Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Peronospora belbahrii
(basil downy mildew)

Crouch J A, 2020. Peronospora belbahrii (basil downy mildew). Invasive Species Compendium. Wallingford, UK: CABI. DOI:10.1079/ISC.118352.20210200694

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Peronospora belbahrii (basil downy mildew)

Summary

  • Last modified
  • 29 June 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Peronospora belbahrii
  • Preferred Common Name
  • basil downy mildew
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Chromista
  •     Phylum: Oomycota
  •       Class: Oomycetes
  •         Order: Peronosporales
  • Summary of Invasiveness
  • Basil downy mildew was first identified from Uganda during 1932 and 1937, resulting in significant crop losses (Hansford, 1933;

  • Principal Source
  • Draft datasheet under review.

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Pictures

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PictureTitleCaptionCopyright
Peronospora belbahrii; symptoms, showing upper leaf surface of a basil plant (Ocimum basilicum) affected by typical dark, angular, vein-delimited lesions.
TitleSymptoms
CaptionPeronospora belbahrii; symptoms, showing upper leaf surface of a basil plant (Ocimum basilicum) affected by typical dark, angular, vein-delimited lesions.
Copyright©Scot Nelson/via flickr - CC BY 2.0
Peronospora belbahrii; symptoms, showing upper leaf surface of a basil plant (Ocimum basilicum) affected by typical dark, angular, vein-delimited lesions.
SymptomsPeronospora belbahrii; symptoms, showing upper leaf surface of a basil plant (Ocimum basilicum) affected by typical dark, angular, vein-delimited lesions.©Scot Nelson/via flickr - CC BY 2.0

Identity

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Preferred Scientific Name

  • Peronospora belbahrii Thines

Preferred Common Name

  • basil downy mildew

Summary of Invasiveness

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Basil downy mildew was first identified from Uganda during 1932 and 1937, resulting in significant crop losses (Hansford, 1933; Hansford, 1939). Following these original outbreaks, the disease was reported sporadically in Africa during the twentieth century: in Tanzania in 1960 (Riley, 1960), then again in Benin during 1998 (Gumedzoe et al., 1998). The disease was first identified outside of Africa in 2001, when it was reported from Switzerland (Belbahri et al., 2005). Unlike the intermittent African outbreaks of the twentieth century, the twenty-first century outbreaks of basil downy mildew are persistent, and the geographic range of P. belbahrii continues to expand. Since 2001, P. belbahrii has spread throughout Europe, North America, Asia, and parts of Africa, Central and South America, and the Caribbean. Losses incurred due to basil downy mildew in the USA alone are estimated to reach tens of millions of dollars (Wyenandt et al., 2015).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Chromista
  •         Phylum: Oomycota
  •             Class: Oomycetes
  •                 Order: Peronosporales
  •                     Family: Peronosporaceae
  •                         Genus: Peronospora
  •                             Species: Peronospora belbahrii

Notes on Taxonomy and Nomenclature

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Peronospora belbahrii is an obligate plant pathogenic oomycete that causes downy mildew disease of basil (Ocimum basilicum) (Thines et al., 2009; Hoffmeister et al., 2020; Salgado-Salazar et al., 2020).

Early reports of P. belbahrii infecting basil referred to the organism as P. lamii (Hansford, 1939; Riley 1960; Gamliel and Yarden, 1998; Gumedzoe et al., 1998; Martini et al., 2003; Liberato et al., 2006) following the broad species concept of Yerkes and Shaw (1959) whereby entire host families were associated with a single pathogen species. However, subsequent investigations showed that P. lamii is restricted to the downy mildew pathogen of Lamium spp. (Choi et al., 2009; Thines et al., 2009) and does not infect the entirety of the Lamiaceae.

As basil downy mildew outbreaks became globally widespread in the early years of the twenty-first century, authors mostly referred to the pathogen as Peronospora sp., reflecting the inability of the existing taxonomy to provide an accurate name for this organism (Belbahri et al., 2005; Garibaldi et al., 2004; Garibaldi et al., 2005; Blomquist et al., 2009; Ronco et al., 2009; Martinez de la Parte et al., 2010; McLeod et al., 2006; Roberts et al., 2009; Voglmayr and Piatek, 2009; Wick and Brazee, 2009). P. belbahrii was described in 2009 on the basis of molecular phylogenetic relationships, morphology and host range (Thines et al., 2009).

Distribution

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Basil downy mildew was first identified from Uganda during 1932 and 1937, resulting in significant crop losses (Hansford, 1933; Hansford, 1939). Following the original outbreaks, the disease was reported sporadically in Africa during the twentieth century: in Tanzania in 1960 (Riley, 1960), then again in Benin during 1998 (Gumedzoe et al., 1998). The disease was first identified outside of Africa in 2001, when it was reported from Switzerland (Belbahri et al., 2005). Unlike the intermittent African outbreaks of the twentieth century, the twenty first century outbreaks are persistent, and the geographic range of P. belbahrii continues to expand. Since 2001, P. belbahrii has spread throughout Europe, North America, Asia, and parts of Africa, Central and South America, and the Caribbean.

