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Datasheet

Pratylenchus coffeae (banana root nematode)

Summary

  • Last modified
  • 20 March 2017
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Pratylenchus coffeae
  • Preferred Common Name
  • banana root nematode
  • Taxonomic Tree
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Nematoda
  •             Family: Pratylenchidae
  •                 Genus: Pratylenchus

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Pictures

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PictureTitleCaptionCopyright
Detail of female nematode isolated from yam tuber in Papua New Guinea.
TitleFemale nematode
CaptionDetail of female nematode isolated from yam tuber in Papua New Guinea.
CopyrightJohn Bridge/CABI BioScience
Detail of female nematode isolated from yam tuber in Papua New Guinea.
Female nematodeDetail of female nematode isolated from yam tuber in Papua New Guinea.John Bridge/CABI BioScience

Identity

Top of page

Preferred Scientific Name

  • Pratylenchus coffeae (Zimmermann 1898) Filipjev & Schuurmans Steckhoven 1941

Preferred Common Name

  • banana root nematode

Other Scientific Names

  • Anguillulina mahogani (Cobb, 1920) Goodey, 1932
  • Pratylenchus mahogani (Cobb, 1920) Filipjev, 1936
  • Pratylenchus musicola
  • Tylenchus coffeae Zimmermann, 1898
  • Tylenchus mahogani Cobb, 1920
  • Tylenchus musicola Cobb, 1919

International Common Names

  • English: nematode, Root lesion
  • Spanish: nematodo de la raiz del platano; nemátodo de la raíz del plátano; nemátodo de las lesiones; nematodo lesionador del cafeto (Mexico)
  • French: nematose des racines

Local Common Names

  • Japan: Negusare-sentyubyo

EPPO code

  • PRATCO (Pratylenchus coffeae)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Nematoda
  •             Family: Pratylenchidae
  •                 Genus: Pratylenchus
  •                     Species: Pratylenchus coffeae

Notes on Taxonomy and Nomenclature

Top of pagePratylenchus coffeae is a widespread and variable species which may represent a species complex. Recent molecular and morphological studies on isolates of P. coffeae and closely related species have demonstrated the difficulties involved in identification (Duncan et al., 1999). The complex has yet to be satisfactorily resolved.

Description

Top of pageMeasurements (after Sher and Allen, 1953): Females: L = 0.45-0.7 mm; a = 25-35; b = 5-7; c = 17-22; V = 54-2676-833.2-8; spear = 15-18 µm. Neotype female: L = 0.59 mm; a = 34; b = 6.3; c = 21; V = 3281.94; spear = 18 µm.

Males: L = 0.45-0.70 mm; a = 26-40; b = 6-7; c = 17-24; T = 45-52; spear = 15-17 µm. (After Loof, 1960): 69 females: L = 0.37-0.70 (0.53) mm; a = 17.7-30.5 (23.7); b = 5.0-7.8 (6.8); c = 13.7-23.9 (19.0); V = 63-2675.8-84.2 (80.1); spear = 14-17 µm.

10 males: L = 0.41-0.56 (0.48) mm; a = 23.8-31.4 (27.4); b = 5.9-7.7 (6.5); c = 17.6-23.3 (19.1); T = 37-58 (48); spear = 14-15 µm.


Description (after Siddiqi, 1972):

Female

Body rather slender in young and fatter in older specimens, distinctly annulated. Lateral fields normally with four, sometimes five or six incisures. Lip region slightly set off, with two distinct annules; occasional specimens have three annules on one side of the lip region.

Basal knobs of spear round to oblong. Post-uterine branch 1.0 to 1.5 times body-width long, but may be up to 90 µm long, with a terminal rudimentary ovary which sometimes has distinct oocytes. Spermathecae large, broadly oval to nearly rounded, often with sperms. Intra-uterine eggs may contain embryos.

Tail 2.0-2.5 times anal body width in young females, 1.5-2.0 times anal body width in old specimens; terminus indented, sometimes appearing smoothly rounded, truncate or irregularly crenate.

Male

Abundant. Spicules slender with well marked manubria and ventrally arcuate shaft, 16-20 µm long; gubernaculum 4-7 µm in length; hypoptygma prominent; bursal margins faintly crenate.


Distribution

Top of pageP. coffeae is very widespread in almost all tropical and subtropical countries. The distribution map includes records based on specimens of P. coffeae identified at CABI Bioscience (Egham, UK); dates of identification are noted in the Table (CABI Bioscience, various dates).

The record for Chile in CABI/EPPO (2000) and previous editions of this Compendium was based on incorrect information. CABI was notified in November 2005 that this species is not present in Chile (I Moreno, Servicio Agrícola y Ganadero (SAG), Chile, personal communication, 2005).

Distribution Table

Top 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.

