Cocksfoot mottle virus (Cocksfoot mottle virus)
- Summary of Invasiveness
- Taxonomic Tree
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Means of Movement and Dispersal
- Seedborne Aspects
- Pathway Causes
- Pathway Vectors
- Vectors and Intermediate Hosts
- Impact Summary
- Economic Impact
- Environmental Impact
- Risk and Impact Factors
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Principal Source
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Cocksfoot mottle virus Tamm and Truve (2000a)
Preferred Common Name
- Cocksfoot mottle virus
Other Scientific Names
- Cocksfoot mottle sobemovirus Mäkinen et al. (1995a, b)
Summary of InvasivenessTop of page
Cocksfoot mottle virus (CfMV) is distributed in the Eastern and Western hemispheres. Invasiveness through introduction outside of Europe is probable.
Taxonomic TreeTop of page
- Domain: Virus
- Family: Unassigned virus family
- Genus: Sobemovirus
- Species: Cocksfoot mottle virus
DescriptionTop of page
Cocksfoot mottle is a positive-strand RNA virus with an isometric particle ~30 nm in diameter (Serjeant, 1964; Serjeant, 1967). Purified virion can remain infective for several months when stored at -10ºC, however, sap infectivity was lost after being heated to 65ºC for 10 minutes (Serjeant, 1967). The virion has an T = 3 symmetry comprised of 180 coat protein monomers (Tars et al., 2003).
CfMV has a viral protein genome (VPg) covalently linked to the 5’ end of a 4082 nucleotide RNA molecule. The genome encodes an open reading frame on the 5’ proximal end that encodes the movement protein/silencing suppressor (12 kDa), of which leaky scanning leads to the expression of polyproteins P2a (71 kDa) and P2b (Tamm et al., 1999). P2b (100 kDa) encodes the RNA dependent RNA polymerase, which is translated through a -1 programmed frameshift mechanism (Mäkinen et al., 1995a; Tamm et al., 1999). A 12-nucleotide stem loop structure in the RNA is indispensable for the -1 ribosomal frameshift (Tamm et al., 2009). The coat protein (34kDa) is expressed via sub-genomic RNA from a 3’ proximal ORF (Mäkinen et al., 1995b; Olspert et al., 2014).
Distribution TableTop of page
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|
|Russia||Present||Detected in Moscow region.|
|United Kingdom||Present||East Anglia, Midlands.|
|-California||Present, Few occurrences||Introduced||2017|
|New Zealand||Present, Few occurrences||Introduced||Invasive||Detected on both North and South Island.|
History of Introduction and SpreadTop of page
Cocksfoot mottle virus (CfMV) was first detected in orchardgrass (Dactylis glomerata), also known as cocksfoot, in the UK in 1964 (Serjeant, 1964). It was subsequently reported in other countries including Denmark (Engsbro, 1978), Germany (Rabenstein and Schmidt, 1979), New Zealand (Mohamed, 1980; Campbell and Guy, 2001), Japan (Toriyama, 1982), Norway (Munthe, 1988; Mäkinen et al., 1995a; Mäkinen et al., 1995b), Russia (Ryabov et al., 1996) and Poland (Trzmiel and Jeżewska, 2017). In North America CfMV is known to be present in Canada (Bittman et al., 2006), but the date of first detection is unknown. Recently, the virus has been detected in the USA in Oregon (Alderman et al., 2016; Gilmore et al., 2017), California (Martin et al., 2018) and Ohio (Hodge et al., 2018).
