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Datasheet

Cercospora zeae-maydis (grey leaf spot)

Summary

  • Last modified
  • 15 May 2008
  • Datasheet Type(s)
  • Pest
  • Preferred Scientific Name
  • Cercospora zeae-maydis
  • Preferred Common Name
  • grey leaf spot
  • Taxonomic Tree
  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Dothideomycetes

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Pictures

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PictureTitleCaptionCopyright
The lower leaves of maize are usually the site of primary infection for C. zeae-maydis.
TitleSite of primary infection
CaptionThe lower leaves of maize are usually the site of primary infection for C. zeae-maydis.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
The lower leaves of maize are usually the site of primary infection for C. zeae-maydis.
Site of primary infectionThe lower leaves of maize are usually the site of primary infection for C. zeae-maydis.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Initial immature lesions of grey leaf spot are pin-point spots surrounded by a yellow halo. These become irregular or extended pale-brown streaks.
TitleLesions
CaptionInitial immature lesions of grey leaf spot are pin-point spots surrounded by a yellow halo. These become irregular or extended pale-brown streaks.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Initial immature lesions of grey leaf spot are pin-point spots surrounded by a yellow halo. These become irregular or extended pale-brown streaks.
LesionsInitial immature lesions of grey leaf spot are pin-point spots surrounded by a yellow halo. These become irregular or extended pale-brown streaks.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Moderately resistant hybrids display chlorotic lesions.
TitleChlorotic lesions
CaptionModerately resistant hybrids display chlorotic lesions.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Moderately resistant hybrids display chlorotic lesions.
Chlorotic lesionsModerately resistant hybrids display chlorotic lesions.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Further lesion expansion results in coalescing of lesions and blighting of the entire leaf.
TitleMature lesions
CaptionFurther lesion expansion results in coalescing of lesions and blighting of the entire leaf.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Further lesion expansion results in coalescing of lesions and blighting of the entire leaf.
Mature lesionsFurther lesion expansion results in coalescing of lesions and blighting of the entire leaf.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot lesions may be seen as dark silhouettes on senescent leaves.
TitleMature lesions
CaptionGrey leaf spot lesions may be seen as dark silhouettes on senescent leaves.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot lesions may be seen as dark silhouettes on senescent leaves.
Mature lesionsGrey leaf spot lesions may be seen as dark silhouettes on senescent leaves.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Symptoms may show on husks, but the lesions are not as clearly defined as on leaf tissue.
TitleSymptoms on husk
CaptionSymptoms may show on husks, but the lesions are not as clearly defined as on leaf tissue.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Symptoms may show on husks, but the lesions are not as clearly defined as on leaf tissue.
Symptoms on huskSymptoms may show on husks, but the lesions are not as clearly defined as on leaf tissue.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot of maize results in increased lodging; unsprayed maize in foreground, sprayed, healthy maize in background.
TitleLodging
CaptionGrey leaf spot of maize results in increased lodging; unsprayed maize in foreground, sprayed, healthy maize in background.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot of maize results in increased lodging; unsprayed maize in foreground, sprayed, healthy maize in background.
LodgingGrey leaf spot of maize results in increased lodging; unsprayed maize in foreground, sprayed, healthy maize in background.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Resistant hybrid (green) shown alongside susceptible hybrid (yellowed).
TitleResistant hybrid
CaptionResistant hybrid (green) shown alongside susceptible hybrid (yellowed).
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Resistant hybrid (green) shown alongside susceptible hybrid (yellowed).
Resistant hybridResistant hybrid (green) shown alongside susceptible hybrid (yellowed).Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Fungicidal treatments of C. zeae-maydis on maize crop. Fungicide sprayed (green crop) and unsprayed (yellowed crop).
TitleFungicidal treatments
CaptionFungicidal treatments of C. zeae-maydis on maize crop. Fungicide sprayed (green crop) and unsprayed (yellowed crop).
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Fungicidal treatments of C. zeae-maydis on maize crop. Fungicide sprayed (green crop) and unsprayed (yellowed crop).
Fungicidal treatmentsFungicidal treatments of C. zeae-maydis on maize crop. Fungicide sprayed (green crop) and unsprayed (yellowed crop).Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot results in smaller kernels. Ears 1 and 2 are from unsprayed, diseased maize, ears 3 and 4 are from healthy, fungicide-sprayed maize.
TitleComparison of maize kernels
CaptionGrey leaf spot results in smaller kernels. Ears 1 and 2 are from unsprayed, diseased maize, ears 3 and 4 are from healthy, fungicide-sprayed maize.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Grey leaf spot results in smaller kernels. Ears 1 and 2 are from unsprayed, diseased maize, ears 3 and 4 are from healthy, fungicide-sprayed maize.
Comparison of maize kernelsGrey leaf spot results in smaller kernels. Ears 1 and 2 are from unsprayed, diseased maize, ears 3 and 4 are from healthy, fungicide-sprayed maize.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Conidia of C. zeae-maydis.
TitleConidia
CaptionConidia of C. zeae-maydis.
CopyrightJulian Ward/KwaZulu-Natal Dept of Agriculture, South Africa
Conidia of C. zeae-maydis.
ConidiaConidia of C. zeae-maydis.Julian Ward/KwaZulu-Natal Dept of Agriculture, South Africa

