of Pseudomonas fluorescens and acibenzolar-S-methyl to control bacterial spot disease of tomato. Crop Protection 27(7), 1118-1124. https://doi.org/10.1016/j.cropro.2008.01.011

Adhikari P, Adhikari TB, Louws FJ & Panthee DR (2020) Advances and challenges in bacterial spot resistance breeding in tomato (Solanum lycopersicum L.). International Journal of Molecular Sciences 21(5), 1734. https://doi.org/10.3390/ijms21051734

Aiello D, Scuderi G, Vitale A, Firrao G, Polizzi G & Cirvilleri G (2013) A pith necrosis caused by Xanthomonas perforans on tomato plants. European Journal of Plant Pathology 137(1), 29-41. https://link.springer.com/article/10.1007/s10658-013-0214-7

Anonymous (2018) Commission Implementing Regulation (EU) 2018/1981 of 13 December 2018 renewing the approval of the active substances copper compounds, as candidates for substitution, in accordance with Regulation (EC) No 1107/2009 of the European Parliament and of the Council concerning the placing of plant protection products on the market, and amending the Annex to Commission Implementing Regulation (EU) No 540/2011. Official Journal of the European Union, L 317, 16-20. http://data.europa.eu/eli/reg_impl/2018/1981/oj

Anonymous (2019) Commission Implementing Regulation (EU) 2019/2072, of 28 November 2019, establishing uniform conditions for the implementation of Regulation (EU) 2016/2031 of the European Parliament and the Council, as regards protective measures against pests of plants, and repealing Commission Regulation (EC) No 690/2008 and amending Commission Implementing Regulation (EU) 2018/2019. Official Journal of the European Union, L 319, 1-279. http://data.europa.eu/eli/reg_impl/2019/2072/oj

Aysan Y & Sahin F (2003) Occurrence of bacterial spot disease, caused by Xanthomonas axonopodis pv. vesicatoria, on pepper in the Eastern Mediterranean region of Turkey. Plant Pathology 52(6), 781. https://doi.org/10.1111/j.1365-3059.2003.00890.x

Baldwin TK, Woudt B, Lastdrager J, Berendsen S & Koenraadt H (2023) Development and validation of real-time PCR tests for specific identification of Xanthomonas species causing bacterial spot disease on tomato (Solanum lycopersicum) and pepper (Capsicum annuum). EPPO Bulletin 53(2), 416-424. https://doi.org/10.1111/epp.12939

Balogh B, Nga NTT & Jones JB (2018) Relative level of bacteriophage multiplication in vitro or in phyllosphere may not predict in planta efficacy for controlling bacterial leaf spot on tomato caused by Xanthomonas perforans. Frontiers in Microbiology 9, 2176. https://doi.org/10.3389/fmicb.2018.02176 (Corrigendum: relative level of bacteriophage multiplication in vitro or in phyllosphere may not predict in planta efficacy for controlling bacterial leaf spot on tomato caused by Xanthomonas perforans. Frontiers in Microbiology, 9, 2647. https://doi.org/10.3389/fmicb.2018.02647

Bashan Y, Azaizeh M, Diab S, Yunis H & Okon Y (1985) Crop loss of pepper plants artificially infected with Xanthomonas campestris pv. vesicatoria in relation to symptom expression. Crop Protection 4(1), 77-84. https://doi.org/10.1016/0261-2194(85)90007-9

Bashan Y, Diab S & Okon Y (1982a) Survival of Xanthomonas campestris pv. vesicatoria in pepper seeds and roots, in symptomless and dry leaves in non-host plants and in the soil. Plant and Soil 68(2), 161-170. https://doi.org/10.1007/BF02373702

Bashan Y, Okon Y & Henis Y (1982b) Long-term survival of Pseudomonas syringae pv. tomato and Xanthomonas campestris pv. vesicatoria in tomato and pepper seeds. Phytopathology 72(9), 1143-1144.