There is evidence based on surveys that CfMV was introduced to New Zealand and became widespread there since 1976 (Mohamed, 1980; Campbell and Guy, 2001). Orchardgrass and Italian ryegrass (Lolium multiflorum), two of the major hosts of CfMV, are both native to Eurasia and North Africa and were introduced to Japan (Holm et al., 1997; Sugiyama, 2003), New Zealand and North America (Beddows, 1959; Lumaret al., 1988; Holm et al., 1997) in the 1600-1800s. Additionally, the cereal leaf beetle (Oulema melanopus), vector of CfMV, is native to Europe and Asia and was introduced to the USA in 1962 (Philips et al., 2011) and became established in North American (Kher et al., 2011). This suggests that CfMV may also be native to Europe and/or Asia and introduced to other parts of the world with hosts and vectors.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|New Zealand||1976-2001||Yes||No||Mohamed (1980); Campbell and Guy (2001)|
Risk of IntroductionTop of page
Major hosts of CfMV, orchardgrass (Dactyllis glomerata) and Italian ryegrass (Lolium multiflorum), are both widespread in most regions of the world (Holm et al., 1997), thus new introductions of CfMV through new introductions of the host is unlikely. The cereal leaf beetle (Oulema melanopus), however, has a reported distribution currently restricted to North America, Canada, Europe, and reported in India, Pakistan and Iran (Bailey et al., 1991; Kher et al., 2011; Philips et al., 2011). It was not reported in New Zealand, where evidence suggests that the virus has spread by means of mechanical inoculation by machinery (Campbell and Guy, 2001). Thus, the risk of introduction of CfMV into new areas follows the risk of introduction of cereal leaf beetle with virus or of the virus alone into new regions. Cereal leaf beetle has received a medium rating of risk of introduction to new areas in California from the US APHIS risk assessment programme, in part because the vector mates after overwintering and only produces one generation per year (Kher et al., 2011; Philips et al., 2011; Leathers, 2015). Canada assigned cereal leaf beetle a low risk of introduction into new areas within Canada due to the beetle’s inability to withstand harsh winters or hot summers (CFIA, 2008). Cereal leaf beetle is not expected to expand its range into areas with warmer climates or harsh winters and therefore expansion of the range of CfMV is unlikely as well. Suggested pathways of spread of cereal leaf beetle include through transportation of grain, forage seed, straw, hay, grass sod, plant litter, used harvest equipment, and cut or balled Christmas trees (CFIA, 2008; Leathers, 2015).
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Principal habitat|
|Terrestrial||Managed||Managed grasslands (grazing systems)||Principal habitat|
|Terrestrial||Managed||Industrial / intensive livestock production systems||Principal habitat|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Principal habitat|
Hosts/Species AffectedTop of page
CfMV has primarily been reported as a pathogen affecting orchardgrass (Dactylis glomerata), but natural infections have been reported on wheat (Serjeant, 1967). The experimental host range of CfMV includes orchardgrass (D. glomerata), wheat (Triticum aestivum), oats (Avena sativa), barley (Hordeum vulgare), rye (Secale cereale), Agrostis puchella [A. tenerrima], Anthoxanthum puelii [A. aristatum], Bromus secalinus, Lagurus ovatus, Lamarckia aurea, Phalaris minor, P. paradoxa, Phleum arenarium and P. paniculatum (Serjeant, 1967; Catherall et al., 1977). Plants identified as non-hosts include rice (Oryza sativa), maize (Zea mays) and the grasses Agropyron repens [Elymus repens], Alopecurus pratensis, Anthoxanthum odoratum, Apera spica-venti, Brachypodium pinnatum, Bromus arvensis, B. mollis [B. hordeaceus], Cynosurus cristatus, Festuca arundinacea, Holcus lanatus, Lolium perenne, Poa pratensis, P. trivialis, Sorghum halepense and S. vulgare [S. bicolor], as well as other monocotyledonous species Lilium formosum [L. sargentiae], L. regale and Musa balbisiana, and dicotyledonous species Chenopodium amaranticolor [C. giganteum], C. hybridum, C. quinoa, Cucumis sativus, Datura stramonium, Nicotiana clevelandii, N. glutinosa, N. tabacum, Petunia hybrida, Phaseolus vulgaris and Vicia faba (Serjeant, 1967; Catherall et al., 1977).
