Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Meloidogyne hapla
(root knot nematode)



Meloidogyne hapla (root knot nematode)


  • Last modified
  • 28 March 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Meloidogyne hapla
  • Preferred Common Name
  • root knot nematode
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Nematoda
  •       Family: Meloidogynidae
  •         Genus: Meloidogyne

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Top of page
Hot spot" in groundnut field infested with M. hapla.
TitleField symptoms
CaptionHot spot" in groundnut field infested with M. hapla.
Copyright©R.A. Motsinger/Nemapix Vol. 1
Hot spot" in groundnut field infested with M. hapla.
Field symptomsHot spot" in groundnut field infested with M. hapla.©R.A. Motsinger/Nemapix Vol. 1


Top of page

Preferred Scientific Name

  • Meloidogyne hapla Chitwood, 1949

Preferred Common Name

  • root knot nematode

International Common Names

  • English: Northern root knot nematode
  • Spanish: nematodo norteno de quiste (Mexico)
  • French: nodosité des racines

Local Common Names

  • Germany: Älchen, Nördliches Wurzelgallen-
  • Japan: kita-nekobu-sentyubyo
  • Netherlands: wortelknobbelaaltje
  • Turkey: kok ur nematodu

EPPO code

  • MELGHA (Meloidogyne hapla)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Nematoda
  •             Family: Meloidogynidae
  •                 Genus: Meloidogyne
  •                     Species: Meloidogyne hapla

Notes on Taxonomy and Nomenclature

Top of page Meloidogyne hapla was first described from the USA by Chitwood (1949). The type host was Solanum tuberosum and the type locality was Long Island, New York, USA. No synonyms are known, although some of the records attributed to this species refer to other species such as Meloidogyne chitwoodi.


Top of page Measurements (after Whitehead, 1968).

Females (n=20): L = 419-845 (612) µm; width = 311-561 (430) µm; spear (9) = 10-13 (11) µm; width spear base (9) = 2-3 (2) µm; dorsal oesophageal gland orifice (8) = 4-6 (5) µm behind spear base; length median bulb (5) = 31-43 (36) µm; width median bulb (6) = 26-37 (31) µm; length median bulb valves (6) = 10-13 (12) µm; width median bulb valves (6) = 9-11 (10) µm.

Males (n=25): L = 791-1432 (1139) µm; a = 33.3-47.0 (41.7); length head (21) = 4.3-7.9 (5.6) µm; spear = 17.3-22.7 (20.0) µm; width spear base = 2.5-5.0 (3.5) µm; dorsal oesophageal gland orifice (8) = 2.5-3.2 (2.9) µm behind spear base; b' (total length divided by distance from anterior end to middle of median bulb) = 12.8-19.2 (15.5); c (24) = 73-283 (118); length median bulb (24) = 15.1-25.9 (19.2) µm; width median bulb (24) = 7.2-12.9 (9.3) µm; length median bulb valves (23) = 3.6-7.2 (5.9) µm; spicules (length of arc) (8) = 21.6-28.1 (25.7) µm; gubernaculum (5) = 7.2-9.4 (8.2) µm.

Infective juveniles (n=20) (J2): L = 312-355 (337) µm; a (18) = 20.1-26.6 (23.9); length tail (15) = 33-48 (43) µm; c (15) = 7.3-10.2 (7.9); c' (15) = 3.7-4.7 (4.4); length body to middle of genital primordium (13) = 177-214 (200) µm; spear (9) = 7.9-10.9 (9.7) µm.

Eggs (unembryonated) (n=20): 71-91 (78) µm 26-40 (31) µm.

Description (after Orton Williams, 1974)

Female: Body pyroid with short neck. Cuticle becoming thicker in posterior half of body, sometimes considerably. Head with two annules behind head-cap. Spear knobs rounded, inconspicuous. Excretory pore 14-20 annules behind head, hemizonid just posterior to pore. Posterior cuticular pattern roughly circular, composed of closely spaced smooth or slightly wavy striae. Dorsal arch low. Lateral fields may be unmarked, may be marked only by slight irregularities in the striae, or dorsal and ventral striae may meet at a slight angle along the fields. Some forking of striae at lateral fields may also occur. In some cases ventral striae may extend laterally on one or both sides to form 'wings' which the dorsal striae meet almost at right angles. Tail with few striae but distinct punctuations forming a stippled area between the anus and tail terminus. Sometimes the stippling may be more diffuse over the inner part of the pattern. Phasmids fairly widely spaced.

Male: Numerous in some populations, absent in others. Head not offset, a truncate cone to hemispherical in outline. Usually only one annule behind head-cap. Spear slender, spear knobs rounded and not offset. Anterior cephalid on second body annule, posterior cephalid just anterior to level of relaxed spear. Hemizonid 45-58 annules behind head, 0-4 annules anterior to excretory pore. Lateral field with four incisures. Tail terminus bluntly rounded; phasmids at about cloacal level. One or two testes. Spicules slightly curved, with small sharp processes projecting from the spicule wall at the junction of head and shaft into the spicule head. Gubernaculum crescentic, proximal end thicker than distal end.