Identity

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Preferred Scientific Name

  • Cercospora zeae-maydis Tehon & E.Y.Daniels

Preferred Common Name

  • grey leaf spot

Other Scientific Names

  • Mycosphaerella sp. (Latterell and Rossi, 1983)

International Common Names

  • English: gray leaf spot; gray leaf spot of corn; grey: maize leaf spot

EPPO code

  • CERCZM (Cercospora zeae-maydis)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Ascomycota
  •             Subphylum: Pezizomycotina
  •                 Class: Dothideomycetes
  •                     Subclass: Dothideomycetidae
  •                         Order: Capnodiales
  •                             Family: Mycosphaerellaceae
  •                                 Genus: Cercospora
  •                                     Species: Cercospora zeae-maydis

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.

CountryDistributionLast ReportedOriginFirst ReportedInvasiveReferencesNotes

ASIA

ChinaRestricted distributionCoates & White, 1994

AFRICA

CameroonPresentNowell, 1997
Congo Democratic RepublicRestricted distributionWard, 1996; Nowell, 1997
EthiopiaWidespreadWard et al., 1999
KenyaWidespreadNowell, 1997
MalawiWidespreadNowell, 1997
MozambiqueWidespreadWard, 1996; Nowell, 1997
NigeriaPresentNowell, 1997
South AfricaWidespreadWard, 1996; Nowell, 1997
SwazilandWidespreadWard, 1996; Nowell, 1997
TanzaniaPresentNowell, 1997
UgandaRestricted distributionNowell, 1997
ZambiaRestricted distributionWard, 1996; Nowell, 1997
ZimbabweWidespreadWard, 1996; Nowell, 1997

NORTH AMERICA

MexicoPresentBoothroyd, 1964
USA
-ColoradoPresentLipps et al., 1998
-IllinoisWidespreadLipps et al., 1998
-IowaWidespreadNutter et al., 1992
-KansasPresentLipps et al., 1998
-KentuckyWidespreadHyre, 1943
-MinnesotaPresentLipps et al., 1998
-North CarolinaRestricted distributionLeonard, 1974
-OhioWidespreadLipps et al., 1998
-South CarolinaRestricted distributionKingsland, 1963
-VirginiaWidespreadRoane, 1950
-WisconsinPresentLipps et al., 1998

CENTRAL AMERICA AND CARIBBEAN

Costa RicaPresentBoothroyd, 1964
Trinidad and TobagoPresentChupp, 1953

SOUTH AMERICA

BrazilPresentChupp, 1953
ColombiaPresentChupp, 1953
EcuadorPresentIMI Herbarium unda
PeruPresentChupp, 1953
VenezuelaPresentBoothroyd, 1964

Host plants/Plants Affected

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Plant nameFamilyContext
Zea mays (maize)PoaceaeUnknown