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
CfMV typically produces mosaic/mottling and stunting symptoms on orchardgrass (Dactylis glomerata) and wheat that turn white or necrotic as the leaves age (Serjeant, 1967). CfMV has been described to commonly cause necrotic leaf tips in orchardgrass, and heavily symptomatic plants are killed by the infection (Rognli et al., 1995; Catherall et al., 1977). There are some reports that infection of wheat seedlings will eventually lead to whole plant necrosis (Serjeant, 1967; Catherall et al., 1977). Symptoms in oats and barley are milder than in orchardgrass or wheat, with mild mottling that develops into necrotic lesions (Serjeant, 1967). Symptoms can take 2-3 weeks after inoculation to appear.
List of Symptoms/SignsTop of page
|Leaves / abnormal colours|
|Leaves / abnormal patterns|
|Whole plant / discoloration|
|Whole plant / dwarfing|
|Whole plant / plant dead; dieback|
Means of Movement and DispersalTop of page
CfMV has not been reported to be seed transmitted but is transmitted efficiently by cereal leaf beetles Oulema melanopus and O. lichensis [O. gallaeciana]. It is also transmitted by mechanical inoculation and is transmissible by cutting implements (Catherall and Potter, 1987), which may be a major mode of spread and establishment even in the absence of insect vectors (Campbell and Guy, 2001).
Vector Transmission (Biotic)
CfMV was not found to be transmitted by a variety of aphid species (Serjeant, 1964). Serjeant (1967) demonstrated transmission of CfMV by a high proportion of exposed adult cereal leaf beetle (O. melanopus), with ability to transmit acquired virus for up to 15 days. Larvae also transmitted the virus, but with much lower efficiency (Serjeant, 1967). Catherall (1970) and A’Brook and Beningo (1972) also reported transmission by O. lichensis [O. gallaeciana]. No latent period prior to transmission was observed after feeding, and transmission occurred only up to about 15 days post-acquisition, suggesting semi-persistent transmission (Catherall, 1970; A’Brook and Beningo, 1972), although A’Brook and Beningo (1972) suggested the possibility of circulation within viruliferous beetles as they reported virus in beetle haemolymph.
Seedborne AspectsTop of page
Incidences up to 55% were reported for CfMV in orchardgrass (Dactylis glomerata) seed cropping fields the East Midlands of England, UK, with virus incidence observed increasing with crop age (Upstone, 1969). In New Zealand, incidences of up to 80% were measured at South Island sites, with higher incidences at mowed than unmowed sites, and an establishment and increase in infection over 3 years (Campbell and Guy, 2001). Incidences high enough to be problematic in production in the Fraser Valley, Canada, were alluded to the resistance screening report by Bittman et al. (2006). Gilmore et al. (2017) reported incidences >45% and increasing incidences of CfMV in orchardgrass in Oregon, USA from 2014-2016.
Pathway CausesTop of page
Pathway VectorsTop of page
|Host and vector organisms||Frequency of introduction not known.||Yes||Yes|
Vectors and Intermediate HostsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
CfMV is reported to have the highest impact on orchardgrass (Dactylis glomerata) forage and seed production. In some areas of the UK, CfMV has been reported to be widespread and cause severe damage in orchardgrass (Upstone, 1969). CfMV is reported to reduce the number of tillers orchardgrass produces and the size of the plants, thus reducing the utilization of orchardgrass for forage (Serjeant, 1967; Upstone 1969). Severe infections of CfMV are lethal, which can impact seed and forage production, and can reduce the volume and germination rate of seed produced by the plants (Upstone, 1969). Upstone (1969) also reported that virus incidence of infection increased over time as the orchardgrass aged, indicating that the spread of CfMV is not restricted by plant development. CfMV has been found in co-infection with Cocksfoot streak virus (Smith, 1952) and Wheat streak mosaic virus (Hodge et al., 2018), but the impact on plant development and yield due to co-infection is unknown. Although natural infection of CfMV in wheat has been reported (Serjeant, 1967; Hodge et al., 2018; Hodge et al., 2020) economic impact in wheat is not reported.