CountryDistributionLast ReportedOriginFirst ReportedInvasiveReferencesNotes

ASIA

AfghanistanPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1985
BangladeshPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1985
BhutanPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1985
Brunei DarussalamWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
ChinaRestricted distributionCABI/EPPO, 2000; EPPO, 2014
-FujianPresentCABI/EPPO, 2000; EPPO, 2014; Liu et al., 1999
-GuangdongPresentCABI/EPPO, 2000; EPPO, 2014
-HunanPresentAnon., 1990; CABI/EPPO, 2000; EPPO, 2014
-JiangsuPresentCABI/EPPO, 2000; Teng et al., 2012; EPPO, 2014; Liu et al., 1999
-YunnanPresentLi et al., 2006
Georgia (Republic of)PresentEliava & Bagaturiya, 1971; CABI/EPPO, 2000; EPPO, 2014
IndiaRestricted distributionCABI/EPPO, 2000; EPPO, 2014
-Andhra PradeshPresentSundararaju, 2006
-BiharPresentCABI/EPPO, 2000; EPPO, 2014
-DelhiRestricted distributionSethi & Swarup, 1971; CABI/EPPO, 2000; EPPO, 2014
-Himachal PradeshWidespreadSethi & Swarup, 1971; Kaur et al., 1989; Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014
-Indian PunjabRestricted distributionSethi & Swarup, 1971; CABI/EPPO, 2000; EPPO, 2014
-KarnatakaPresentCABI/EPPO, 2000; EPPO, 2014
-KeralaPresentReni & Mohandas, 2004
-Madhya PradeshPresentTiwari et al., 2000
-ManipurWidespreadAnandi & Dhanachand, 1992; CABI/EPPO, 2000; EPPO, 2014
-OdishaPresentCABI/EPPO, 2000; EPPO, 2014
-RajasthanWidespreadSethi & Swarup, 1971; Mahajan & Kaur, 1991; CABI/EPPO, 2000; EPPO, 2014
-SikkimPresentSrivastava et al., 1998; CABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1995
-Tamil NaduPresentSundaram & Vadivelu, 1995; Sundaram, 1997; CABI/EPPO, 2000; EPPO, 2014
-TripuraPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1986
-Uttar PradeshPresentSiddiqi, 1964; CABI/EPPO, 2000; EPPO, 2014
-West BengalPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1981
IndonesiaWidespreadCABI/EPPO, 2000; EPPO, 2014
-JavaWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
-SumatraPresentCABI/EPPO, 2000; EPPO, 2014
IranPresentPourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014
JapanRestricted distributionGotoh, 1974; Inagaki, 1984; CABI/EPPO, 2000; EPPO, 2014
-KyushuPresentGotoh, 1972; Oashi, 1984; CABI/EPPO, 2000; EPPO, 2014
-Ryukyu ArchipelagoWidespreadFukudome, 1978; CABI/EPPO, 2000; EPPO, 2014
-ShikokuRestricted distributionZinno & Igarashi, 1972; CABI/EPPO, 2000; EPPO, 2014
Korea, DPRPresentKornobis & Dobosz, 1997; CABI/EPPO, 2000; EPPO, 2014
Korea, Republic ofPresentCABI/EPPO, 2000; EPPO, 2014; Kim & Minagawa, 1996
MalaysiaWidespreadMuhammad, 1992; CABI/EPPO, 2000; EPPO, 2014
OmanPresentMani et al., 1997; CABI/EPPO, 2000; EPPO, 2014
PakistanWidespreadKhan et al., 1989; Maqbool, 1992; CABI/EPPO, 2000; EPPO, 2014
PhilippinesWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
Sri LankaPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1987
TaiwanRestricted distributionHuang & Chiang, 1976; Tsay et al., 1995; CABI/EPPO, 2000; EPPO, 2014
ThailandWidespreadSiddiqi, 1972; Chunram, 1972; CABI/EPPO, 2000; EPPO, 2014
TurkeyPresentAkgül & Ökten, 2001
VietnamWidespreadRyss & Fam-Tkhan-Bin, 1989; Chau et al., 1997; CABI/EPPO, 2000; EPPO, 2014

AFRICA

CameroonRestricted distributionBridge et al., 1995; CABI/EPPO, 2000; EPPO, 2014
Congo Democratic RepublicWidespreadGeraert, 1962; CABI/EPPO, 2000; EPPO, 2014
Côte d'IvoireRestricted distributionFargette & Quénéhervé, 1988; CABI/EPPO, 2000; EPPO, 2014
GhanaPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1984
KenyaPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1992
MadagascarWidespreadWhitehead, 1968; CABI/EPPO, 2000; EPPO, 2014
MalawiWidespreadSaka & Siddiqi, 1979; Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014
MauritiusPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1992
MozambiquePresentOever & Mangane, 1992; CABI/EPPO, 2000; EPPO, 2014
NigeriaPresentKhan & Misari, 1992; CABI/EPPO, 2000; EPPO, 2014
SeychellesWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
South AfricaWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
SpainCABI/EPPO, 2000; EPPO, 2014
-Canary IslandsWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
SwazilandPresentDaneel et al., 2003
TanzaniaWidespreadCABI/EPPO, 2000; EPPO, 2014
UgandaPresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1983
ZambiaWidespreadKeetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014
ZimbabweWidespreadKeetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014

NORTH AMERICA

MexicoRestricted distributionKnobloch & Laughlin, 1973; CABI/EPPO, 2000; EPPO, 2014
USAWidespreadCABI/EPPO, 2000; EPPO, 2014
-ArkansasPresentRiggs et al., 1956; CABI/EPPO, 2000; EPPO, 2014
-CaliforniaPresentCABI/EPPO, 2000; EPPO, 2014
-FloridaPresentCABI/EPPO, 2000; EPPO, 2014
-HawaiiWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
-South CarolinaPresentCABI/EPPO, 2000; EPPO, 2014

CENTRAL AMERICA AND CARIBBEAN

BarbadosWidespreadSiddiqi, 1972; Braithwaite, 1977b; CABI/EPPO, 2000; EPPO, 2014
BelizePresentCABI/EPPO, 2000; EPPO, 2014; CABI Bioscience, 1993
Costa RicaWidespreadLopez & Salazar, 1990; CABI/EPPO, 2000; EPPO, 2014
CubaPresentStoyanov, 1967; CABI/EPPO, 2000; EPPO, 2014
DominicaPresentSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
Dominican RepublicWidespreadCABI/EPPO, 2000; EPPO, 2014
El SalvadorWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
GrenadaPresentCABI/EPPO, 2000; EPPO, 2014
GuadeloupePresentKermarrec et al., 1988; CABI/EPPO, 2000; EPPO, 2014
GuatemalaWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
HondurasWidespreadPinochet et al., 1978; CABI/EPPO, 2000; EPPO, 2014
JamaicaWidespreadThompson et al., 1973; CABI/EPPO, 2000; EPPO, 2014
MartiniqueWidespreadCadet et al., 1993; CABI/EPPO, 2000; EPPO, 2014
NicaraguaWidespreadEdwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014
PanamaWidespreadEdwards & Wehunt, 1973; Pinochet & Duarte, 1986; CABI/EPPO, 2000; EPPO, 2014
Puerto RicoPresentSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
Trinidad and TobagoWidespreadBraithwaite, 1980; CABI/EPPO, 2000; EPPO, 2014

SOUTH AMERICA

BrazilWidespreadCafe-Filho & Huang, 1988; CABI/EPPO, 2000; EPPO, 2014
-Minas GeraisPresentCastro et al., 2008
-PernambucoPresentMoura et al., 2005
-Rio de JaneiroPresentSilva & Inomoto, 2002
-Rio Grande do NortePresentMoura et al., 2004
-Sao PauloPresentMonteiro & Lordello, 1974; Rossi et al., 1995; CABI/EPPO, 2000; EPPO, 2014
ChileAbsent, invalid recordCABI/EPPO, 2000; EPPO, 2014
ColombiaWidespreadWehunt & Edwards, 1968; CABI/EPPO, 2000; EPPO, 2014
EcuadorWidespreadBridge, 1975; CABI/EPPO, 2000; EPPO, 2014
French GuianaPresentCABI/EPPO, 2000; EPPO, 2014
SurinamePresentCABI/EPPO, 2000; EPPO, 2014
VenezuelaWidespreadSiddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014

EUROPE

AustriaPresentFauna Europaea, 2014
BulgariaWidespreadKatalan-Gateva & Budurova, 1979; CABI/EPPO, 2000; EPPO, 2014
ItalyPresentCABI/EPPO, 2000; EPPO, 2014
SloveniaPresentFauna Europaea, 2014
SpainWidespreadCABI/EPPO, 2000; EPPO, 2014

OCEANIA

AustraliaWidespreadCSIRO, 1973; CABI/EPPO, 2000; EPPO, 2014
-New South WalesPresentHarding & Wicks, 2007
-QueenslandPresentBlair et al., 1999; Colbran, 1964; CABI/EPPO, 2000; EPPO, 2014
Cook IslandsPresentGrandison, 1990; CABI/EPPO, 2000; EPPO, 2014
FijiPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
KiribatiPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
NiuePresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
PalauPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
Papua New GuineaPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
SamoaPresentBridge, 1988; Grandison, 1996; CABI/EPPO, 2000; EPPO, 2014
Solomon IslandsPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
TongaPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014
VanuatuPresentBridge, 1988; CABI/EPPO, 2000; EPPO, 2014

Risk of Introduction

Top of pageP. coffeae is a phytosanitary risk in all tropical and subtropical countries.

