Peronospora belbahrii (basil downy mildew)
Index
- Pictures
- Identity
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution
- Distribution Table
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- Symptoms
- List of Symptoms/Signs
- Biology and Ecology
- Means of Movement and Dispersal
- Seedborne Aspects
- Economic Impact
- References
- Principal Source
- Contributors
- Distribution Maps
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Top of pageIdentity
Top of pagePreferred Scientific Name
- Peronospora belbahrii Thines
Preferred Common Name
- basil downy mildew
Summary of Invasiveness
Top of pageBasil 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
Top of page- Domain: Eukaryota
- Kingdom: Chromista
- Phylum: Oomycota
- Class: Oomycetes
- Order: Peronosporales
- Family: Peronosporaceae
- Genus: Peronospora
- Species: Peronospora belbahrii
Notes on Taxonomy and Nomenclature
Top of pagePeronospora 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
Top of pageBasil 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
Top of pageThe 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: 12 May 2022Continent/Country/Region | Distribution | Last Reported | Origin | First Reported | Invasive | Reference | Notes |
---|---|---|---|---|---|---|---|
Africa |
|||||||
Benin | Present | 1998 | Invasive | ||||
Cameroon | Present | 2007 | Invasive | Plantations in Obala town (Centre Province), Cameroon. | |||
South Africa | Present | 2005 | Invasive | Western Cape Province. | |||
Tanzania | Present | ||||||
Uganda | Present | 1932 | |||||
Asia |
|||||||
China | Present, Localized | ||||||
-Beijing | Present, Localized | 2016 | Invasive | Shunyi and Daxing districts of Beijing. | |||
-Hainan | Present | 2014 | Invasive | Sanya City (18°N) of Hainan Province. | |||
Iran | Present, Widespread | 2006 | Invasive | ||||
Israel | Present, Widespread | 2011: Found in two greenhouses in the northern Jordan Valley, then at the southwest and southeast Israeli borders. Summer 2012: disease widespread in Israel. | |||||
Japan | Present | ||||||
-Honshu | Absent, Invalid presence record(s) | This report of Peronospora belbahrii from coleus is most likely P. choii (see Hoffmeister et al., 2020). | |||||
South Korea | Present, Localized | 2015 | Invasive | ||||
Taiwan | Present | 2009 | Invasive | Nantu and Yunlin. | |||
Europe |
|||||||
Austria | Present | 2005 | Invasive | Styria: East Styrian Hills, Graz, Ragnitztal on the eastern outskirts. | |||
Belgium | Present | 2004 | Invasive | ||||
Croatia | Present, Widespread | 2012 | Invasive | Dubrovnik-Neretva, Varaždin, Krapina-Zagorje, Medimurje, Split-Dalmatian and Zagreb Counties. | |||
Cyprus | Present, Localized | 2012 | Invasive | ||||
Czechia | Present, Widespread | 2012 | Invasive | Central Bohemia. | |||
France | Present, Few occurrences | 2004 | Invasive | ||||
Germany | Present | 2005 | Invasive | Lankreis, Heilbronn. | |||
Hungary | Present, Few occurrences | 2003 | Invasive | Albertirsa, Budapest-Soroksár, Tordas. | |||
Italy | Present | 2003 | Invasive | ||||
Malta | Present | 2005 | Invasive | ||||
Spain | Present | 2016 | Invasive | ||||
-Canary Islands | Present | ||||||
Switzerland | Present | 2002 | Invasive | Eastern Switzerland, island of Reichenau. | |||
United Kingdom | Present, Localized | 2005 | Invasive | ||||
-England | Present, Localized | ||||||
North America |
|||||||
Canada | Present, Localized | ||||||
-British Columbia | Present | 2011 | Invasive | ||||
-Ontario | Present, Few occurrences | 2011 | Invasive | ||||
-Quebec | Present | 2011 | Invasive | ||||
Cuba | Present | 2009 | Invasive | Horticulture Research Institute, Liliana Dimitrova in Quivican, Havana province and Siboney, Havana City. | |||
Mexico | Present, Localized | 2009 | Invasive | ||||
United States | Present, Widespread | ||||||
-Alabama | Present | 2010 | Invasive | ||||
-Alaska | Present | 2011 | Invasive | ||||
-Arkansas | Present | 2008 | Invasive | ||||
-California | Present, Widespread | 2008 | Invasive | ||||
-Colorado | Present, Widespread | 2012 | Invasive | ||||
-Connecticut | Present | 2009 | Invasive | ||||
-Delaware | Present | 2009 | Invasive | ||||
-District of Columbia | Present | 2009 | Invasive | ||||
-Florida | Present, Widespread | 2007 | Invasive | Collier, Hendry, Miami-Dade, and Palm Beach counties in south Florida; Hillsborough County in west-central north Florida. | |||
-Georgia | Present | 2012 | Invasive | ||||
-Hawaii | Present, Widespread | 2010 | Invasive | Wai?anac Oahu Maui Molokai Hawaii | |||
-Illinois | Present | 2009 | Invasive | ||||
-Indiana | Present | 2009 | Invasive | ||||
-Iowa | Present | 2014 | Invasive | ||||
-Kansas | Present | 2008 | Invasive | ||||
-Kentucky | Present, Widespread | 2010 | Invasive | ||||