In the USA, after the disease was first identified from Florida in 2007, scientists began monitoring the spread of P. belbahrii using the online reporting tool Basil Ag Pest Monitor (https://basil.agpestmonitor.org/), as described in Wyenandt et al. (2015). Between 2009 and 2019, 1386 reports of basil downy mildew have been made from 44 states and the District of Columbia (http://blogs.cornell.edu/livegpath/extension/basil-downy-mildew/where-in-the-usa-is-basil-downy-mildew/).

The origin of P. belbahrii is unknown. Thines et al. (2009) speculated that P. belbahrii might be of African origin, given the fact that the biodiversity centre of Ocimum is Africa (Patton, 1992). This interpretation is consistent with the fact that the pathogen was only known from Africa for almost 70 years.

There is a risk of intentional introduction of this pathogen. There is evidence that the pathogen is transmitted through seeds, providing a means for long-distance transport (Cohen et al., 2017). The pathogen may also be present in asymptomatic stems and leaves, facilitating unintentional transport on host cuttings (Cohen et al., 2017).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 17 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

BeninPresent1998Invasive
CameroonPresent2007InvasivePlantations in Obala town (Centre Province), Cameroon.
South AfricaPresent2005InvasiveWestern Cape Province.
TanzaniaPresent
UgandaPresent1932

Asia

ChinaPresent, Localized
-BeijingPresent, Localized2016InvasiveShunyi and Daxing districts of Beijing.
-HainanPresent2014InvasiveSanya City (18°N) of Hainan Province.
IranPresent, Widespread2006Invasive
IsraelPresent, Widespread2011: Found in two greenhouses in the northern Jordan Valley, then at the southwest and southeast Israeli borders. Summer 2012: disease widespread in Israel.
JapanPresent
-HonshuAbsent, Invalid presence record(s)This report of Peronospora belbahrii from coleus is most likely P. choii (see Hoffmeister et al., 2020).
South KoreaPresent, Localized2015Invasive
TaiwanPresent2009InvasiveNantu and Yunlin.

Europe

AustriaPresent2005InvasiveStyria: East Styrian Hills, Graz, Ragnitztal on the eastern outskirts.
BelgiumPresent2004Invasive
CroatiaPresent, Widespread2012InvasiveDubrovnik-Neretva, Varaždin, Krapina-Zagorje, Medimurje, Split-Dalmatian and Zagreb Counties.
CyprusPresent, Localized2012Invasive
CzechiaPresent, Widespread2012InvasiveCentral Bohemia.
FrancePresent, Few occurrences2004Invasive
GermanyPresent2005InvasiveLankreis, Heilbronn.
HungaryPresent, Few occurrences2003InvasiveAlbertirsa, Budapest-Soroksár, Tordas.
ItalyPresent2003Invasive
MaltaPresent2005Invasive
SpainPresent2016Invasive
-Canary IslandsPresent
SwitzerlandPresent2002InvasiveEastern Switzerland, island of Reichenau.
United KingdomPresent, Localized2005Invasive
-EnglandPresent, Localized

North America

CanadaPresent, Localized
-British ColumbiaPresent2011Invasive
-OntarioPresent, Few occurrences2011Invasive
-QuebecPresent2011Invasive
CubaPresent2009InvasiveHorticulture Research Institute, Liliana Dimitrova in Quivican, Havana province and Siboney, Havana City.
MexicoPresent, Localized2009Invasive
United StatesPresent, Widespread
-AlabamaPresent2010Invasive
-AlaskaPresent2011Invasive
-ArkansasPresent2008Invasive
-CaliforniaPresent, Widespread2008Invasive
-ColoradoPresent, Widespread2012Invasive
-ConnecticutPresent2009Invasive
-DelawarePresent2009Invasive
-District of ColumbiaPresent2009Invasive
-FloridaPresent, Widespread2007InvasiveCollier, Hendry, Miami-Dade, and Palm Beach counties in south Florida; Hillsborough County in west-central north Florida.
-GeorgiaPresent2012Invasive
-HawaiiPresent, Widespread2010InvasiveWai?anac Oahu Maui Molokai Hawaii
-IllinoisPresent2009Invasive
-IndianaPresent2009Invasive
-IowaPresent2014Invasive
-KansasPresent2008Invasive
-KentuckyPresent, Widespread2010Invasive
-LouisianaPresent2009Invasive
-MainePresent2012Invasive
-MarylandPresent, Widespread2009Invasive
-MassachusettsPresent, Widespread2008Invasive
-MichiganPresent2010Invasive
-MinnesotaPresent2011Invasive
-MississippiPresent2016Invasive
-MissouriPresent, Widespread2012Invasive
-MontanaPresent, Widespread2010Invasive
-NebraskaPresent2014Invasive
-New HampshirePresent2012Invasive
-New JerseyPresent, Widespread2008Invasive
-New MexicoPresent2013Invasive
-New YorkPresent, Widespread2008Invasive
-North CarolinaPresent2008Invasive
-North DakotaPresent2008Invasive
-OhioPresent2010Invasive
-OklahomaPresent2016Invasive
-OregonPresent2012Invasive
-PennsylvaniaPresent, Widespread2009Invasive
-Rhode IslandPresent2012Invasive
-South CarolinaPresent2009Invasive
-TennesseePresent2013Invasive
-TexasPresent2009Invasive
-VermontPresent, Widespread2009Invasive
-VirginiaPresent2010Invasive
-WashingtonPresent2011Invasive
-West VirginiaPresent2012Invasive
-WisconsinPresent2009Invasive