Habitat

Top of pageThis nematode is a migratory root endoparasite found in soil and the roots of plants growing in subtropical and tropical uplands.

Hosts/Species Affected

Top of pageP. coffeae is a polyphagous species with a large host range of over 250 plant species covering almost all plant families. No attempt, therefore, has been made to list all host species; the listing has been limited to the most important crops which are amongst its hosts. Hosts of the nematode include Citrus spp., vegetables, ornamental foliar plants and weeds (Edwards and Wehunt, 1973; O'Bannon and Esser, 1975; Das and Das, 1986).

Host Plants/Plants Affected

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Plant nameFamilyContext
Aesculus chinensisHippocastanaceaeOther
Anacardium occidentale (cashew nut)AnacardiaceaeHabitat/association
AraceaeAraceaeMain
Areca catechu (betelnut palm)ArecaceaeHabitat/association
CitrusRutaceaeMain
Cocos nucifera (coconut)ArecaceaeHabitat/association
Coffea (coffee)RubiaceaeMain
Colocasia esculenta (taro)AraceaeMain
Cucurbitaceae (cucurbits)CucurbitaceaeMain
Curcuma longa (turmeric)ZingiberaceaeMain
Dioscorea (yam)DioscoreaceaeMain
Dioscorea batatas (Chinese yam)DioscoreaceaeOther
Elettaria cardamomum (cardamom)ZingiberaceaeMain
Euonymus alatus (winged spindle)CelastraceaeOther
Ficus carica (fig)MoraceaeMain
Musa (banana)MusaceaeMain
Musa textilis (manila hemp)MusaceaeMain
Musa x paradisiaca (plantain)MusaceaeMain
Piper nigrum (black pepper)PiperaceaeMain
Solanum tuberosum (potato)SolanaceaeMain
Theobroma cacao (cocoa)SterculiaceaeMain
Zea mays (maize)PoaceaeMain
Zingiber officinale (ginger)ZingiberaceaeMain

Growth Stages

Top of pageFlowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

Top of pageP. coffeae only infects the roots, tubers, corms and rhizomes of host plants but referred symptoms are produced on other parts of the plant.

Bananas, Plantains and Musa textilis

P. coffeae causes damage symptoms similar to those observed with other root lesion nematodes such as Radopholus similis: stunting; lengthening of the vegetative cycle; reduction in the size and number of leaves, and in bunch weight; reduction of the productive life of the plantation; toppling. Roots heavily infested by P. coffeae have extensive black or purple necrosis of epidermal and cortical tissue, often accompanied by secondary rotting and root breakage. Similar necrosis can be observed on the outer parts of the corm.

Yams

P. coffeae causes dry rot symptoms in yam tubers. Brown, irregular dry rot extends 1-2 cm into the outer tissues of D. rotundata tubers, but can occur as deep as 5 cm in tubers of D. alata. The dry rot can be more pronounced in the oldest apical portions of the tubers, adjacent to the vines or may even be restricted to these portions in newly harvested tubers.

External symptoms on tubers of D. alata, D. cayenensis and D. rotundata are deep cracks, a corky appearance, exposed dark-brown rotted areas and diseased tubers, which are spongy to the touch. Necrosis or rotting, caused by P. coffeae, has also been observed in tubers of D. esculenta and D. trifida. Above-ground symptoms of damage are not as obvious. Vines from tubers which are severely infected with P. coffeae are unthrifty and shorter than normal plants.

Using planting material with a high proportion of dry rot can result in tubers not sprouting and poor stands in yam fields (Acosta, 1974; Acosta and Ayala, 1975; Bridge and Page, 1984; Coates-Beckford and Brathwaite, 1977; Thompson et al., 1973).

Coffee

Coffee roots infected by P. coffeae turn yellow, then brown and most of the lateral roots are rotten. Infected plants appear stunted and have few small, chlorotic leaves. The earliest symptoms of infection in newly transplanted trees are yellowing of the leaves, loss of young primary branches and stunting of the shoot. A gradual wilt sets in, followed by death of the whole tree. Severely infected plants may die prematurely.

In the field, symptoms may occur in patches with yields reducing according to disease severity. Lesions occur on roots and, as a consequence, the whole root system is destroyed (Campos et al., 1990; Monteiro and Lordello, 1974; Whitehead, 1969).

Ginger and Turmeric

P. coffeae causes rhizome rot and leaf yellowing of ginger, and is associated with discoloration and rotting of the rhizomes of turmeric. At the advanced stages of infection, the rhizomes of turmeric become deep-red to brown in colour, less turgid, and wrinkled with rot symptoms. Internally, affected rhizomes have dark-brown, necrotic lesions (Sarma et al., 1974; Koshy and Bridge, 1990).

Symptoms List

Top of page
SignLife StagesType

Leaves

abnormal colours

Roots

cortex with lesions
necrotic streaks or lesions
reduced root system

Stems

stunting or rosetting

Vegetative organs

internal rotting or discoloration
surface cracking
surface lesions or discoloration

Whole plant

dwarfing
uprooted or toppled

Biology and Ecology

Top of pageP. coffeae is a migratory endoparasite of the root cortex; also of the corms of banana, plantain and Musa textilis; tubers of yam and caladium; and rhizomes of ginger, cardamom and turmeric, where it feeds and multiplies. It causes lesions and is often associated with primary and secondary rot in the root cortex, corms, tubers and rhizomes.

The developmental stages of P. coffeae infesting coffee in Java are as follows: the first moult takes place within the egg, three moults occur outside, and the eggs hatch in 6-8 days at 28-30°C.

In Japan, adult P. coffeae appeared about 2 weeks after hatching in potato tubers; the average lifespan was about 27 days at 25-30°C (Siddiqi, 1972).

P. coffeae has a life cycle of 21-28 days in yam roots and tubers. It can survive in moist soil for up to 8 months in the absence of host plants (Colbran, 1954).