Growth Stages

Top of pageFlowering stage, Fruiting stage, Vegetative growing stage

List of Symptoms

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SignLife StagesType

Fruit

reduced size

Leaves

abnormal colours
fungal growth
necrotic areas
yellowed or dead

Seeds

shrivelled

Stems

lodging; broken stems

Whole plant

early senescence
plant dead; dieback

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Leaveshyphae; sporesYesYesPest or symptoms usually visible to the naked eye
Stems (above ground), Shoots, Trunks, Brancheshyphae; sporesYesYesPest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs, Tubers, Corms, Rhizomes
Fruits (inc. pods)
Growing medium accompanying plants
Roots
Seedlings, Micropropagated plants
True seeds (inc. grain)
Wood

References

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Allison JCS, Watson DJ, 1996. The production and distribution of dry matter in maize after flowering. Annals of Botany (London), 30:365-381.

Anderson BM, 1995. Gray leaf spot resistance - why do we need it? How do we get it? Proceedings of the 31st Annual Conference of AL Illinois Corn Breeders School, University of Illinois, 6-7 March, 1995, USA.

Anon., 1993. KwaZulu-Natal Dept. of Agric. Private Bag X9059, Pietermaritzburg, South Africa.

Anon., 1994. KwaZulu-Natal Dept. of Agric. Private Bag X9059, Pietermaritzburg, South Africa.

Anon., 1995, Fungal disease is threatening the US. Wall Street Journal, 21 August, Page C1.

Anon., 1995. KwaZulu-Natal Dept. of Agric. Private Bag X9059, Pietermaritzburg, South Africa.

Anon., 1996. Garst establishes a GLS standard. Garst Seeds. On-line publication, 980318.

Ayers JE, Johnson, MW, Hill RR, 1985. Identifying resistance to gray leaf spot. In: Proceedings of the Maize Sorghum Research Conference. 39th. Washington DC, USA: American Seed Trade Association, 157-175.

Bair W, Ayers JE, 1986. Variability in isolates of Cercospora zeae-maydis. Phytopathology, 76(2):129-132

Beckman PM, Payne GA, 1982. External growth, penetration, and development of Cercospora zeae-maydis in corn leaves. Phytopathology, 72(7):810-815

Beckman PM, Payne GA, 1983. Cultural techniques and conditions influencing growth and sporulation of Cercospora zeae-maydis and lesion development in corn. Phytopathology, 73(2):286-289

Boothroyd CW, 1964. Effect of elevation, rainfall and temperature upon the incidence of corn diseases in Costa Rica. (Abstr.) Phytopathology, 54:127.

Bubeck DM, Goodman MM, Beavis WD, Grant D, 1993. Quantitative trait loci controlling resistance to gray leaf spot in maize. Crop Science, 33(4):838-847

Cackett R, 2000. Sales data. Glenanil, South Africa: Farmers Agricare Pvt. Ltd.

Chupp C, 1953. A monograph of the fungus genus Cercospora. Ithaca, New York, USA: Cornell University Press.

Coates ST, White DG, 1994. Sources of resistance to gray leaf spot of corn. Plant Disease, 78(12):1153-1155

Crous PW, Braun U, 2003. Mycosphaerella and its anamorphs: 1. Names published in Cercospora and Passalora. CBS Biodiversity Series, 1:1-571.

Crous PW, Groenewald JZ, Groenewald M, Caldwell P, Braun U, Harrington TC, 2006. Species of Cercospora associated with grey leaf spot of maize. Studies in Mycology, 55:189-197.

de Nazareno NRX, Madden LV, Lipps PE, 1993. Characterization of gray leaf spot epidemics in maize. Journal of Plant Disease Protection, 100:410-425.

Dodd JL, 1980. The role of plant stresses in development of corn stalk rots. Plant Disease, 64(6):533-537

Eik K, Hanway JJ, 1966. Leaf area in relation to yield of corn grain. Agronomy Journal, 58:16-18.

Elwinger GF, Johnson MW, Hill RR Jr, Ayers JE, 1990. Inheritance of resistance to gray leaf spot of corn. Crop Science, 30(2):350-358

Freppon JT, Lipps PE, Pratt RC, 1994. Characterization of the chlorotic lesion response by maize to Cercospora zeae-maydis. Plant Disease, 78(10):945-949

Freppon JT, Pratt RC, Lipps PE, 1996. Chlorotic lesion response of maize to Cercospora zeae-maydis and its effect on gray leaf spot disease. Phytopathology, 86(7):733-738; 29 ref.