Oceania

New ZealandPresent

South America

ArgentinaPresent2008InvasiveLa Plata
BrazilPresent
-Minas GeraisAbsent, Unconfirmed presence record(s)This report of Peronospora belbahrii from coleus is most likely P. choii (see Hoffmeister et al., 2020).
-Sao PauloPresent, Few occurrences

Risk of Introduction

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There is a risk of intentional introduction of this pathogen. There is evidence that the pathogen is transmitted through seeds, providing a means for long-distance transport (Cohen et al., 2017). The pathogen may also be present in asymptomatic stems and leaves, facilitating unintentional transport on host cuttings (Cohen et al., 2017).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Present, no further details
Terrestrial ManagedProtected agriculture (e.g. glasshouse production) Present, no further details

Hosts/Species Affected

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Prior to 2020, the name P. belbahrii was sometimes used to refer to Peronospora causing newly emergent downy mildew diseases affecting two other herbaceous plants in the Lamiaceae: namely, agastache (Agastache spp.) and coleus (Plectranthus scuttarioides). However, as early as 2009, many researchers recognized that the pathogens associated with these three different hosts were diverse, and exhibited variation between one another in DNA marker sequences. The publication originally describing P. belbahrii first introduced the idea that the coleus pathogen was most likely distinct from P. belharii on basil (Thines et al., 2009). As a result, while the literature shows some authors referring to the causal pathogens of coleus and agastache as P. belbahrii (Henricot et al., 2010; Denton et al., 2015; Ito et al., 2015), many others identified these organisms as Peronospora sp. or P. belbahrii sensu lato (Daughtrey et al., 2006; Palmateer et al., 2008; Thines et al., 2009; Rivera et al., 2016a; Rivera et al., 2016b; Gorayeb et al., 2020). In 2020, the pathogen of coleus was described as the new species, P. choii (Hoffmeister et al., 2020), while the pathogen of agastache was described as the new species, P. monardae (Salgado-Salazar et al., 2020). Peronospora belbahrii, P. choi and P. monardae are reliably distinguished from one another on the basis of molecular and morphological criteria (Hoffmeister et al., 2020; Salgado-Salazar et al., 2020).

There is some evidence that P. balbahrii isolates from basil can infect other plants in the Lamiaceae under experimental conditions (Micromeria fruticosa, Nepeta curviflora, Rosmarinus officinalis, Salvia pinnata and S. fruticosa; Cohen et al., 2017) but it is unknown whether any of these plants play any role in natural populations.

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Ocimum basilicum (basil)LamiaceaeMain

    Growth Stages

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    Vegetative growing stage

    Symptoms

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    Initial symptoms typically occur on lower leaves starting with yellowing of the leaf while the leaf vein remains green. This stage of the disease is easily misdiagnosed as a nutritional deficiency. Eventually necrosis occurs in the centre of chlorotic lesions. Leaves may become slightly curved. Necrotic lesions may be brown to black, can be variable in size, with irregular shapes bordered by leaf veins. Greyish-brown growth of the pathogen (sporangia) can be seen on the abaxial leaf surface, and sometimes on the adaxial surface.

    List of Symptoms/Signs

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    SignLife StagesType
    Leaves / abnormal colours
    Leaves / abnormal forms
    Leaves / abnormal leaf fall
    Leaves / fungal growth
    Leaves / necrotic areas
    Leaves / yellowed or dead

    Biology and Ecology

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    P. belbahrii is an obligate plant pathogen, requiring a living host to survive. The pathogen has been shown to colonize basil leaves, stems and seeds (Farahani-Kofoet et al., 2012). However, it is not clear to what extent P. belbahrii colonizes seeds or systemically infects host plants during natural infections (Cohen et al., 2017).

    Although environmental conditions influence the timing of specific events in the life-cycle of P. belbahrii, there is a general understanding of how the pathogen colonizes and reproduces in the host plant. Cohen et al. (2017) produces a comprehensive account of the infection cycle. Briefly, P. belbahrii reproduces asexually, producing dark-brown, purplish conidia that germinate to form germ tubes (Thines et al., 2009; Koroch et al., 2013; Zhang et al., 2020) between 3-6 days after coming into contact with a susceptible host (Zhang et al., 2020). At least 2 hours of exposure to water and temperatures of 15-20°C is required for germination (Cohen and Ben Naim, 2016), although some reports show 6 h leaf wetness is required (Garibaldi et al., 2007). Host tissue is penetrated directly (Cohen and Ben-Naim, 2016; Cohen et al., 2017; Zhang et al., 2020) or the pathogen may enter via stomata reported (Koroch et al., 2013; Cohen et al., 2017). A latent period can last for 5-10 days and is influenced by light and temperature (Cohen et al., 2017). Between 6 and 14 days after infection, the pathogen produces conidiophores that emerge through stomata on the abaxial leaf surface, or 1-2 days later through stomata on the adaxial surface (Zhang et al., 2020). New conidia can be produced as soon as 10 days after the initial infection (Zhang et al., 2020). Sporulation is inhibited by light (López-López et al., 2014) and may be inhibited by the lack of water although not all studies agree on this point (Garibaldi et al., 2007; Cohen and Ben-Naim, 2016; Cohen et al., 2017).