The optimum temperature for reproduction in citrus roots is 29.5°C (Radewald et al., 1971). P. coffeae reproduces and multiplies in stored yams and is disseminated in seed tubers. It can also be introduced into yam fields in the roots and tissues of other crops. The nematodes can survive in field soil between yam crops on other hosts.

Notes on Natural Enemies

Top of pageNone known.

Plant Trade

Top of page
Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Barkadults; eggs; juvenilesYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Bulbs, Tubers, Corms, Rhizomesadults; eggs; juvenilesYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Growing medium accompanying plantsadults; eggs; juvenilesNoYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Rootsadults; eggs; juvenilesYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Seedlings, Micropropagated plantsadults; eggs; juvenilesYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Stems (above ground), Shoots, Trunks, Branchesadults; eggs; juvenilesYesPest or symptoms not visible to the naked eye but usually visible under light microscope
Plant parts not known to carry the pest in trade/transport
Flowers, Inflorescences, Cones, Calyx
Fruits (inc. pods)
Leaves
True seeds (inc. grain)
Wood

Impact

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Introduction

P. coffeae is a major pest of bananas, plantains and abaca, yams, coffee, ginger, citrus and other crops.

Bananas

P. coffeae is of particular importance on bananas in the Pacific island countries (Bridge, 1988) and parts of Africa (Sarah 1989), and on bananas, plantains and abaca (Musa textilis) in Cuba, Central and South America (Gowen and Quénéhervé, 1990). P. coffeae rarely occurs as the only nematode pest on the roots of Musa species. Although localized in South Africa, where it is found as the main nematode pest, the estimated crop losses can be as high as 80% (Sarah, 1989). In Cuba, high population densities cause toppling of the plants and limit production of the crop; economic threshold densities of the nematode have been estimated at 5000-10,000 nematodes per 100 g roots in good soils but as low as 1000-5000 nematodes per 100 g roots in soils with low fertility (Fernández and Ortega, 1998).

Yams (Dioscorea spp.)

P. coffeae has been recorded as a parasite of yams in Barbados, Jamaica and Puerto Rico, and in the Pacific islands of Papua New Guinea, Fiji, Niue, Tonga, Vanuatu and Solomon Islands. P. coffeae is the cause of tuber dry rot disease of yams, known locally in Jamaica as burn. (Ayala and Acosta, 1971; Brathwaite, 1977; Coates-Beckford and Brathwaite, 1977; Bridge, 1988).

P. coffeae is important as a parasite of the tubers reducing their edible portions, marketable value and, particularly, their storage qualities. In the regions of the world where the nematode occurs it can be very widespread. In Jamaica, 67 to 100% of D. rotundata and D. cayenensis tubers were found to be infected with P. coffeae (Thompson et al., 1973), and over 50% of D. alata tubers examined in Papua New Guinea had obvious signs of dry rot and were infested with P. coffeae sometimes in numbers in excess of 60,000 nematodes per 50 g tissues (Bridge and Page, 1984). Where there are obvious signs of dry rot in the tubers the likelihood of complete rot of tubers during storage is high. Dry rot of yams alone causes a marked reduction in the quality, marketable value and edible portions of tubers, and these reductions are more severe in stored yams.

Soil populations of 600 P. coffeae per vine of D. rotundata can produce significant tuber damage, and 1000 nematodes/plant can cause complete deterioration and severe reduction in tuber quality (Acosta and Ayala, 1975). However, neither of these populations causes reduction in total weight of harvested tubers. If seed tubers are badly affected by dry rot but survive storage, they can be so weakened that sprouting does not occur (Acosta and Ayala, 1976; Coates-Beckford and Brathwaite, 1977). Disinfesting tuber planting material with hot water treatment to control P. coffeae can increase tuber yields by 23% (Hutton et al., 1982).

Coffee

P. coffeae is found as pest of coffee in Dominican Republic, El Salvador, Guatemala, Puerto Rico, Costa Rica, Cuba and Brazil. P. coffeae also occurs on coffee in India, Southeast Asia, Barbados, Martinique, and Tanzania, Madagascar and in Indochina. In Java and India it is a very damaging and major pest of coffee (Campos et al., 1990). P. coffeae is the most destructive nematode of Coffea arabica in South India (Palanichamy, 1973). The use of nematicides to control P. coffeae can increase coffee yields by 28% (Figueroa, 1978).

Ginger, Turmeric

P. coffeae is widely distributed on ginger and turmeric in Kerala and Himachal Pradesh in India (Koshy and Bridge, 1990) where it is the cause of 'ginger yellows' and infects both roots and rhizomes (Kaur and Sharma, 1990). It has also been found in Sikkim associated with rotting of ginger rhizomes (J Bridge, CABI Bioscience, Egham, UK, personal communication).

Citrus

P. coffeae is not universally found on citrus but, where it occurs, field damage can be severe depending on the rootstock. Growth reduction in young trees can be 49-80% in the United States and fruit yields on rough lemon and sour orange rootstocks can be 143% and 231% higher, respectively, than trees infected with P. coffeae in the first bearing year, and 220% and 271% more in the second year (O'Bannon and Tomerlin, 1973).

Ramie (Boehmeria nivea)

P. coffeae has been shown to cause a serious root rot of this fibre crop in China resulting in significant yield decrease. Use of nematicides can give yield increases ranging from 11 to 60% (China, 1990).

Diagnosis

Top of pageNematodes are extracted from plant tissues and soils using standard nematological extraction techniques. In bananas and related crops, nematode damage is sometimes assessed by recording the incidence of uprooting per hectare per month. This may also be correlated with assessments of necrosis on primary roots and corms. These techniques can be used by those who are familiar with nematode symptoms but care should be taken not to confuse lesions caused by plant parasitic nematodes with those resulting from other root-infesting pests and pathogens.

In yams, the incidence and extent of dry rot disease, caused by P. coffeae, in tubers, can be assessed by direct observation. In tubers without obvious external symptoms of damage, the surface layers must be removed, or the tubers sectioned, to determine the presence of dry rot. Nematodes are found in yam soil and roots which can be sampled, particularly at the end of the growing season. However, most nematodes are found in tuber tissues and sampling these tissues is the most appropriate means of assessing populations.

Peelings of known thickness (1 or 2 cm) are cut from tubers. These are chopped finely, teased apart or, preferably, macerated before placing on a support tissue or sieve in water. Between 30 and 50% of nematodes emerge from tissues in the first 3 days but they continue migrating from the tissues for over 20 days.

Detection and Inspection

Top of pageP. coffeae can be detected by extraction from root, corm, tuber and rhizome tissues of plants showing damage symptoms. It can also be found in soil extracts.