-Louisiana | Present | 2009 | Invasive | ||||
-Maine | Present | 2012 | Invasive | ||||
-Maryland | Present, Widespread | 2009 | Invasive | ||||
-Massachusetts | Present, Widespread | 2008 | Invasive | ||||
-Michigan | Present | 2010 | Invasive | ||||
-Minnesota | Present | 2011 | Invasive | ||||
-Mississippi | Present | 2016 | Invasive | ||||
-Missouri | Present, Widespread | 2012 | Invasive | ||||
-Montana | Present, Widespread | 2010 | Invasive | ||||
-Nebraska | Present | 2014 | Invasive | ||||
-New Hampshire | Present | 2012 | Invasive | ||||
-New Jersey | Present, Widespread | 2008 | Invasive | ||||
-New Mexico | Present | 2013 | Invasive | ||||
-New York | Present, Widespread | 2008 | Invasive | ||||
-North Carolina | Present | 2008 | Invasive | ||||
-North Dakota | Present | 2008 | Invasive | ||||
-Ohio | Present | 2010 | Invasive | ||||
-Oklahoma | Present | 2016 | Invasive | ||||
-Oregon | Present | 2012 | Invasive | ||||
-Pennsylvania | Present, Widespread | 2009 | Invasive | ||||
-Rhode Island | Present | 2012 | Invasive | ||||
-South Carolina | Present | 2009 | Invasive | ||||
-Tennessee | Present | 2013 | Invasive | ||||
-Texas | Present | 2009 | Invasive | ||||
-Vermont | Present, Widespread | 2009 | Invasive | ||||
-Virginia | Present | 2010 | Invasive | ||||
-Washington | Present | 2011 | Invasive | ||||
-West Virginia | Present | 2012 | Invasive | ||||
-Wisconsin | Present | 2009 | Invasive | ||||
Oceania |
|||||||
New Zealand | Present | ||||||
South America |
|||||||
Argentina | Present | 2008 | Invasive | La Plata | |||
Brazil | Present | ||||||
-Minas Gerais | Absent, Unconfirmed presence record(s) | This report of Peronospora belbahrii from coleus is most likely P. choii (see Hoffmeister et al., 2020). | |||||
-Sao Paulo | Present, Few occurrences |
Risk of Introduction
Top of pageThere 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
Top of pageCategory | Sub-Category | Habitat | Presence | Status |
---|---|---|---|---|
Terrestrial | Managed | Cultivated / agricultural land | Present, no further details | |
Terrestrial | Managed | Protected agriculture (e.g. glasshouse production) | Present, no further details |
Hosts/Species Affected
Top of pagePrior 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
Top of pagePlant name | Family | Context | References |
---|---|---|---|
Ocimum basilicum (basil) | Lamiaceae | Main |
Symptoms
Top of pageInitial 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
Top of pageSign | Life Stages | Type |
---|---|---|
Leaves / abnormal colours | ||
Leaves / abnormal forms | ||
Leaves / abnormal leaf fall | ||
Leaves / fungal growth | ||
Leaves / necrotic areas | ||
Leaves / yellowed or dead |
Biology and Ecology
Top of pageP. 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
Top of pageThere 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
Top of pageThere is evidence that the pathogen is transmitted through seeds (Cohen et al., 2017).
Economic Impact
Top of pageBasil 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. 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
Top of pageEPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
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.
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.
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.
Patton A, 1992. A synthesis of Ocimum L. (labiate) in Africa. Kew Bulletin, 47, 403-435.
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
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.
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
CABI, 2020. CABI Distribution Database: Status as determined by CABI editor. Wallingford, UK: CABI
CABI, Undated. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
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.
Coosemans J, 2004. First report of Peronospora lamii, downy mildew on basil (Ocimum basilicum) in Belgium. Parasitica. 27.
Cornell University, 2020. Where in the USA is Basil Downy Mildew? Long Island, New York, USA: Cornell University, College of Agriculture & Life Sciences. http://blogs.cornell.edu/livegpath/extension/basil-downy-mildew/where-in-the-usa-is-basil-downy-mildew/
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.
Joshi V, Jeffries M, Jesperson G, 2013. 2013 Canadian plant disease survey. Canadian Plant Disease Survey. 7-32.
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.
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.
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.
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
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.
Contributors
Top of page12/05/20 Original text by:
Jo Anne Crouch, USDA-ARS, 10300 Baltimore Avenue, Beltsville, MD 20715, USA.
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