    Although downy mildew pathogens overwinter by producing sexual reproductive structures called oospores, to date these structures are only known for P. belbahrii from Israel (Cohen et al., 2013; Elad et al., 2016; Cohen et al., 2017). No information is available about whether P. belbahrii oospores are produced through a homothallic or heterothallic mating system.

    Means of Movement and Dispersal

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    There is evidence that the pathogen is transmitted through seeds, providing a means for long-distance transport (Cohen et al., 2017). The pathogen may also be present in asymptomatic stems and leaves, facilitating unintentional transport on host cuttings (Cohen et al., 2017).

    Seedborne Aspects

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    ​There is evidence that the pathogen is transmitted through seeds (Cohen et al., 2017).

    Economic Impact

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    Basil downy mildew was first identified from Uganda during 1932 and 1937, resulting in significant crop losses (Hansford, 1933Hansford, 1939). Following these original outbreaks, the disease was reported sporadically in Africa during the twentieth century. However, the twenty-first century outbreaks of basil downy mildew are persistent, and the geographic range of P. belbahrii continues to expand. Since 2001, P. belbahrii has spread throughout Europe, North America, Asia, and parts of Africa, Central and South America, and the Caribbean. Losses incurred due to basil downy mildew in the USA alone are estimated to reach tens of millions of dollars (Wyenandt et al., 2015).

    References

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    Belbahri, L., Calmin, G., Pawlowski, J., Lefort, F., 2005. Phylogenetic analysis and Real Time PCR detection of a presumbably undescribed Peronospora species on sweet basil and sage. Mycological Research, 109(11), 1276-1287. doi: 10.1017/S0953756205003928

    Blomquist, C. L., Rooney-Latham, S., Nolan, P. A., 2009. First report of downy mildew on field-grown sweet basil caused by a Peronospora sp. in San Diego County, California. Plant Disease, 93(9), 968. doi: 10.1094/PDIS-93-9-0968A

    CABI/EPPO, 2015. Peronospora belbahrii. [Distribution map]. Distribution Maps of Plant Diseases, No.October. Wallingford, UK: CABI, Map 1177 (Edition 1)

    Chen CH, Huang JH, Hsieh TF, 2010. First report of Peronospora belbahrii causing downy mildew on basil. Plant Pathology Bulletin, 19(3):177-180

    Choi YoungJoon, Shin HyeonDong, Thines, M., 2009. Two novel Peronospora species are associated with recent reports of downy mildew on sages. Fungal Biology, 113(12), 1340-1350. doi: 10.1016/j.mycres.2009.08.010

    Cohen Y, Vaknin M, Ben-Naim Y, Rubin AE, Galperin M, Silverman D, Bitton S, Adler U, 2013. First report of the occurrence and resistance to mefenoxam of Peronospora belbahrii, causal agent of downy mildew of basil (Ocimum basilicum) in Israel. Plant Disease, 97(5):692. http://apsjournals.apsnet.org/loi/pdis

    Cohen, Y., Ben-Naim, Y., 2016. Nocturnal fanning suppresses downy mildew epidemics in sweet basil. PLoS ONE, 11(5), e0155330. doi: 10.1371/journal.pone.0155330

    Cohen, Y., Naim, Y. B., Falach, L., Rubin, A. E., 2017. Epidemiology of basil downy mildew. Phytopathology, 107(10), 1149-1160. http://apsjournals.apsnet.org/loi/phyto

    Daughtrey, M. L., Holcomb, G. E., Eshenaur, B., Palm, M. E., Belbahri, L., Lefort, F., 2006. First report of downy mildew on greenhouse and landscape coleus caused by a Peronospora sp. in Louisiana and New York. Plant Disease, 90(8), 1111. doi: 10.1094/PD-90-1111B

    Denton GJ, Beal E, Denton JO, Clover G, 2015. First record of downy mildew, caused by Peronospora belbahrii, on Solenostemon scutellarioides in the UK. New Disease Reports, 31:14. http://www.ndrs.org.uk/article.php?id=031014

    Elad, Y., Omer, C., Nisan, Z., Harari, D., Goren, H., Adler, U., Silverman, D., Biton, S., 2016. Passive heat treatment of sweet basil crops suppresses Peronospora belbahrii downy mildew. Annals of Applied Biology, 168(3), 373-389. doi: 10.1111/aab.12269

    EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

    Farahani-Kofoet, R. D., Römer, P., Grosch, R., 2012. Systemic spread of downy mildew in basil plants and detection of the pathogen in seed and plant samples. Mycological Progress, 11(4), 961-966. doi: 10.1007/s11557-012-0816-z

    Gamliel, A., Yarden, O., 1998. Diversification of diseases affecting herb crops in Israel accompanies the increase in herb crop production. Phytoparasitica, 26(1), 53-58.