Bernal RF & Berger RD (1996) The spread of epiphytic populations of Xanthomonas campestris pv. vesicatoria on pepper in the field. Journal of Phytopathology 144(9-10), 479-484. https://doi.org/10.1111/j.1439-0434.1996.tb00328.x

Bouzar H, Jones JB, Stall RE, Hodge NC, Minsavage GV, Benedict AA & Alvarez AM (1994) Physiological, chemical, serological, and pathogenic analyses of a worldwide collection of Xanthomonas campestris pv. vesicatoria strains. Phytopathology 84(7), 663‐671. https://doi.org/10.1094/phyto-84-663

Burlakoti RR, Hsu CF, Chen J, & Wang J (2018) Population dynamics of xanthomonads associated with bacterial spot of tomato and pepper during 27 years across Taiwan. Plant Disease 102(7), 1348-1356. https://doi.org/10.1094/PDIS-04-17-0465-RE

Constantin EC, Cleenwerck I, Maes M, Baeyen S, Van Malderghem C, De Vos P & Cottyn B (2016) Genetic characterization of strains named as Xanthomonas axonopodis pv. dieffenbachiae leads to a taxonomic revision of the X. axonopodis species complex. Plant Pathology 65(5), 792-806. https://doi.org/10.1111/ppa.12461

Diab S, Bashan Y, Okon Y & Henis Y (1982) Effects of relative humidity on bacterial scab caused by Xanthomonas campestris pv. vesicatoria on pepper. Phytopathology 72(9), 1257-1260.

Doidge EM (1920) A tomato canker. Journal of the Department of Agriculture, Union of South Africa 1, 718-721.

Doidge EM (1921) A tomato canker. Annals of Applied Biology 7(4), 407-430. https://doi.org/10.1111/j.1744-7348.1921.tb05528.x

Dougherty DE (1978) Yield reduction in tomato caused by bacterial spot and disease control with copper sprays. Proceedings of the Florida State Horticultural Society 91, 291-293.

Dowson WJ (1939) On the systematic position and generic names of the Gram negative bacterial plant pathogens. Zentralblatt fur Bakteriologie, Parasitenkunde und Infektionskrankheiten 100, 177-193.

Dutta B, Gitaitis R, Sanders RH, Booth C, Smith S & Langston DB, Jr (2013) Role of blossom colonization in pepper seed infestation by Xanthomonas euvesicatoria. Phytopathology 104(3), 232-239. https://doi.org/10.1094/PHYTO-05-13-0138-R

EFSA (2014) EFSA PLH Panel (EFSA Panel on Plant Health) (2014) Scientific Opinion on the pest categorisation of Xanthomonas campestris pv. vesicatoria (Doidge) Dye. EFSA Journal 12(6), 3720, 26 pp. https://doi.org/10.2903/j.efsa.2014.3720

EPPO (2021) EPPO Standards Phytosanitary procedures PM 3/80 (2). Consignment inspection of seed of Solanum lycopersicum and its hybrids. EPPO Bulletin 51, 397-403. https://doi.org/10.1111/epp.12773

EPPO (2023) EPPO Standard Diagnostics PM 7/110 (2) Xanthomonas spp. (Xanthomonas euvesicatoria pv. euvesicatoria, Xanthomonas hortorum pv. gardneri, Xanthomonas euvesicatoria pv. perforans, Xanthomonas vesicatoria) causing bacterial spot of tomato and sweet pepper. EPPO Bulletin 53, 558-579. https://doi.org/10.1111/epp.12960

Gardner MW & Kendrick JB (1921) Bacterial spot of tomato. Journal of Agricultural Research 21, 123-156.

Gardner MW & Kendrick JB (1923) Bacterial spot of tomato and pepper. Phytopathology 13, 307-315.

Gašić K, Kuzmanović N, Ivanović M, Prokić A, Šević M & Obradović A (2018) Complete genome of the Xanthomonas euvesicatoria specific bacteriophage KΦ1, its survival and potential in control of pepper bacterial spot. Frontiers in Microbiology 9, 2021. https://doi.org/10.3389/fmicb.2018.02021

Goode MJ & Sasser M (1980) Prevention - the key to controlling bacterial speck and bacterial speck of tomato. Plant Disease 64, 831-834.

Holt JG (1994) Bergey’s manual of determinative bacteriology. 9th Edition, Lippincott Williams and Wilkins, Baltimore. Philadelphia, Pennsylvania, 787 pp.

Ignjatov M, Gasić K, Ivanović M, Šević M, Obradović A & Milošević M (2010) Characterisation of Xanthomonas euvesicatoria strains pathogens of pepper in Serbia. Pesticidi i Fitomedicina 25,139-149. English summary (In Serbian).