Environmental ImpactTop of page
Due to CfMV causing lethality in orchardgrass (Dactylis glomerata) it can change the ecology and biodiversity in grasslands. It has been reported that in an orchardgrass-only sward that is infected with CfMV, the plants that die are filled in by neighbouring healthy plants to recolonize the vacant space (Catherall et al., 1977). However, in multi-species grasslands this may lead to a modification of the ecological composition of the sward. Catherall (1966) also reported that orchardgrass/clover leys examined at the Grassland Research Institute in Hurley, UK, in 1958 and 1950 were taken over by the clover following natural epidemics of CfMV. Upstone (1969) reports a similar story of CfMV spreading through leys through grazing/cutting of the grass.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Abundant in its native range
- Has propagules that can remain viable for more than one year
- Reproduces asexually
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Host damage
- Increases vulnerability to invasions
- Monoculture formation
- Negatively impacts agriculture
- Negatively impacts animal health
- Negatively impacts livelihoods
- Threat to/ loss of native species
- Antagonistic (micro-organisms)
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
DiagnosisTop of page
Virus symptoms including mosaic suggest the possibility of CfMV but are not sufficient to determine the pathogenic agent. CfMV can be definitively diagnosed using serological assays such as enzyme-linked immunosorbent assays (ELISA) or sequence-based amplification assays.
Similarities to Other Species/ConditionsTop of page
Cocksfoot streak virus produces symptoms very similar to Cocksfoot mottle virus and is also manually transmitted; however, Cocksfoot streak virus has filamentous particles, is transmitted by aphids (Myzus persicae and Hyalopterus humilis), and cannot be transmitted to oats, barley or wheat (Serjeant, 1964). Ryegrass mosaic virus is similar to CfMV in that it infects oats and orchardgrass; however, it does not infect wheat and is mite-transmitted (Serjeant, 1964). CfMV may be confused with Cocksfoot mild mosaic virus partially due to similar symptomology and isometric particles; however, Cocksfoot mild mosaic virus can readily be transmitted by aphids to Setaria italica (Huth, 1968). CfMV was found to be co-infected with Wheat streak mosaic virus (WSMV) in Ohio (Hodge et al., 2018), but WSMV does not infect orchardgrass.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
There are no currently recommended measures for CfMV infection prevention.
Cultural Control and Sanitary Measures
Mowing may efficiently spread CfMV even in the absence of cereal leaf beetle vectors (Catherall and Potter, 1987; Campbell and Guy, 2001) such that equipment sanitation may help reduce transmission. Host-plant resistance has been reported to decrease the impact of virus infection. Crop rotation with non-hosts may also reduce disease loads.
Host Resistance (Incl. Vaccination)
Orchardgrass resistant to CfMV has been selected for improved outcomes under infection conditions (Catherall and Potter, 1987; Rognli et al., 1995; Bittman et al., 2006). Catherall and Potter (1987) reported that some cultivars were less likely to become infected and exhibited better outcomes. Most resistance to CfMV in orchardgrass was reported to exhibit quantitative variation rather than single gene-conferred immunity to virus infection (Rognli et al., 1995). Such resistance to CfMV reported is alternatively termed ‘tolerance’ because virus replication is supported but yields are better under virus infection pressure than comparison genotypes (Rognli et al., 1995). Resistance selected by Bittman et al. (2006) improved yields up to 23% after artificial inoculation compared to non-resistant controls.
Gaps in Knowledge/Research NeedsTop of page
Most published data on CfMV incidence and yield impact are from previous decades and may not reflect current conditions. Field data-supported management recommendations are needed if current disease impacts warrant improved management. There is no information available regarding the optimal temperature requirements for CfMV infection and symptom development in its hosts. Further research is needed to examine the potential impact that CfMV has on wheat production in areas where it has been found, such as in Ohio, USA (Hodge et al., 2018; Hodge et al., 2020).