    Garibaldi, A., Bertetti, D., Gullino, M. L., 2007. Effect of leaf wetness duration and temperature on infection of downy mildew (Peronospora sp.) of basil. Journal of Plant Diseases and Protection, 114(1), 6-8. http://www.ulmer.de

    Garibaldi, A., Minuto, A., Gullino, M. L., 2005. First report of downy mildew caused by Peronospora sp. on basil (Ocimum basilicum) in France. Plant Disease, 89(6), 683. doi: 10.1094/PD-89-0683C

    Garibaldi, A., Minuto, A., Minuto, G., Gullino, M. L., 2004. First report of downy mildew on basil (Ocimum basilicum) in Italy. Plant Disease, 88(3), 312. doi: 10.1094/PDIS.2004.88.3.312A

    Gorayeb, E. S., Pieroni, L. P., Cruciol, G. C. D., Pieri, C. de, Dovigo, L. H., Pavan, M. A., Kurozawa, C., Krause-Sakate, R., 2020. First report of downy mildew on coleus (Plectranthus spp.) caused by Peronospora belbahrii sensu lato in Brazil. Plant Disease, 104(1), 294-294. doi: 10.1094/PDIS-07-19-1551-PDN

    Gumedzoe, MYD, Hemou, P, Lepage, A, Lecat, N, 1998. (Inventaire et identification de Peronospora lamii (Al. Br.) de By, agent causal du mildiou du basilic (Ocimum basilicum) dans les exploitations de Gyma Cultures). In: Les VIIIe Journe´es Scientifiques de l’Universite´ du Be´nin, 11–15 mai 1998 . Lome, Benin

    Hansford, CG, 1933. Annual report of the mycologist. Rev. Appl. Mycol, 12, 421-422.

    Hansford, CG, 1939. Annual report of the mycologist. Rev. Appl. Mycol, 17, 345-346.

    Henricot, B., Denton, J., Scrace, J., Barnes, A. V., Lane, C. R., 2010. Peronospora belbahrii causing downy mildew disease on Agastache in the UK: a new host and location for the pathogen. Plant Pathology, 59(4), 801. doi: 10.1111/j.1365-3059.2010.02264.x

    Hoffmeister M, Ashrafi S, Thines M, Maier W, 2020. Two new species of the Peronospora belbahrii species complex, Pe. choii sp. nov. and Pe. salvia-pratensis sp. nov., and a new host for Pe. salviae-officinalis. Fungal Systematics and Evolution, 6, 39-53.

    Hu, B., Li, Z., Hu, M., Sun, H., Zheng, J., Diao, Y., 2018. Outbreak of downy mildew caused by Peronospora belbahrii on Ocimum basilicum var. polosum in China. New Disease Reports, 37, 1. doi: 10.5197/j.2044-0588.2018.037.001

    Ito, Y., Takeuchi, T., Matsushita, Y., Chikuo, Y., Satou, M., 2015. Downy mildew of coleus caused by Peronospora belbahrii in Japan. Journal of General Plant Pathology, 81(4), 328-330.

    Kanetis L, Vasiliou A, Neophytou G, Samouel S, Tsaltas D, 2014. First report of downy mildew caused by Peronospora belbahrii on sweet basil (Ocimum basilicum) in Cyprus. Plant Disease, 98(2):283. http://apsjournals.apsnet.org/loi/pdis

    Koroch, A. R., Villani, T. S., Pyne, R. M., Simon, J. E., 2013. Rapid staining method to detect and identify downy mildew (Peronospora belbahrii) in basil. Applications in Plant Sciences, 1(7), 1300032. doi: 10.3732/apps.1300032

    Liberato, J. R., Forsberg, L., Grice, K. R., Shivas, R. G., 2006. Peronospora lamii on Lamiaceae in Australia. Australasian Plant Pathology, 35(3), 367-368. doi: 10.1071/AP06027

    López-López, A., Koller, M., Herb, C., Schärer, H. J., 2014. Influence of light management on the sporulation of downy mildew on sweet basil. Acta Horticulturae, (No.1041), 213-219. http://www.actahort.org/books/1041/1041_24.htm

    Martínez de la Parte, E., Pérez-Vicente, L., Bernal, B., García, D., 2010. First report of Peronospora sp. on sweet basil (Ocimum basilicum) in Cuba. Plant Pathology, 59(4), 800. doi: 10.1111/j.1365-3059.2009.02249.x

    Martini, P, Rapetti, S, Bozzano, G, Bassetti, G, 2003. (Segnalazione in Italia di Peronospora lamii su basilico (Ocimum basilicum L.)). In: Atti del Convegno ‘Problemi fitopatologici emergenti e implicazioni per la difesa delle colture’ . 79–82.

    McLeod, A., Coertze, S., Mostert, L., 2006. First report of a Peronospora species on sweet basil in South Africa. Plant Disease, 90(8), 1115.

    Nagy G, Horváth A, 2011. Occurrence of downy mildew caused by Peronospora belbahrii on sweet basil in Hungary. Plant Disease, 95(8):1034. http://apsjournals.apsnet.org/loi/pdis

    Palmateer, A. J., Harmon, P. F., Schubert, T. S., 2008. Downy mildew of coleus (Solenostemon scutellarioides) caused by Peronospora sp. in Florida. Plant Pathology, 57(2), 372. doi: 10.1111/j.1365-3059.2007.01763.x

    Patton A, 1992. A synthesis of Ocimum L. (labiate) in Africa. Kew Bulletin, 47, 403-435.