ISF (2017) Method for the Detection of Xanthomonas spp. in Tomato seed (available online). https://worldseed.org/our-work/seed-health/ishi-methods/

Jibrin MO, Timilsina S, Minsavage GV, Vallad GE, Roberts PD, Goss EM & Jones JB (2022) Bacterial spot of tomato and chili pepper in Africa: diversity, emergence of T5 race, and management. Frontiers in Microbiology 13, 835647. https://doi.org/10.3389/fmicb.2022.835647

Jones JB, Bouzar H. Stall RE, Almira EC, Roberts P, Bowen BW, Jones JB, Bouzar H, Stall RE, Almira EC, Roberts PD, Bowen BW, Sudberry J, Strickler PM & Chun J (2000) Systematic analysis of xanthomonads (Xanthomonas spp.) associated with pepper and tomato lesions. International Journal of Systematic Bacteriology 50(3), 1211-1219. https://doi.org/10.1099/00207713-50-3-1211

Jones JB, Lacy GH, Bouzar H, Stall RE & Schaad NW (2004a) Reclassification of the xanthomonads associated with bacterial spot disease of tomato and pepper. Systematic and Applied Microbiology 27(6), 755-762. https://doi.org/10.1078/0723202042369884

Jones JB, Lacy GH, Bouzar H, Minsavage GV, Stall R & Schaad N (2004b) Bacterial spot—worldwide distribution, importance and review. 1st International Symposium on Tomato Diseases. Acta Horticulturae 695, 27-33. https://doi.org/10.17660/ActaHortic.2005.695.1

Jones J, Stall R, Scott J, Somodi G, Bouzar H & Hodge N (1995) A third tomato race of Xanthomonas campestris pv. vesicatoria. Plant Disease 79, 395-398.

Jones JB, Vallad GE, Iriarte FB, Obradović A, Wernsing MH, Jackson LE, Balogh B., Hong JC & Momol T (2012) Considerations for using bacteriophages for plant disease control. Bacteriophage 2(4), 208-214. https://doi.org/10.4161/bact.23857

Kizheva Y, Vancheva T, Hristova P, Stoyanova M, Bogatzevska N & Moncheva P (2011) Diversity of Xanthomonas spp. causal agents of bacterial spot on pepper and tomato plants in Bulgaria. Biotechnology & Biotechnological Equipment 25(4), (Suppl. 4), 98-104. https://doi.org/10.5504/BBEQ.2011.0126

Klein-Gordon JM, Xing Y, Garrett KA, Abrahamian P, Paret ML, Minsavage GV, Strayer-Scherer AL, Fulton JC, Timilsina S, Jones JB, Goss EM & Vallad GE (2021) Assessing changes and associations in the Xanthomonas perforans population across Florida commercial tomato fields via a statewide survey. Phytopathology 111(6), 1029-1041. https://doi.org/10.1094/PHYTO-09-20-0402-R

Lamichhane JR, Osdaghi E, Behlau F, Köhl J, Jones JB, & Aubertot JN (2018) Thirteen decades of anti-microbial copper compounds applied in agriculture. A review. Agronomy for Sustainable Development 38, 28. https://doi.org/10.1007/s13593-018-0503-9

Ma X, Ivey MLL, & Miller SA (2011) First report of Xanthomonas gardneri causing bacterial spot of tomato in Ohio and Michigan. Plant Disease 95(12), 1584. https://doi.org/10.1094/PDIS-05-11-0448

Liu K, McInroy JA, Hu CH & Kloepper JW (2018) Mixtures of plant-growth-promoting rhizobacteria enhance biological control of multiple plant diseases and plant-growth promotion in the presence of pathogens. Plant Disease 102(1), 67-72. https://doi.org/10.1094/PDIS-04-17-0478-RE

McInnes TB, Gitaitis RD, McCarter SM, Jaworski CA & Phatak SC (1988) Airborne dispersal of bacteria in tomato and pepper transplant fields. Plant Disease 72, 575-579.