ReferencesTop of page
A'Brook, J., Benigno, D. A., 1972. The transmission of cocksfoot mottle and phleum mottle viruses by Oulema melanopa and O. lichenis. Annals of Applied Biology, 72(2), 169-176. doi: 10.1111/j.1744-7348.1972.tb01282.x
Alderman, S. C., Martin, R. C., Gilmore, B. S., Martin, R. R., Hoffman, G. D., Sullivan, C. S., Anderson, N. P., 2016. First report of Cocksfoot mottle virus infecting Dactylis glomerata in Oregon and the United States. Plant Disease, 100(5), 1030-1031. doi: 10.1094/PDIS-09-15-1017-PDN
Bailey, W. C., Carlson, C. E., Puttler, B., Stoltenow, C. R., 1991. Expansion of the range of the cereal leaf beetle, Oulema melanopus (L.) (Coleoptera: Chrysomelidae), in Missouri and Iowa. Journal of the Kansas Entomological Society, 64(4), 455-457.
Catherall PL, 1970. Cocksfoot mottle virus. In: Descriptions of Plant Viruses, (23) , UK: Association of Applied Biologists.http://www.dpvweb.net/dpv/showdpv.php?dpvno=023
Catherall, P. L., Andrews, P. A., Chamberlain, J. A., 1977. Host-ranges of cocksfoot mottle and cynosurus mottle viruses. Annals of Applied Biology, 87(2), 233-235. doi: 10.1111/j.1744-7348.1977.tb01880.x
Catherall, P. L., Potter, L. R., 1987. Effect of cocksfoot mottle virus on resistant and susceptible cocksfoot cultivars grown alone or in combination with healthy or ryegrass mosaic virus-infected Italian ryegrass. Annals of Applied Biology, 111(2), 345-351. doi: 10.1111/j.1744-7348.1987.tb01461.x
CFIA, 2008. Cereal leaf beetle (Oulema melanopus). In: Risk Management Document, CFIA Guidance Documents , Canada: Canadian Food Inspection Agency.
Gilmore, B. S., Martin, R. C., Dombrowski, J. E., Alderman, S. C., Martin, R. R., Mosier, N. J., Sullivan, C. S., Anderson, N. P., Hoffman, G. D., Guy, P. L., 2017. Virus incidence in orchardgrass (Dactylis glomerata L.) seed production fields in the Willamette Valley. Crop, Forage & Turfgrass Management, 3(1), cftm2016.12.0087. https://dl.sciencesocieties.org/publications/cftm/tocs/3/1
Hodge BA, Paul PA, Stewart LR, 2020. Occurrence and high throughput sequencing of viruses in Ohio wheat. Plant Disease, doi: Doi.org/10.1094/PDIS-08-19-1724-RE
Hodge, B. A., Paul, P. A., Stewart, L. R., 2018. First report of Cocksfoot mottle virus infecting wheat (Triticum aestivum) in Ohio. Plant Disease, 102(2), 464. http://apsjournals.apsnet.org/loi/pdis doi: 10.1094/PDIS-08-17-1224-PDN
Huth, W., 1968. Studies on a new virus of D. glomerata: Cocksfoot mild mosaic virus. (Untersuchungen über ein neues Virus von Dactylis glomerata: Cocksfoot mild mosaic virus). Phytopathologische Zeitschrift, 62(3), 300-303.
Kher SV, Dosdall LM, Carcamo HA, 2011. The cereal leaf beetle: biology, distribution, and prospects for control. Prairie Soil and Crops Journal, 4, 32-41.