Top of page M. hapla is extremely widely distributed, particularly in temperate regions and the cooler, higher altitude areas of the tropics. According to Whitehead (1969), M. hapla only flourishes at high altitudes above 6000 feet in East Africa (Kenya, Tanzania and Uganda), despite the abundance of host plants at lower altitudes. In Queensland, Australia, M. hapla was not found as far north as M. javanica (Colbran, 1958). Taylor and Buhrer (1958) reported that in the USA, M. hapla was the commonest root-knot nematode north of 39°N.

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes


ArmeniaPresentCABI/EPPO, 2002
ChinaPresentHu et al., 1997; CABI/EPPO, 2002
-ChongqingPresentCABI/EPPO, 2002
-FujianPresentCABI/EPPO, 2002
-HebeiPresentCABI/EPPO, 2002
-HenanPresentCABI/EPPO, 2002
-JiangsuPresentCABI/EPPO, 2002
-Nei MengguPresentCABI/EPPO, 2002
-ShandongPresentCABI/EPPO, 2002
-ShanxiPresentCABI/EPPO, 2002
-SichuanPresentCABI/EPPO, 2002
-YunnanPresentCABI/EPPO, 2002; Dong et al., 2015
IndiaPresentGoyal et al., 1976; CABI/EPPO, 2002
-Himachal PradeshPresentCABI/EPPO, 2002
-Jammu and KashmirPresentCABI/EPPO, 2002
-Tamil NaduPresentCABI/EPPO, 2002
-Uttar PradeshPresentCABI/EPPO, 2002
-West BengalPresentCABI/EPPO, 2002
IndonesiaPresentCABI/EPPO, 2002
-JavaPresentCABI/EPPO, 2002
IranPresentMaafi and Mahdavian, 1997; CABI/EPPO, 2002; Gharabadiyan et al., 2012
IsraelPresentMinz, 1956; CABI/EPPO, 2002
JapanPresentMitsui et al., 1976; CABI/EPPO, 2002
-HokkaidoPresentCABI/EPPO, 2002
-HonshuPresentCABI/EPPO, 2002
-KyushuPresentCABI/EPPO, 2002
-Ryukyu ArchipelagoPresentCABI/EPPO, 2002
-ShikokuPresentCABI/EPPO, 2002
KazakhstanPresentCABI/EPPO, 2002
Korea, Republic ofPresentChoi, 1981; CABI/EPPO, 2002
KyrgyzstanPresentCABI/EPPO, 2002
MalaysiaPresentMuhammad, 1992; CABI/EPPO, 2002
MongoliaPresentCABI/EPPO, 2002
PakistanPresentGul and Saeed, 1990; CABI/EPPO, 2002
PhilippinesPresentCABI/EPPO, 2002
Sri LankaPresentCABI/EPPO, 2002
TaiwanPresentRuelo, 1981; CABI/EPPO, 2002
TajikistanPresentCABI/EPPO, 2002
ThailandPresentRatanaprapa and Chunram, 1988; CABI/EPPO, 2002
TurkeyPresentCABI/EPPO, 2002
TurkmenistanPresentArutyunov, 1992; CABI/EPPO, 2002
UzbekistanPresentNarbaev, 1976; CABI/EPPO, 2002


AlgeriaPresent, few occurrencesCABI/EPPO, 2002
Côte d'IvoirePresentKouame et al., 1997; CABI/EPPO, 2002
EgyptPresentCABI/EPPO, 2002
EthiopiaPresentMeressa et al., 2014
KenyaPresentWhitehead, 1969; Parlevliet, 1971; CABI/EPPO, 2002; Karuri et al., 2017
LibyaPresentDabaj and Jenser, 1987; CABI/EPPO, 2002
MalawiPresentSaka, 1990; CABI/EPPO, 2002
MoroccoPresentCABI/EPPO, 2002
NigeriaPresentCABI/EPPO, 2002
South AfricaPresentVan, 1956; Kleynhans, 1991; CABI/EPPO, 2002
TanzaniaPresentWhitehead, 1969; Swai et al., 1996; CABI/EPPO, 2002
UgandaPresentWhitehead, 1969; CABI/EPPO, 2002
ZimbabweRestricted distributionMartin, 1961; Shepherd and Coombs, 1981; CABI/EPPO, 2002