Gevers HO, Lake JK, 1994. GLS1 - a major gene for resistance to grey leaf spot in maize. South African Journal of Science, 90(7):377-379

Hilty JW, Hadden CH, Garden FT, 1979. Response of maize hybrids and inbred lines to gray leaf spot disease and the effects on yield in Tennessee. Plant Disease Reporter, 63(6):515-518

Huff CA, Ayers JE, Hill RR, 1988. Inheritance of resistance in corn (Zea mays) to gray leaf spot. Phytopathology, 78(6):790-794

Hyre RA, 1943. New records and unusual occurrence of plant diseases. Plant Disease Reporter, 27:553-554.

IMI Herbarium, 1900-. Herbarium specimen. International Mycological Institute (now CABI Bioscience) Herbarium. Egham, UK: CABI Bioscience.

Jenco JH, 1995. Epidemiology of Cercospora zeae-maydis on Zea mays in Iowa. MSc Thesis. Ames, Iowa, USA: Iowa State University.

Jenco JH, Nutter FW, Jr, 1992. Diurnal and seasonal periodicity of Cerospora zeae-maydis in Iowa. (Abstr.) Phytopathology, 82:992.

Juan Wang, Levy M, Dunkle LD, 1998. Sibling species of Cercospora associated with gray leaf spot of maize. Phytopathology, 88(12):1269-1275; 49 ref.

Kingsland GC, 1963. Cercospora blight of maize: A case history of a local epiphytotic in South Carolina. Plant Disease Reporter, 47:724-725.

Latterell FM, Rossi p, 1983. Gray leaf spot of corn: a disease on the move. Plant Disease, 67(8):842-847

Leonard KJ, 1974. Foliar pathogens of corn in North Carolina. Plant Disease Reporter, 58(6):532-534

Lipps PE, 1998. Gray Leaf Spot: A Global Threat to Corn Production. APSnet Feature. APSnet: Plant Pathology On-Line. American Phytopathological Society, St Paul, USA. World Wide Web page at http://www.scisoc.org/.

Lipps PE, Pratt RC, 1989. Maize hybrid evaluations for gray leaf spot. Plant Pathol. Ser. 81. Wooster, Ohio, USA: Department of Plant Pathology, Agricultural Research and Development Centre.

Lipps PE, Thomison PR, Pratt RC, 1996. Reaction of corn hybrids to gray leaf spot. In: 51st Annual Maize and Sorghum Research Conference, 163-180.

Lipps PE, White DG, Ayers JE, Dunkle LD, 1998. Gray Leaf Spot of Corn: Update. A Report from NCR-25 Technical Committee on Corn and Sorghum Diseases. American Phytopathological Society. On-line publication. May feature, full report.

Maroof MAS, Scoyoc SW van, Yu YG, Stromberg EL, 1993. Gray leaf spot disease of maize: rating methodology and inbred line evaluation. Plant Disease, 77(6):583-587

McGee DC, 1988. Maize diseases. A reference source for seed technologists. St. Paul, Minnesota, USA; APS Press, 149 pp.

Nazareno NRX de, Lipps PE, Madden LV, 1992. Survival of Cercospora zeae-maydis in corn residue in Ohio. Plant Disease, 76(6):560-563

Nazareno NRX de, Lipps PE, Madden LV, 1993. Effect of levels of corn residue on the epidemiology of gray leaf spot of corn in Ohio. Plant Disease, 77(1):67-70

Nowell DC, 1997. Studies on ear rot and gray leaf spot of maize in South Africa. PhD thesis. University of Natal, Pietermaritzburg, South Africa.

Nutter FW Jr, Teng PS, Royer MH, 1993. Terms and concepts for yield, crop loss, and disease thresholds. Plant Disease, 77(2):211-215

Nutter FW, Jr., Jenco JH, 1992. Development of critical-point yield loss models to estimate yield losses in corn caused by Cercospora zeae-maydis. (Abstr.) Phytopathology, 82:994.