Leathers J, 2015. Oulema melanopus: Cereal leaf beetle. California, USA: California Department of Food and Agriculture.https://blogs.cdfa.ca.gov/Section3162/?p=232
Martin, R. C., Martin, R. R., Putnam, M. L., 2018. First report of cocksfoot mottle virus infecting Dactylis glomerata in forage production fields in California. Plant Disease, 102(10), 2050-2051. doi: 10.1094/PDIS-02-18-0277-PDN
Mäkinen, K., Næss, V., Tamm, T., Truve, E., Aaspõllu, A., Saarma, M., 1995. The putative replicase of the cocksfoot mottle sobemovirus is translated as a part of the polyprotein by -1 ribosomal frameshift. Virology (New York), 207(2), 566-571. doi: 10.1006/viro.1995.1118
Mäkinen, K., Tamm, T., Næss, V., Truve, E., Puurand, Ü., Munthe, T., Saarma, M., 1995. Characterization of cocksfoot mottle sobemovirus genomic RNA and sequence comparison with related viruses. Journal of General Virology, 76(11), 2817-2825. doi: 10.1099/0022-1317-76-11-2817
Olspert, A., Kamsol, K., Sarmiento, C., Gerassimenko, J., Truve, E., 2014. Cocksfoot mottle virus coat protein is dispensable for the systemic infection. Virology Journal, 11(19), (4 February 2014). http://www.virologyj.com/content/pdf/1743-422X-11-19.pdf
Philips, C. R., Herbert, D. A., Kuhar, T. P., Reisig, D. D., Thomason, W. E., Malone, S., 2011. Fifty years of cereal leaf beetle in the U.S.: an update on its biology, management, and current research. Journal of Integrated Pest Management, 2(2), C1-C5. doi: 10.1603/IPM11014
Rabenstein, F., Schmidt, H. B., 1979. Identification of cocksfoot mottle virus in the GDR. (Nachweis des Knaulgrasscheckungs-Virus (cocksfoot mottle virus) in der DDR). Archiv fur Phytopathologie und Pflanzenschutz, 15(5), 351-354. doi: 10.1080/03235407909440482
Ryabov, E. V., Krutov, A. A., Novikov, V. K., Zheleznikova, O. V., Morozov, S. Yu., Zavriev, S. K., 1996. Nucleotide sequence of RNA from the Sobemovirus found in infected cocksfoot shows a luteovirus-like arrangement of the putative replicase and protease genes. Phytopathology, 86(4), 391-397. doi: 10.1094/Phyto-86-391
Tamm T, Suurväli, Luccesi J, Olspert A, Truve E, 2009. Stem-loop structure of Cocksfoot mottle virus RNA is indispensable for programmed -1 ribosomal frameshifting. Virus Research, 146, 73-80.
Alderman S C, Martin R C, Gilmore B S, Martin R R, Hoffman G D, Sullivan C S, Anderson N P, 2016. First report of Cocksfoot mottle virus infecting Dactylis glomerata in Oregon and the United States. Plant Disease. 100 (5), 1030-1031. DOI:10.1094/PDIS-09-15-1017-PDN
CABI, 2020. CABI Distribution Database: Status as determined by CABI editor. Wallingford, UK: CABI
Hodge B A, Paul P A, Stewart L R, 2018. First report of Cocksfoot mottle virus infecting wheat (Triticum aestivum) in Ohio. Plant Disease. 102 (2), 464. http://apsjournals.apsnet.org/loi/pdis DOI:10.1094/PDIS-08-17-1224-PDN
Martin R C, Martin R R, Putnam M L, 2018. First report of cocksfoot mottle virus infecting Dactylis glomerata in forage production fields in California. Plant Disease. 102 (10), 2050-2051. DOI:10.1094/PDIS-02-18-0277-PDN
Rabenstein F, Schmidt H B, 1979. Identification of cocksfoot mottle virus in the GDR. (Nachweis des Knaulgrasscheckungs-Virus (cocksfoot mottle virus) in der DDR.). Archiv fur Phytopathologie und Pflanzenschutz. 15 (5), 351-354. DOI:10.1080/03235407909440482
Ryabov E V, Krutov A A, Novikov V K, Zheleznikova O V, Morozov S Yu, Zavriev S K, 1996. Nucleotide sequence of RNA from the Sobemovirus found in infected cocksfoot shows a luteovirus-like arrangement of the putative replicase and protease genes. Phytopathology. 86 (4), 391-397. DOI:10.1094/Phyto-86-391
Principal SourceTop of page
Draft datasheet under review
ContributorsTop of page
01/04/20 Original text by:
Brian A. Hodge, The Ohio State University, Department of Plant Pathology. 1680 Madison Ave., Wooster, Ohio, USA.
Lucy R. Stewart, United States Department of Agriculture, Agricultural Research Service. Corn, Soybean and Wheat Quality Research Unit. 023 Selby Hall, 1680 Madison Ave., Wooster, Ohio, USA.
Distribution MapsTop of page
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