Similarities to Other Species/Conditions

Top of pageP. coffeae is similar to several other species of Pratylenchus and may be confused with Radopholus similis by a non-expert. Recent approaches to the taxonomy of the genus involve molecular methodologies in an attempt to more accurately characterize the many described species (Duncan et al., 1999; Orui and Mizukubo, 1999; Uehara et al., 1999).

The nematodes, and the symptoms produced, are similar to those of other lesion nematodes of the genus Pratylenchus, and also those of the burrowing nematode, R. similis. Symptoms of dry rot disease in yam tubers are identical to those caused by the nematode Scutellonema bradys.

Prevention and Control

Top of pageCultural Control

In yams, the use of plant material which is free of nematodes and dry rot disease is an effective means of controlling or reducing the damage caused by P. coffeae. Central or distal tuber pieces, which generally contain lowest number of P. coffeae, are recommended as propagative material. Seed tubers showing symptoms of dry rot (cracking and flaking) should not be used for planting.

Any foliar material used for propagation should be completely free of P. coffeae. Yams, such as Dioscorea bulbifera and some forms of D. alata, can be readily propagated from bulbils or aerial tubers. A number of yams, such as D. alata, D. rotundata and D. dumetorum, can be produced from vine cuttings. Even true seed can be used for propagating D. rotundata.

Although these methods of propagation are not a practical means of producing ware tubers, they can be used to produce nematode-free seed tubers. Producing large numbers of seed tubers from relatively few yams by growing 'microsetts' or 'minisetts' cut from mature tubers (IITA, 1984) effectively produces nematode-free propagation material if clean, healthy 'mother seed yams' are selected (Jatala and Bridge, 1990).

In bananas, planting nematode disease-free suckers is recommended. Some growers or organizations maintain 'disease-free' nurseries from which new planting material, usually sword suckers, is collected. More commonly, planting material is taken from existing banana fields and these are more likely to be infested with nematodes. If the external tissue of the corm has purple or reddish-brown lesions, these, together with root stumps and adhering soil should be removed with a machete (pared) until only white corm tissue is exposed.

Paring suckers should take place away from the field, and corms with severe lesions should be discarded. Although useful, the paring technique may never be totally effective in removing all nematode infection; this treatment is often complemented by dipping suckers in a nematicidal solution or, more effectively, by coating them with a nematicidal mud.

In Côte d'Ivoire, it is recommended practice to store large corms in the sun for 2 weeks prior to planting; populations of Radopholus similis in the corm tissue can decline by as much as 80%. The most reliable means of avoiding the introduction of the nematode is to use completely nematode-free banana plantlets grown from meristem cultures (Quénéhervé and Cadet, 1985; Gowen and Quénéhervé, 1990).

Hot-Water Treatment

Bananas

The immersion of banana suckers in water held at a constant 55°C for periods of 15-25 minutes has been a commercial practice in Australia and Central and South America (Stover, 1972). However, the technique is difficult to manage because of the critical balance that is required to achieve a temperature that is lethal to nematodes in the corm tissue but does not cause permanent damage to the plant. This factor can also be important if suckers are not of uniform size.

Yams

In yam tubers, theoretical, but not always practical, control of P. coffeae can be achieved by hot water treatment. The immersion of tubers in hot water can reduce tuber populations of P. coffeae considerably, but rarely eliminates them without damage to the tuber.

Hot water at 46-52°C for 15-30 minutes is recommended for control of P. coffeae in D. rotundata tubers. The use of seed tubers with extreme dry rot should be avoided as treatment of these tubers is less effective. Treatments in water at 51°C for 15-45 minutes also effectively suppresses populations of P. coffeae and dry rot in D. rotundata tubers, and also increases vine growth. However, hot water treatment can cause severe physiological damage (Acosta and Ayala, 1976; Coates-Beckford and Brathwaite, 1977; Thompson et al., 1973).

Chemical Control

Nematicides are widely used by commercial banana growers producing fruit for the international export trade. Less specialized production, serving local markets, may not justify the high cost of chemical treatment. A number of organophosphate, oxime carbamate and carbamate nematicides are used on bananas either as granular or emulsifiable concentrate formulations.

Nematicides have been successfully used to control P. coffeae in coffee nurseries in Central America. Effective control of P. coffeae can be achieved with organocarbamate and organophosphate nematicides under field conditions.

Resistant Varieties

Increasing work is being done on assessing varietal resistance to P. coffeae. Price et al. (1996) reviewed techniques for field screening of Musa germplasm. Binks and Gowen (1996) also did field evaluation of Musa germplasm against P. coffeae and Radopholus similis. Banana cultivars were screened by Collingborn and Gowen (1997), the varieties Kunnan and Paka being considered highly resistant whereas Pisang Sipulu, Pisang Tongat and Pisang Mas were not considered to be resistant to the nematode. Collingborne et al. (1998) screened Indian cultivars of Musa for resistance or tolerance to P. coffeae and found that Yamgambi km5, Kunnan and Paka were most resistant. Stoffelen et al. (1999) investigated the host plant response of 13 Musa genotypes from Malaysia and Vietnam. Although differences in susceptibility between cultivars were observed, no resistance was detected. Stoffelen et al. (1999) also screened 32 Papua New Guinea banana varieties against P. coffeae, but none showed resistance to P. coffeae. Wiryadiputra (1996) tested 22 genotypes of Robusta coffee for resistance and found that BP961 was the most resistant with BP959, BP973, BP991 and BP993 also showing resistance. BP42, BP358 and BP409 were considered susceptible.

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Tsay TT, Cheng YH, Cheng HY, Lin YY, Wu WS, 1995. Occurrence and control of nematode diseases on bulbous flowers. Plant Pathology Bulletin, 4(4):180-192; 42 ref.

Uehara T, Kushida A, Momota Y, 1999. Rapid and sensitive identification of Pratylenchus spp. using reverse dot blot hybridization. Nematology, 1(5):549-555; 25 ref.

Wehunt EJ, Edwards DJ, 1968. Radopholus similis and other nematode species on banana. In: Smart GC, Perry VG, eds. Tropical Nematology. Gainesville, USA: University of Florida Press.

Whitehead AG, 1968. Nematodea. In: Le Pelley RH, ed. Pests of coffee. Longmans, Green and Co.

WHITEHEAD AG, 1969. Nematodes attacking coffee, tea and cocoa, and their control. Nematodes of tropical crops, 40:238-250.

Wiryadiputra S, 1996. Resistance of Robusta coffee to coffee root lesion nematode, Pratylenchus coffeae. Pelita Perkebunan, 12(3):137-148; 19 ref.