    Petrzelová I, Kitner M, Dolezalová I, Ondrej V, Lebeda A, 2015. First report of basil downy mildew caused by Peronospora belbahrii in the Czech Republic. Plant Disease, 99(3):418. http://apsjournals.apsnet.org/loi/pdis

    Riley, E. A., 1960. A revised list of plant diseases in Tanganyika Territory. In: Mycological Papers , (75) . 42 pp.

    Rivera, Y., Salgado-Salazar, C., Creswell, T. C., Ruhl, G., Crouch, J. A., 2016. First report of downy mildew caused by Peronospora sp. on Agastache sp. in the United States. Plant Disease, 100(6), 1249-1250. doi: 10.1094/PDIS-10-15-1119-PDN

    Rivera, Y., Salgado-Salazar, C., Windham, A. S., Crouch, J. A., 2016. Downy mildew on coleus (Plectranthus scutellarioides) caused by Peronospora belbahrii sensu lato in Tennessee. Plant Disease, 100(3), 655. doi: 10.1094/PDIS-10-15-1120-PDN

    Roberts, P. D., Raid, R. N., Harmon, P. F., Jordan, S. A., Palmateer, A. J., 2009. First report of downy mildew caused by a Peronospora sp. on Basil in Florida and the United States. Plant Disease, 93(2), 199. doi: 10.1094/PDIS-93-2-0199B

    Ronco, L., Rollán, C., Choi, Y. J., Shin, H. D., 2009. Downy mildew of sweet basil (Ocimum basilicum) caused by Peronospora sp. in Argentina. Plant Pathology, 58(2), 395. doi: 10.1111/j.1365-3059.2008.02006.x

    Salgado-Salazar C, LeBlanc N, Wallace EC, Daughtrey ML, Crouch JA, 2020. Peronospora monardae, Hyaloperonospora daughtreyae and H. iberidis: New species associated with downy mildew diseases affecting ornamental plants in the United States. European Journal of Plant Pathology, doi: 10.1007/s10658-020-01989-9

    Saude C, Westerveld S, Filotas M, McDonald MR, 2013. First report of downy mildew caused by Peronospora belbahrii on basil (Ocimum spp.) in Ontario. Plant Disease, 97(9):1248. http://apsjournals.apsnet.org/loi/pdis

    Smith, S., Urrea, K., 2018. First report of downy mildew caused by Peronospora belbahrii on sweet basil (Ocimum basilicum) in Arkansas. Plant Disease, 102(3), 686. http://apsjournals.apsnet.org/loi/pdis doi: 10.1094/pdis-10-17-1586-pdn

    Thines, M., Telle, S., Ploch, S., Runge, F., 2009. Identity of the downy mildew pathogens of basil, coleus, and sage with implications for quarantine measures. Fungal Biology, 113(5), 532-540. doi: 10.1016/j.mycres.2008.12.005

    Voglmayr, H., Piątek, M., 2009. Peronospora causing downy mildew disease of sweet basil newly reported in Cameroon. Plant Pathology, 58(4), 805.

    Wick, R. L., Brazee, N. J., 2009. First report of downy mildew caused by a Peronospora species on sweet basil (Ocimum basilicum) in Massachusetts. Plant Disease, 93(3), 318. doi: 10.1094/PDIS-93-3-0318B

    Wyenandt, CA, Simon, JE, Pyne, RM, Homa, K, McGrath, MT, Zhang, S, Raid, RN, Ma, L-J, Wick, R, Guo, L, Madeiras, A, 2015. Basil downy mildew (Peronospora belbahrii): Discoveries and challenges relative to its control. Phytopathology, 105, 885-894.

    Yerkes, W. D. , Shaw, C. G. , 1959. Taxonomy of the Peronospora species on Cruci-ferae and Chenopodiaceae. Phytopathology, 49(8), 499-507 pp.

    Distribution References

    Anon, 2011. Basil downy mildew Peronospora belbahrii. In: Alien Pest Alert! Hawaii, College of Tropical Agriculture and Human Resources, University of Hawai'i at Manoa. https://www.ctahr.hawaii.edu/site/downloads/BDM/factsheet.pdf

    Belbahri L, Calmin G, Pawlowski J, Lefort F, 2005. Phylogenetic analysis and Real Time PCR detection of a presumbably undescribed Peronospora species on sweet basil and sage. Mycological Research. 109 (11), 1276-1287. DOI:10.1017/S0953756205003928

    Blomquist C L, Rooney-Latham S, Nolan P A, 2009. First report of downy mildew on field-grown sweet basil caused by a Peronospora sp. in San Diego County, California. Plant Disease. 93 (9), 968. DOI:10.1094/PDIS-93-9-0968A

    CABI, 2020. CABI Distribution Database: Status as determined by CABI editor. Wallingford, UK: CABI

    CABI, EPPO, 2015. Peronospora belbahrii. [Distribution map]. In: Distribution Maps of Plant Diseases, Wallingford, UK: CABI. Map 1177 (Edition 1). DOI:10.1079/DMPD/20153399811

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

    Chen C H, Huang J H, Hsieh T F, 2010. First report of Peronospora belbahrii causing downy mildew on basil. Plant Pathology Bulletin. 19 (3), 177-180.

    Choi Y-J, Choi IY, Lee KJ, Shin H-D, 2016. First report of downy mildew caused by Peronospora belbahrii on sweet basil (Ocimum basilicum) in Korea. Plant Disease. 2335.