Morinière L, Burlet A, Rosenthal ER, Nesme X, Portier P, Bull CT, Lavire C, Fischer-Le Saux M & Bertolla F (2020) Clarifying the taxonomy of the causal agent of bacterial leaf spot of lettuce through a polyphasic approach reveals that Xanthomonas cynarae Trébaol et al. 2000 emend. Timilsina et al. 2019 is a later heterotypic synonym of Xanthomonas hortorum Vauterin et al. 1995. Systematic and Applied Microbiology 43(4), 126087. https://doi.org/10.1016/j.syapm.2020.126087

Naue CR, Rocha DJA & Moura AB (2014) Biological control of tomato bacterial spot by seed microbiolization. Tropical Plant Pathology 39(5), 413–416. https://doi.org/10.1590/S1982-56762014000500009

Newberry EA, Bhandari R, Minsavage GV, Timilsina S, Jibrin MO, Kemble J, Sikora EJ, Jones JB, Potnis N 2019) Independent evolution with the gene flux originating from multiple Xanthomonas species explains genomic heterogeneity in Xanthomonas perforans. Applied and Environmental Microbiology 85(20), e00885-19. https://doi.org/10.1128/AEM.00885-19

Obradovic A, Jones JB, Momol MT, Olson SM, Jackson LE, Balogh B, Guven K & Iriarte FB (2005) Integration of biological control agents and systemic acquired resistance inducers against bacterial spot on tomato. Plant Disease 89(7), 712-716. https://doi.org/10.1094/PD-89-0712

Osdaghi E, Jones JB, Sharma A, Goss EM, Abrahamian P, Newberry EA, Potnis N, Carvalho R, Choudhary M, Paret ML, Timilsina S & Vallad GE (2021) A centenary for bacterial spot of tomato and pepper. Molecular Plant Pathology 22(12),1500-1519. https://doi.org/10.1111/mpp.13125

de Paula Kuyat Mates A, de Carvalho Pontes N & de Almeida Halfeld-Vieira B (2019) Bacillus velezensis GF267 as a multi-site antagonist for the control of tomato bacterial spot. Biological Control 137, 104013. https://doi.org/10.1016/j.biocontrol.2019.104013

Picard C, Afonso T, Benko‐Beloglavec A, Karadjova O, Matthews‐Berry S, Paunovic SA, Pietsch M, Reed P, Van Der Gaag DJ, Ward M (2018) Recommended regulated non‐quarantine pests (RNQPs), associated thresholds and risk management measures in the European and Mediterranean region. EPPO Bulletin 48(3), 552-568. https://doi.org/10.1111/epp.12500

Pohronezny K, Moss MA, Dankers W & Schenk J (1990) Dispersal and management of Xanthomonas campestris pv. vesicatoria during thinning of direct-seeded tomato. Plant Disease 74, 800-805.

Pohronezny K & Volin RB (1983) The effect of bacterial spot on yield and quality of fresh market tomatoes. HortScience 18(1), 69-70. https://doi.org/10.21273/HORTSCI.18.1.69

Potnis N, Timilsina S, Strayer A, Shantharaj D, Barak JD, Paret ML, Vallad GE & Jones JB (2015) Bacterial spot of tomato and pepper: Diverse Xanthomonas species with a wide variety of virulence factors posing a worldwide challenge. Molecular Plant Pathology 16(9), 907-920. https://doi.org/10.1111/mpp.12244

Príncipe A, Fernandez M, Torasso M, Godino A & Fischer S (2018) Effectiveness of tailocins produced by Pseudomonas fluorescens SF4c in controlling the bacterial-spot disease in tomatoes caused by Xanthomonas vesicatoria. Microbiological Research 212–213, 94–102. https://doi.org/10.1016/j.micres.2018.05.010

Ríos-Sandoval M, Quiñones-Aguilar EE, Solís-Sánchez GA, Enríquez-Vara JN & Rincón-Enríquez G (2020) Complete genome sequence of Xanthomonas vesicatoria bacteriophage ΦXaF18, a contribution to the biocontrol of bacterial spot of pepper in Mexico. Microbiology Resource Announcements 9(16), e00213-20. https://doi.org/10.1128/mra.00213-20

Stall RE, Beaulieu C, Egel D, Hodge NC, Leite RP, Minsavage GV, Bouzar H, Jones JB, Alvarez AM & Benedict AA (1994) Two genetically diverse groups of strains are included in Xanthomonas campestris pv. vesicatoria. International Journal of Systematic Bacteriology 44(1), 47-53. https://doi.org/10.1099/00207713-44-1-47

Stall RE, Jones JB & Minsavage GV (2009) Durability of resistance in tomato and pepper to xanthomonads causing bacterial spot. Annual Review of Phytopathology 47, 265‐284. https://doi.org/10.1146/annurev-phyto-080508-081752