Nutter FW, Jr., Martinson CA, Jenco I, Wegulo SN, 1994. Epidemiological studies concerning Cercospora zeae-maydis on maize. Proceedings of the 32nd Congress South African Society of Plant Pathology.

Payne GA, Waldron JK, 1983. Overwintering and spore release of Cercospora zeae-maydis in corn debris in North Carolina. Plant Disease, 67(1):87-89

Perkins JM, Smith DR, Kinsey JG, Dowden DC, 1995. Prevalence and control of gray leaf spot. In: Proceedings of the Annual Conference. Ill. Maize breeders School 31. Urbana, Illinois, USA: University of Illinois, 177-185.

Ringer CE, Grybauskas AP, 1995. Infection cycle components and disease progress of gray leaf spot on field corn. Plant Disease, 79(1):24-28

Roane CW, 1950. Observations on corn disease in Virginia from 1947 to 1950. Plant Disease Reporter, 34:394-396.

Roane CW, Harrison RL, Genter CF, 1974. Observations on gray leaf spot of maize in Virginia. Plant Disease Reporter, 58(5):456-459.

Rupe JC, Siegel MR, Hartman JR, 1982. Influence of environment and plant maturity on gray leaf spot of corn caused by Cercospora zeae-maydis. Phytopathology, 72(12):1587-1591

Saghai Maroof MA, Yue YG, Xiang ZX, Stromberg EL, Rufener GK, 1996. Identification of quantitative trait loci controlling resistance to gray leaf spot disease in maize. Theoretical and Applied Genetics, 93(4):539-546; 27 ref.

Shurtleff MC, ed., 1980. Compendium of corn diseases. Compendium of corn diseases. American Phytopathological Society St. Paul, Minn. USA, Edition 2:105 pp.

Shurtleff ML, Pederson M, 1991. Gray leaf spot, eyespot and yellow leaf blight of corn. Co-op Ext. Services Report on plant diseases. RPD No. 204. Illinois, USA: Department of Plant Pathology, University of Illinois Urbana-Champaign.

Smith KL, Grybauskas AP, 1985. Incidence of and tillage methods associated with gray leaf spot of field corn in Maryland. (Abstr.) Phytopathology, 75:1366.

Stromberg EL, 1986. Gray leaf spot disease of maize. Virginia Co-op. Extension Service Publication 450-072, Blacksburg, Virginia, USA: Virginia Polytechnic Institute and State University.

Stromberg EL, Donahue PJ, 1986. Hybrid performance of yield losses associated with gray leaf spot disease. In: Proceedings of the Maize Sorghum Research Conference, 41st American Seed Trade Association, Washington DC, USA, 92-104.

Stromberg EL, Flinchum LE, 1998. Evaluation of foliar fungicides for the control of gray leaf spot in Virginia, 1997. Fungic. Nematicide Test, 53:143-144.

Tehon LR, Daniels E, 1925. Notes on the parasitic fungi of Illinois. Mycologia, 17:240-249.

Thorson PR, Martinson CA, 1993. Development and survival of Cercospora zeae-maydis germlings in different relative humidity environments. Phytopathology, 83:153-157.

Ulrich JF, Hawk JA, Carroll RB, 1990. Diallel analysis of maize inbreds for resistance to gray leaf spot. Crop Science, 30(6):1198-1200

Ward JMJ, 1996. Epidemiology and management of gray leaf spot: A new disease of maize in South Africa. PhD thesis. University of Natal, Pietermaritzburg 3200, South Africa.

Ward JMJ, Hohls T, Laing MD, Rijkenberg FHJ, 1996. Fungicide responses of maize hybrids to grey leaf spot. European Journal of Plant Pathology, 102(8):765-771; 16 ref.

Ward JMJ, Laing MD, Cairns ALP, 1997. Management practices to reduce gray leaf spot of maize. Crop Science, 37(4):1257-1262; 19 ref.

Ward JMJ, Nowell DC, 1998. Integrated management for the control of maize gray leaf spot. Integrated Pest Management Review, 3:1-12.

Ward JMJ, Stromberg EL, Nowell DC, Nutter FWJr, 1999. Gray leaf spot: a disease of global importance in maize production. Plant Disease, 83(10):884-895; 80 ref.