    Cohen Y, Vaknin M, Ben-Naim Y, Rubin A E, Galperin M, Silverman D, Bitton S, Adler U, 2013. First report of the occurrence and resistance to mefenoxam of Peronospora belbahrii, causal agent of downy mildew of basil (Ocimum basilicum) in Israel. Plant Disease. 97 (5), 692. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-12-12-1126-PDN

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    Denton G J, Beal E, Denton J O, Clover G, 2015. First record of downy mildew, caused by Peronospora belbahrii, on Solenostemon scutellarioides in the UK. New Disease Reports. 14. http://www.ndrs.org.uk/article.php?id=031014 DOI:10.5197/j.2044-0588.2015.031.014

    EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. https://gd.eppo.int/

    Garibaldi A, Minuto A, Gullino M L, 2005. First report of downy mildew caused by Peronospora sp. on basil (Ocimum basilicum) in France. Plant Disease. 89 (6), 683. DOI:10.1094/PD-89-0683C

    Garibaldi A, Minuto A, Minuto G, Gullino M L, 2004. First report of downy mildew on basil (Ocimum basilicum) in Italy. Plant Disease. 88 (3), 312. DOI:10.1094/PDIS.2004.88.3.312A

    Gómez Tenorio MA, Lupión Rodríguez B, Boix Ruiz A, Ruiz Olmos C, Moreno Díaz A, Marín Guirao JI, Pérez Molina G, García Raya P, Tello Maarquina JC, 2016. (El mildiu nueva enfermedad de la albahaca en España). Phytoma. 48-52.

    Gumedzoe MYD, Hemou P, Lepage A, Lecat N, 1998. (Inventaire et identification de Peronospora lamii (Al. Br.) de By, agent causal du mildiou du basilic (Ocimum basilicum) dans les exploitations de Gyma Cultures). In: Les VIIIe Journe´es Scientifiques de l’Universite´ du Be´nin, 11–15 mai 1998 [Les VIIIe Journe´es Scientifiques de l’Universite´ du Be´nin, 11–15 mai 1998], Lome, Benin:

    Hafellner J, 2006. (Der Falsche Mehltaupilz auf Basilikum nun auch in Österreich eingeschleppt). Fritschiana (Graz). 29-34.

    Hansford CG, 1933. Annual report of the mycologist. Rev. Appl. Mycol. 421-422.

    Hansford CG, 1939. Annual report of the mycologist. Rev. Appl. Mycol. 345-346.

    Heller W, Baroffio C, 2003. (Le mildiou (Peronospora lamii) dubasilic progresse!). Der Gemüsebau/Le Maraîcher. 12-13.

    Hu B, Li Z, Hu M, Sun H, Zheng J, Diao Y, 2018. Outbreak of downy mildew caused by Peronospora belbahrii on Ocimum basilicum var. polosum in China. New Disease Reports. 1. DOI:10.5197/j.2044-0588.2018.037.001

    Ito Y, Takeuchi T, Matsushita Y, Chikuo Y, Satou M, 2015. Downy mildew of coleus caused by Peronospora belbahrii in Japan. Journal of General Plant Pathology. 81 (4), 328-330. http://link.springer.com/article/10.1007%2Fs10327-015-0601-3

    Joshi V, Jeffries M, Jesperson G, 2013. 2013 Canadian plant disease survey. Canadian Plant Disease Survey. 7-32.

    Kanetis L, Vasiliou A, Neophytou G, Samouel S, Tsaltas D, 2014. First report of downy mildew caused by Peronospora belbahrii on sweet basil (Ocimum basilicum) in Cyprus. Plant Disease. 98 (2), 283. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-07-13-0759-PDN

    Khateri H, Calmin G, Moarrefzadeh N, Belbahri L, Lefort F, 2007. First report of downy mildew caused by Peronospora sp. on basil in Northern Iran. Journal of Plant Pathology. 89 (Suppl.3), S70. http://www.agr.unipi.it/sipav/jpp/index.html

    Kong X Y, Wang S, Wan S L, Xiao C L, Luo F, Liu Y, 2015. First report of downy mildew on basil (Ocimum basilicum) in China. Plant Disease. 99 (11), 1642-1643. DOI:10.1094/PDIS-01-15-0077-PDN

    Lefort F, Gigon V, Amos B, 2003. (Le mildiou s’étend. Déjà détecté dans de nombreux pays européens, Peronospora lamii responsable du mildiou du basilic, a été observé en Suisse dans larégion lémanique). Réussir Fruits et Légumes. 66.

    Martínez de la Parte E, Pérez-Vicente L, Bernal B, García D, 2010. First report of Peronospora sp. on sweet basil (Ocimum basilicum) in Cuba. Plant Pathology. 59 (4), 800. DOI:10.1111/j.1365-3059.2009.02249.x

    Martini P, Rapetti S, Bozzano G, Bassetti G, 2003. (Segnalazione in Italia di Peronospora lamii su basilico (Ocimum basilicum L.)). In: Atti del Convegno ‘Problemi fitopatologici emergenti e implicazioni per la difesa delle colture’ [Atti del Convegno ‘Problemi fitopatologici emergenti e implicazioni per la difesa delle colture’, Sanremo IM 27–29 Novembre 2003], 79–82.