Strayer AL, Jeyaprakash A, Minsavage GV, Timilsina S, Vallad GE, Jones JB &Paret ML (2016) A multiplex real-time PCR assay differentiates four Xanthomonas species associated with bacterial spot of tomato. Plant Disease 100(8), 1660-1668. https://doi.org/10.1094/PDIS-09-15-1085-RE

Šutic D (1957) Bakterioze crvenog patlidzana (Tomato bacteriosis). Posebna Izd. Inst. Zasht. Bilja Beograd (Spec. Edit. Inst. Plant Prot. Beograd) 6, 1-65. English summary: Review of Applied Mycology 36, 734-735.

Timilsina S, Jibrin MO, Potnis N, Minsavage GV, Kebede M, Schwartz A, Bart R, Staskawicz B, Boyer C, Vallad GE, Pruvost O, Jones JB & Goss EM (2015) Multilocus sequence analysis of xanthomonads causing bacterial spot of tomato and pepper plants reveals strains generated by recombination among species and recent global spread of Xanthomonas gardneri. Applied and Environmental Microbiology 81(4), 1520-1529. https://doi.org/10.1128/AEM.03000-14

Vallad GE, Timilsina S, Adkison H, Potnis N, Minsavage G, Jones J, & Goss EA (2013) A recent survey of Xanthomonads causing bacterial spot of tomato in Florida provides insights into management strategies. Proceedings of the 2013 Florida Tomato Institute (eds Ozores-Hampton M & Snodgrass C), pp. 25-27. University of Florida, Naples, FL (USA).

Vancheva T, Bogatzevska N, Moncheva P, Mitrev S, Vernière C & Koebnik R (2021) Molecular epidemiology of Xanthomonas euvesicatoria strains from the Balkan Peninsula revealed by a new multiple-locus variable-number tandem-repeat analysis scheme. Microorganisms 9(3), 536. https://doi.org/10.3390/microorganisms9030536

Vauterin L, Hoste B, Kersters K & Swings J (1995) Reclassification of Xanthomonas. International Journal of Systematic Bacteriology 45(3), 472-489. https://doi.org/10.1099/00207713-45-3-472

Vlajić S, Iličić R, Maširević S, Feldeždi M, & Jošič D (2017) Appearance of Xanthomonas euvesicatoria on pepper in Vojvodina during 2016. Proceedings of 69th International Symposium on Crop Protection, p. 159. University of Ghent, Faculty of Bioscience Engineering. Ghent, Belgium.

Young JM, Dye DW, Bradbury JF, Panagopoulos CG & Robbs CF (1978) A proposed nomenclature and classification for plant pathogenic bacteria. New Zealand Journal of Agricultural Research 21(1), 153-177. https://doi.org/10.1080/00288233.1978.10427397

This datasheet was extensively revised in 2024 by Ana Palacio-Bielsa (CITA, Centro de Investigación y Tecnología Agroalimentaria de Aragón, Spain / Instituto Agroalimentario de Aragón - IA2 (CITA-Universidad de Zaragoza, Spain), Jaime Cubero (INIA/CSIC, Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria/Consejo Superior De Investigaciones Científicas, Spain) and Jerson Garita-Cambronero (ANOVE, Asociación Nacional de Obtentores Vegetales, Spain / ITACyL, Instituto Tecnológico Agrario de Castilla y León, Spain). Their valuable contribution is gratefully acknowledged.

EPPO (2024) Xanthomonas vesicatoria. EPPO datasheets on pests recommended for regulation. https://gd.eppo.int (accessed 2024-12-23)

This datasheet was first published in the EPPO Bulletin in 1988 and revised in the two editions of 'Quarantine Pests for Europe’ in 1992 and 1997, as well as in 2024. It is now maintained in an electronic format in the EPPO Global Database. The sections on 'Identity', ‘Hosts’, and 'Geographical distribution' are automatically updated from the database. For other sections, the date of last revision is indicated on the right.

CABI/EPPO (1992/1997) Quarantine Pests for Europe (2^nd edition). CABI, Wallingford (GB).

EPPO (1988) Data sheets on quarantine organisms No. 157, Xanthomonas campestris pv. vesicatoria. EPPO Bulletin 18, 521-526. https://onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2338.1988.tb00409.x