Zinno Y, Igarashi Y, 1972. Distribution of plant parasitic nematodes associated with coniferous seedlings in Shikoku, Japan. Bulletin of the Government Forestry Experimental Station, Tokyo, No. 246:11-20

Distribution Maps

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Distribution map Afghanistan: Present
CABI/EPPO, 2000; EPPO, 2014Austria: Present
Fauna Europaea, 2014Australia: Widespread
CSIRO, 1973; CABI/EPPO, 2000; EPPO, 2014Australia
See regional map for distribution within the countryAustralia
See regional map for distribution within the countryBarbados: Widespread
Siddiqi, 1972; Braithwaite ,1977b; CABI/EPPO, 2000; EPPO, 2014Barbados: Widespread
Siddiqi, 1972; Braithwaite ,1977b; CABI/EPPO, 2000; EPPO, 2014Bangladesh: Present
CABI/EPPO, 2000; EPPO, 2014Bulgaria: Widespread
Katalan-Gateva & Budurova, 1979; CABI/EPPO, 2000; EPPO, 2014Brunei Darussalam: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Brazil: Widespread
Cafe-Filho & Huang, 1988; CABI/EPPO, 2000; EPPO, 2014Brazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBhutan: Present
CABI/EPPO, 2000; EPPO, 2014Belize: Present
CABI/EPPO, 2000; EPPO, 2014Belize: Present
CABI/EPPO, 2000; EPPO, 2014Côte d'Ivoire: Restricted distribution
Fargette & Quénéhervé, 1988; CABI/EPPO, 2000; EPPO, 2014Cook Islands: Present
Grandison, 1990; CABI/EPPO, 2000; EPPO, 2014Cameroon: Restricted distribution
Bridge et al., 1995; CABI/EPPO, 2000; EPPO, 2014China: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014China: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014China
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryColombia: Widespread
Wehunt & Edwards, 1968; CABI/EPPO, 2000; EPPO, 2014Colombia: Widespread
Wehunt & Edwards, 1968; CABI/EPPO, 2000; EPPO, 2014Costa Rica: Widespread
Lopez & Salazar, 1990; CABI/EPPO, 2000; EPPO, 2014Costa Rica: Widespread
Lopez & Salazar, 1990; CABI/EPPO, 2000; EPPO, 2014Cuba: Present
Stoyanov, 1967; CABI/EPPO, 2000; EPPO, 2014Cuba: Present
Stoyanov, 1967; CABI/EPPO, 2000; EPPO, 2014Dominica: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Dominican Republic: Widespread
CABI/EPPO, 2000; EPPO, 2014Dominican Republic: Widespread
CABI/EPPO, 2000; EPPO, 2014Ecuador: Widespread
Bridge, 1975; CABI/EPPO, 2000; EPPO, 2014Spain: Widespread
CABI/EPPO, 2000; EPPO, 2014Spain: Widespread
CABI/EPPO, 2000; EPPO, 2014Spain
See regional map for distribution within the countryFiji: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Grenada: Present
CABI/EPPO, 2000; EPPO, 2014Georgia (Republic of): Present
Eliava & Bagaturiya, 1971; CABI/EPPO, 2000; EPPO, 2014Georgia (Republic of): Present
Eliava & Bagaturiya, 1971; CABI/EPPO, 2000; EPPO, 2014French Guiana: Present
CABI/EPPO, 2000; EPPO, 2014Ghana: Present
CABI/EPPO, 2000; EPPO, 2014Guadeloupe: Present
Kermarrec et al., 1988; CABI/EPPO, 2000; EPPO, 2014Guatemala: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Guatemala: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Honduras: Widespread
Pinochet et al., 1978; Pinochet et al., 1978; CABI/EPPO, 2000; EPPO, 2014Honduras: Widespread
Pinochet et al., 1978; Pinochet et al., 1978; CABI/EPPO, 2000; EPPO, 2014Indonesia: Widespread
CABI/EPPO, 2000; EPPO, 2014Indonesia: Widespread
CABI/EPPO, 2000; EPPO, 2014Indonesia
See regional map for distribution within the countryIndonesia
See regional map for distribution within the countryIndia: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014India
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Iran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Iran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Italy: Present
CABI/EPPO, 2000; EPPO, 2014Jamaica: Widespread
Thompson et al., 1973; CABI/EPPO, 2000; EPPO, 2014Jamaica: Widespread
Thompson et al., 1973; CABI/EPPO, 2000; EPPO, 2014Japan: Restricted distribution
Gotoh, 1974; Inagaki, 1984; CABI/EPPO, 2000; EPPO, 2014Japan
See regional map for distribution within the countryJapan
See regional map for distribution within the countryJapan
See regional map for distribution within the countryKenya: Present
CABI/EPPO, 2000; EPPO, 2014Kiribati: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Korea, DPR: Present
Kornobis & Dobosz, 1997; CABI/EPPO, 2000; EPPO, 2014Korea, Republic of: Present
CABI/EPPO, 2000; EPPO, 2014Sri Lanka: Present
CABI/EPPO, 2000; EPPO, 2014Madagascar: Widespread
Whitehead, 1968; CABI/EPPO, 2000; EPPO, 2014Martinique: Widespread
Cadet et al., 1993; CABI/EPPO, 2000; EPPO, 2014Mauritius: Present
CABI/EPPO, 2000; EPPO, 2014Malawi: Widespread
Saka & Siddiqi, 1979; Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014Mexico: Restricted distribution
Knobloch & Laughlin, 1973; CABI/EPPO, 2000; EPPO, 2014Mexico: Restricted distribution
Knobloch & Laughlin, 1973; CABI/EPPO, 2000; EPPO, 2014Malaysia: Widespread
Muhammad, 1992; CABI/EPPO, 2000; EPPO, 2014Mozambique: Present
Oever & Mangane, 1992; CABI/EPPO, 2000; EPPO, 2014Nigeria: Present
Khan & Misari, 1992; CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Niue: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Oman: Present
Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014Oman: Present
Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014Panama: Widespread
Edwards & Wehunt, 1973; Pinochet & Duarte, 1986; CABI/EPPO, 2000; EPPO, 2014Panama: Widespread
Edwards & Wehunt, 1973; Pinochet & Duarte, 1986; CABI/EPPO, 2000; EPPO, 2014Papua New Guinea: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Papua New Guinea: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Philippines: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Philippines: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Pakistan: Widespread