    McLeod A, Coertze S, Mostert L, 2006. First report of a Peronospora species on sweet basil in South Africa. Plant Disease. 90 (8), 1115. DOI:10.1094/PD-90-1115A

    Nagy G, Horváth A, 2011. Occurrence of downy mildew caused by Peronospora belbahrii on sweet basil in Hungary. Plant Disease. 95 (8), 1034. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-04-11-0329

    Novak A, Sever Z, Ivić D, Čajkulić A M, 2016. Basil downy mildew (Peronospora belbahrii) - destructive disease in basil production. (Plamenjača bosiljka (Peronospora belbahrii) - destruktivna bolest u proizvodnji bosiljka.). Glasilo Biljne Zaštite. 16 (6), 544-547. http://www.hdbz.hr

    Petrželová I, Kitner M, Doležalová I, Ondřej V, Lebeda A, 2015. First report of basil downy mildew caused by Peronospora belbahrii in the Czech Republic. Plant Disease. 99 (3), 418. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-06-14-0613-PDN

    Porta-Puglia A, Mifsud D, 2006. First record of downy mildew caused by Peronospora sp. on basil in Malta. Journal of Plant Pathology. 88 (1), 124. http://www.agr.unipi.it/sipav/jpp/index.html

    Riley E A, 1960. A revised list of plant diseases in Tanganyika Territory. In: Mycological Papers, 42 pp.

    Roberts P D, Raid R N, Harmon P F, Jordan S A, Palmateer A J, 2009. First report of downy mildew caused by a Peronospora sp. on Basil in Florida and the United States. Plant Disease. 93 (2), 199. DOI:10.1094/PDIS-93-2-0199B

    Romero Bastidas M, Saucedo Picazo L, Murillo Amador B, Nieto Garibay A, Latisnere Barragan H, Hernandez Montiel L G, 2016. First report of Peronospora belbahrii on sweet basil in Baja California Sur, México. Journal of Phytopathology. 164 (2), 122-124. DOI:10.1111/jph.12391

    Ronco L, Rollán C, Choi Y J, Shin H D, 2009. Downy mildew of sweet basil (Ocimum basilicum) caused by Peronospora sp. in Argentina. Plant Pathology. 58 (2), 395. DOI:10.1111/j.1365-3059.2008.02006.x

    Šafránková I, Holková L, 2014. The first report of downy mildew caused by Peronospora belbahrii on sweet basil in greenhouses in the Czech Republic. Plant Disease. 98 (11), 1579. DOI:10.1094/PDIS-07-13-0747-PDN

    Saude C, Westerveld S, Filotas M, McDonald M R, 2013. First report of downy mildew caused by Peronospora belbahrii on basil (Ocimum spp.) in Ontario. Plant Disease. 97 (9), 1248. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-01-13-0026-PDN

    Silva A L, Mansur P S C, Barreto R W, Salcedo-Sarmiento S, 2019. Peronospora belbahrii causes downy mildew on Ocimum basilicum in Brazil. Australasian Plant Disease Notes. 14 (1), 37. https://link.springer.com/article/10.1007/s13314-019-0368-z

    Smith S, Urrea K, 2018. First report of downy mildew caused by Peronospora belbahrii on sweet basil (Ocimum basilicum) in Arkansas. Plant Disease. 102 (3), 686. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/pdis-10-17-1586-pdn

    Thines M, Telle S, Ploch S, Runge F, 2009. Identity of the downy mildew pathogens of basil, coleus, and sage with implications for quarantine measures. Fungal Biology. 113 (5), 532-540. DOI:10.1016/j.mycres.2008.12.005

    Topolovec-Pintarić S, Martinko K, 2020. Downy mildew of basil: A new destructive disease worldwide. In: Plant Disease - Current Threats and Management Trends. [ed. by Topolovec-Pintarić P]. UK, London: InTechOpen. 1-15.

    Voglmayr H, Piątek M, 2009. Peronospora causing downy mildew disease of sweet basil newly reported in Cameroon. Plant Pathology. 58 (4), 805. DOI:10.1111/j.1365-3059.2009.02047.x

    Webb K, Sansford C, McLeod A, Matthews-Berry S, 2012. Rapid assessment of the need for a detailed Pest Risk Analysis for Peronospora belbahrii. In: UK Risk Register Details for Peronospora belbahrii. UK Plant Health Risk Register (DEFRA), https://secure.fera.defra.gov.uk/phiw/riskRegister/viewPestRisks.cfm?cslref=26813

    Wick R L, Brazee N J, 2009. First report of downy mildew caused by a Peronospora species on sweet basil (Ocimum basilicum) in Massachusetts. Plant Disease. 93 (3), 318. DOI:10.1094/PDIS-93-3-0318B

    Wyenandt CA, Simon JE, Pyne RM, Homa K, McGrath MT, Zhang S, Raid RN, Ma L-J, Wick R, Guo L, Madeiras A, 2015. Basil downy mildew (Peronospora belbahrii): Discoveries and challenges relative to its control. Phytopathology. 885-894.

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    12/05/20 Original text by:

    Jo Anne Crouch, USDA-ARS, 10300 Baltimore Avenue, Beltsville, MD 20715, USA.

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