White DG, Lipps PE, Ayers JE, 1996. Gray leaf spot of maize. Rep. NCR-25 Technical Comm. Maize Sorghum Diseases, St. Louis, MO, USA.

Distribution Maps

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Distribution map Brazil: Present
Chupp, 1953Cameroon: Present
Nowell, 1997China: Restricted distribution
Coates & White, 1994China: Restricted distribution
Coates & White, 1994Colombia: Present
Chupp, 1953Colombia: Present
Chupp, 1953Costa Rica: Present
Boothroyd, 1964Costa Rica: Present
Boothroyd, 1964Ecuador: Present
Ethiopia: Widespread
Ward et al., 1999Kenya: Widespread
Nowell, 1997Malawi: Widespread
Nowell, 1997Mexico: Present
Boothroyd, 1964Mexico: Present
Boothroyd, 1964Mozambique: Widespread
Ward, 1996; Nowell, 1997Nigeria: Present
Nowell, 1997Peru: Present
Chupp, 1953Swaziland: Widespread
Ward, 1996; Nowell, 1997Trinidad and Tobago: Present
Chupp, 1953Trinidad and Tobago: Present
Chupp, 1953Tanzania: Present
Nowell, 1997Uganda: Restricted distribution
Nowell, 1997USA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryVenezuela: Present
Boothroyd, 1964Venezuela: Present
Boothroyd, 1964South Africa: Widespread
Ward, 1996; Nowell, 1997Zambia: Restricted distribution
Ward, 1996; Nowell, 1997Congo Democratic Republic: Restricted distribution
Ward, 1996; Nowell, 1997Zimbabwe: Widespread
Ward, 1996; Nowell, 1997
  • = Present, no further details
  • = Evidence of pathogen
  • = Widespread
  • = Last reported
  • = Localised
  • = Presence unconfirmed
  • = Confined and subject to quarantine
  • = See regional map for distribution within the country
  • = Occasional or few reports
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Distribution map (asia) China: Restricted distribution
Coates & White, 1994
Distribution map (europe)
Distribution map (africa) Cameroon: Present
Nowell, 1997Ethiopia: Widespread
Ward et al., 1999Kenya: Widespread
Nowell, 1997Malawi: Widespread
Nowell, 1997Mozambique: Widespread
Ward, 1996; Nowell, 1997Nigeria: Present
Nowell, 1997Swaziland: Widespread
Ward, 1996; Nowell, 1997Tanzania: Present
Nowell, 1997Uganda: Restricted distribution
Nowell, 1997South Africa: Widespread
Ward, 1996; Nowell, 1997Zambia: Restricted distribution
Ward, 1996; Nowell, 1997Congo Democratic Republic: Restricted distribution
Ward, 1996; Nowell, 1997Zimbabwe: Widespread
Ward, 1996; Nowell, 1997
Distribution map (north america) Mexico: Present
Boothroyd, 1964Colorado: Present
Lipps et al., 1998Iowa: Widespread
Nutter et al., 1992Illinois: Widespread
Lipps et al., 1998Kansas: Present
Lipps et al., 1998Kentucky: Widespread
Hyre, 1943Minnesota: Present
Lipps et al., 1998North Carolina: Restricted distribution
Leonard, 1974Ohio: Widespread
Lipps et al., 1998South Carolina: Restricted distribution
Kingsland, 1963Virginia: Widespread
Roane, 1950Wisconsin: Present
Lipps et al., 1998
Distribution map (central america) Colombia: Present
Chupp, 1953Costa Rica: Present
Boothroyd, 1964Mexico: Present
Boothroyd, 1964Trinidad and Tobago: Present
Chupp, 1953Venezuela: Present
Boothroyd, 1964
Distribution map (south america) Brazil: Present
Chupp, 1953Colombia: Present
Chupp, 1953Costa Rica: Present
Boothroyd, 1964Ecuador: Present
Peru: Present
Chupp, 1953Trinidad and Tobago: Present
Chupp, 1953Venezuela: Present
Boothroyd, 1964
Distribution map (pacific) China: Restricted distribution
Coates & White, 1994