Khan et al., 1989; Maqbool, 1992; CABI/EPPO, 2000; EPPO, 2014Puerto Rico: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Puerto Rico: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Palau: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Palau: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Solomon Islands: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Seychelles: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Slovenia: Present
Fauna Europaea, 2014Suriname: Present
CABI/EPPO, 2000; EPPO, 2014Suriname: Present
CABI/EPPO, 2000; EPPO, 2014El Salvador: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014El Salvador: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Swaziland: Present
Daneel et al., 2003Thailand: Widespread
Siddiqi, 1972; Chunram, 1972; CABI/EPPO, 2000; EPPO, 2014Tonga: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Turkey: Present
Akgül & Ökten, 2001Turkey: Present
Akgül & Ökten, 2001Turkey: Present
Akgül & Ökten, 2001Trinidad and Tobago: Widespread
Braithwaite, 1980; CABI/EPPO, 2000; EPPO, 2014Trinidad and Tobago: Widespread
Braithwaite, 1980; CABI/EPPO, 2000; EPPO, 2014Taiwan: Restricted distribution
Huang & Chiang, 1976; Tsay et al., 1995; CABI/EPPO, 2000; EPPO, 2014Taiwan: Restricted distribution
Huang & Chiang, 1976; Tsay et al., 1995; CABI/EPPO, 2000; EPPO, 2014Tanzania: Widespread
CABI/EPPO, 2000; EPPO, 2014Uganda: Present
CABI/EPPO, 2000; EPPO, 2014USA: Widespread
CABI/EPPO, 2000; EPPO, 2014USA: Widespread
CABI/EPPO, 2000; EPPO, 2014USA
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: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Venezuela: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Vietnam: Widespread
Ryss & Fam-Tkhan-Bin, 1989; Chau et al., 1997; CABI/EPPO, 2000; EPPO, 2014Vanuatu: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Samoa: Present
Bridge, 1988; Grandison, 1996; CABI/EPPO, 2000; EPPO, 2014South Africa: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Zambia: Widespread
Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014Congo Democratic Republic: Widespread
Geraert, 1962; CABI/EPPO, 2000; EPPO, 2014Zimbabwe: Widespread
Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014
  • = 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) Afghanistan: Present
CABI/EPPO, 2000; EPPO, 2014Bangladesh: Present
CABI/EPPO, 2000; EPPO, 2014Brunei Darussalam: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Bhutan: Present
CABI/EPPO, 2000; EPPO, 2014China: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014Fujian: Present
CABI/EPPO, 2000; EPPO, 2014Guangdong: Present
CABI/EPPO, 2000; EPPO, 2014Hunan: Present
Anon., 1990; CABI/EPPO, 2000; EPPO, 2014Jiangsu: Present
CABI/EPPO, 2000; Teng et al., 2012; EPPO, 2014Yunnan: Present
Li et al., 2006Georgia (Republic of): Present
Eliava & Bagaturiya, 1971; CABI/EPPO, 2000; EPPO, 2014Indonesia: Widespread
CABI/EPPO, 2000; EPPO, 2014Java: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Sumatra: Present
CABI/EPPO, 2000; EPPO, 2014India: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014Andhra Pradesh: Present
Sundararaju, 2006Bihar: Present
CABI/EPPO, 2000; EPPO, 2014Delhi: Restricted distribution
Sethi & Swarup, 1971; CABI/EPPO, 2000; EPPO, 2014Himachal Pradesh: Widespread
Sethi & Swarup, 1971; Kaur et al., 1989; Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014Kerala: Present
Reni & Mohandas, 2004Karnataka: Present
CABI/EPPO, 2000; EPPO, 2014Manipur: Widespread
Anandi & Dhanachand, 1992; CABI/EPPO, 2000; EPPO, 2014Madhya Pradesh: Present
Tiwari et al., 2000Odisha: Present
CABI/EPPO, 2000; EPPO, 2014Indian Punjab: Restricted distribution
Sethi & Swarup, 1971; CABI/EPPO, 2000; EPPO, 2014Rajasthan: Widespread
Sethi & Swarup, 1971; Mahajan & Kaur, 1991; CABI/EPPO, 2000; EPPO, 2014Sikkim: Present
Srivastava et al., 1998; CABI/EPPO, 2000; EPPO, 2014Tamil Nadu: Present
Sundaram & Vadivelu, 1995; Sundaram, 1997; CABI/EPPO, 2000; EPPO, 2014Tripura: Present
CABI/EPPO, 2000; EPPO, 2014Uttar Pradesh: Present
Siddiqi, 1964; CABI/EPPO, 2000; EPPO, 2014West Bengal: Present
CABI/EPPO, 2000; EPPO, 2014Iran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Japan: Restricted distribution
Gotoh, 1974; Inagaki, 1984; CABI/EPPO, 2000; EPPO, 2014Kyushu: Present
Gotoh, 1972; Oashi, 1984; CABI/EPPO, 2000; EPPO, 2014Ryukyu Archipelago: Widespread
Fukudome, 1978; CABI/EPPO, 2000; EPPO, 2014Shikoku: Restricted distribution
Zinno & Igarashi, 1972; CABI/EPPO, 2000; EPPO, 2014Korea, DPR: Present
Kornobis & Dobosz, 1997; CABI/EPPO, 2000; EPPO, 2014Korea, Republic of: Present
CABI/EPPO, 2000; EPPO, 2014Sri Lanka: Present
CABI/EPPO, 2000; EPPO, 2014Malaysia: Widespread
Muhammad, 1992; CABI/EPPO, 2000; EPPO, 2014Oman: Present
Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014Papua New Guinea: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Philippines: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Pakistan: Widespread
Khan et al., 1989; Maqbool, 1992; CABI/EPPO, 2000; EPPO, 2014Palau: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Thailand: Widespread
Siddiqi, 1972; Chunram, 1972; CABI/EPPO, 2000; EPPO, 2014Turkey: Present
Akgül & Ökten, 2001Taiwan: Restricted distribution
Huang & Chiang, 1976; Tsay et al., 1995; CABI/EPPO, 2000; EPPO, 2014Vietnam: Widespread
Ryss & Fam-Tkhan-Bin, 1989; Chau et al., 1997; CABI/EPPO, 2000; EPPO, 2014
Distribution map (europe) Austria: Present
Fauna Europaea, 2014Bulgaria: Widespread
Katalan-Gateva & Budurova, 1979; CABI/EPPO, 2000; EPPO, 2014Spain: Widespread
CABI/EPPO, 2000; EPPO, 2014Georgia (Republic of): Present
Eliava & Bagaturiya, 1971; CABI/EPPO, 2000; EPPO, 2014Iran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Italy: Present
CABI/EPPO, 2000; EPPO, 2014Slovenia: Present
Fauna Europaea, 2014Turkey: Present
Akgül & Ökten, 2001
Distribution map (africa) Côte d'Ivoire: Restricted distribution
Fargette & Quénéhervé, 1988; CABI/EPPO, 2000; EPPO, 2014Cameroon: Restricted distribution
Bridge et al., 1995; CABI/EPPO, 2000; EPPO, 2014Spain: Widespread
CABI/EPPO, 2000; EPPO, 2014Canary Islands: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Ghana: Present
CABI/EPPO, 2000; EPPO, 2014Iran: Present
Pourjam et al., 1997; CABI/EPPO, 2000; EPPO, 2014Kenya: Present
CABI/EPPO, 2000; EPPO, 2014Madagascar: Widespread
Whitehead, 1968; CABI/EPPO, 2000; EPPO, 2014Mauritius: Present
CABI/EPPO, 2000; EPPO, 2014Malawi: Widespread
Saka & Siddiqi, 1979; Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014Mozambique: Present
Oever & Mangane, 1992; CABI/EPPO, 2000; EPPO, 2014Nigeria: Present
Khan & Misari, 1992; CABI/EPPO, 2000; EPPO, 2014Oman: Present
Mani et al., 1997; CABI/EPPO, 2000; EPPO, 2014Seychelles: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Swaziland: Present
Daneel et al., 2003Turkey: Present
Akgül & Ökten, 2001Tanzania: Widespread
CABI/EPPO, 2000; EPPO, 2014Uganda: Present
CABI/EPPO, 2000; EPPO, 2014South Africa: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Zambia: Widespread
Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014Congo Democratic Republic: Widespread
Geraert, 1962; CABI/EPPO, 2000; EPPO, 2014Zimbabwe: Widespread
Keetch & Buckley, 1984; CABI/EPPO, 2000; EPPO, 2014
Distribution map (north america) Belize: Present
CABI/EPPO, 2000; EPPO, 2014Cuba: Present
Stoyanov, 1967; CABI/EPPO, 2000; EPPO, 2014Dominican Republic: Widespread
CABI/EPPO, 2000; EPPO, 2014Guatemala: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Honduras: Widespread
Pinochet et al., 1978; Pinochet et al., 1978; CABI/EPPO, 2000; EPPO, 2014Jamaica: Widespread
Thompson et al., 1973; CABI/EPPO, 2000; EPPO, 2014Mexico: Restricted distribution
Knobloch & Laughlin, 1973; CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Puerto Rico: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014El Salvador: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014USA: Widespread
CABI/EPPO, 2000; EPPO, 2014Arkansas: Present
Riggs et al., 1956; CABI/EPPO, 2000; EPPO, 2014California: Present
CABI/EPPO, 2000; EPPO, 2014Florida: Present
CABI/EPPO, 2000; EPPO, 2014Hawaii: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014South Carolina: Present
CABI/EPPO, 2000; EPPO, 2014
Distribution map (central america) Barbados: Widespread
Siddiqi, 1972; Braithwaite ,1977b; CABI/EPPO, 2000; EPPO, 2014Belize: Present
CABI/EPPO, 2000; EPPO, 2014Colombia: Widespread
Wehunt & Edwards, 1968; CABI/EPPO, 2000; EPPO, 2014Costa Rica: Widespread
Lopez & Salazar, 1990; CABI/EPPO, 2000; EPPO, 2014Cuba: Present
Stoyanov, 1967; CABI/EPPO, 2000; EPPO, 2014Dominica: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Dominican Republic: Widespread
CABI/EPPO, 2000; EPPO, 2014Grenada: Present
CABI/EPPO, 2000; EPPO, 2014Guadeloupe: Present
Kermarrec et al., 1988; CABI/EPPO, 2000; EPPO, 2014Guatemala: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Honduras: Widespread
Pinochet et al., 1978; Pinochet et al., 1978; CABI/EPPO, 2000; EPPO, 2014Jamaica: Widespread
Thompson et al., 1973; CABI/EPPO, 2000; EPPO, 2014Martinique: Widespread
Cadet et al., 1993; CABI/EPPO, 2000; EPPO, 2014Mexico: Restricted distribution
Knobloch & Laughlin, 1973; CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Panama: Widespread
Edwards & Wehunt, 1973; Pinochet & Duarte, 1986; CABI/EPPO, 2000; EPPO, 2014Puerto Rico: Present
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Suriname: Present
CABI/EPPO, 2000; EPPO, 2014El Salvador: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Trinidad and Tobago: Widespread
Braithwaite, 1980; CABI/EPPO, 2000; EPPO, 2014USA: Widespread
CABI/EPPO, 2000; EPPO, 2014Florida: Present
CABI/EPPO, 2000; EPPO, 2014Venezuela: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
Distribution map (south america) Barbados: Widespread
Siddiqi, 1972; Braithwaite ,1977b; CABI/EPPO, 2000; EPPO, 2014Brazil: Widespread
Cafe-Filho & Huang, 1988; CABI/EPPO, 2000; EPPO, 2014Minas Gerais: Present
Castro et al., 2008Pernambuco: Present
Moura et al., 2005Rio de Janeiro: Present
Silva & Inomoto, 2002Rio Grande do Norte: Present
Moura et al., 2004Sao Paulo: Present
Monteiro & Lordello, 1974; Rossi et al., 1995; CABI/EPPO, 2000; EPPO, 2014Colombia: Widespread
Wehunt & Edwards, 1968; CABI/EPPO, 2000; EPPO, 2014Costa Rica: Widespread
Lopez & Salazar, 1990; CABI/EPPO, 2000; EPPO, 2014Ecuador: Widespread
Bridge, 1975; CABI/EPPO, 2000; EPPO, 2014French Guiana: Present
CABI/EPPO, 2000; EPPO, 2014Nicaragua: Widespread
Edwards & Wehunt, 1973; CABI/EPPO, 2000; EPPO, 2014Panama: Widespread
Edwards & Wehunt, 1973; Pinochet & Duarte, 1986; CABI/EPPO, 2000; EPPO, 2014Suriname: Present
CABI/EPPO, 2000; EPPO, 2014Trinidad and Tobago: Widespread
Braithwaite, 1980; CABI/EPPO, 2000; EPPO, 2014Venezuela: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014
Distribution map (pacific) Australia: Widespread
CSIRO, 1973; CABI/EPPO, 2000; EPPO, 2014New South Wales: Present
Harding & Wicks, 2007Queensland: Present
Blair et al., 1999; Colbran, 1964; CABI/EPPO, 2000; EPPO, 2014Cook Islands: Present
Grandison, 1990; CABI/EPPO, 2000; EPPO, 2014China: Restricted distribution
CABI/EPPO, 2000; EPPO, 2014Fiji: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Indonesia: Widespread
CABI/EPPO, 2000; EPPO, 2014Kiribati: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Niue: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Papua New Guinea: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Philippines: Widespread
Siddiqi, 1972; CABI/EPPO, 2000; EPPO, 2014Palau: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Solomon Islands: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Tonga: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Taiwan: Restricted distribution
Huang & Chiang, 1976; Tsay et al., 1995; CABI/EPPO, 2000; EPPO, 2014Vanuatu: Present
Bridge, 1988; CABI/EPPO, 2000; EPPO, 2014Samoa: Present
Bridge, 1988; Grandison, 1996; CABI/EPPO, 2000; EPPO, 2014