North America

CanadaWidespreadPotter et al., 1972; CABI/EPPO, 2002
-New BrunswickRestricted distributionCABI/EPPO, 2002
-OntarioPresentCABI/EPPO, 2002
-Prince Edward IslandPresentCABI/EPPO, 2002
-QuebecPresentCABI/EPPO, 2002
MexicoPresentCABI/EPPO, 2002
USAWidespreadChitwood, 1949; CABI/EPPO, 2002
-CaliforniaPresentCABI/EPPO, 2002
-ConnecticutPresentLaMondia, 2002
-FloridaPresentCABI/EPPO, 2002
-HawaiiPresentHandoo et al., 2005
-IdahoPresentCABI/EPPO, 2002
-MainePresentCABI/EPPO, 2002
-MichiganPresentCABI/EPPO, 2002
-MinnesotaPresentCABI/EPPO, 2002
-NevadaPresentCABI/EPPO, 2002
-New JerseyPresentCABI/EPPO, 2002
-New YorkPresentCABI/EPPO, 2002
-North CarolinaPresentCABI/EPPO, 2002
-North DakotaPresentCABI/EPPO, 2002
-OhioPresentCABI/EPPO, 2002
-OklahomaPresentCABI/EPPO, 2002
-OregonPresentCABI/EPPO, 2002
-PennsylvaniaPresentCABI/EPPO, 2002
-Rhode IslandPresentMennan et al., 2006
-TennesseePresentCABI/EPPO, 2002
-TexasPresentWheeler et al., 2000
-UtahPresentCABI/EPPO, 2002
-VirginiaPresentCABI/EPPO, 2002
-WashingtonPresentCABI/EPPO, 2002
-WisconsinPresentMennan et al., 2006
-WyomingPresentCABI/EPPO, 2002

Central America and Caribbean

Costa RicaPresentLopez, 1991; CABI/EPPO, 2002
GuatemalaPresentHernandez et al., 2004
PanamaPresentCABI/EPPO, 2002

South America

ArgentinaPresentChaves and Torres, 1993; CABI/EPPO, 2002
BrazilPresentLordello and Monteiro, 1974; CABI/EPPO, 2002
-BahiaPresentCABI/EPPO, 2002
-CearaPresentCABI/EPPO, 2002
-GoiasRestricted distributionCABI/EPPO, 2002
-Minas GeraisPresentCABI/EPPO, 2002
-ParaPresentCABI/EPPO, 2002
-ParanaPresentCABI/EPPO, 2002
-PernambucoPresentCABI/EPPO, 2002
-Rio Grande do NortePresentCABI/EPPO, 2002
-Rio Grande do SulPresentCABI/EPPO, 2002
-Santa CatarinaRestricted distributionCABI/EPPO, 2002
-Sao PauloRestricted distributionCABI/EPPO, 2002
ChilePresentPhilippi et al., 1996; CABI/EPPO, 2002
ColombiaPresentCABI/EPPO, 2002
EcuadorWidespreadCABI/EPPO, 2002
ParaguayPresentCABI/EPPO, 2002
PeruPresentVargas and Pajuelo, 1973; CABI/EPPO, 2002
UruguayPresentRobertson et al., 2006
VenezuelaPresentCABI/EPPO, 2002


BelarusPresentGladkaya, 1983; CABI/EPPO, 2002
BelgiumPresentCoolen and Hendrickx, 1972; CABI/EPPO, 2002
BulgariaPresentStoyanov, 1980; CABI/EPPO, 2002
Czech RepublicPresentZouhar et al., 2003
EstoniaPresentKrall, 1970; CABI/EPPO, 2002
FinlandPresentTiilikkala, 1991; CABI/EPPO, 2002
Former USSRPresent
FrancePresentBerge et al., 1972; CABI/EPPO, 2002
GermanyPresentSturhan, 1976; CABI/EPPO, 2002
GreecePresentPyrowolakis, 1975; CABI/EPPO, 2002
HungaryPresentBudai, 1979; CABI/EPPO, 2002
ItalyPresentAmbrogioni, 1969; CABI/EPPO, 2002
LatviaPresentErenfelde, 1984; CABI/EPPO, 2002
LithuaniaPresentEfremenko and Klimakova, 1972; CABI/EPPO, 2002
MacedoniaPresentCABI/EPPO, 2002
MoldovaPresentCABI/EPPO, 2002
NetherlandsPresentBrinkman, 1975; CABI/EPPO, 2002
NorwayPresentStoeen, 1974; CABI/EPPO, 2002
PolandPresentBerbec, 1972; CABI/EPPO, 2002
PortugalPresentSantos et al., 1987; CABI/EPPO, 2002
RomaniaPresentRomascu et al., 1974; CABI/EPPO, 2002
Russian FederationWidespreadPokharel and Kruchina, 1991; CABI/EPPO, 2002
-Central RussiaPresentCABI/EPPO, 2002
-Southern RussiaPresentCABI/EPPO, 2002
-Western SiberiaPresentCABI/EPPO, 2002
SlovakiaPresentCABI/EPPO, 2002
SloveniaPresent?irca and Urek, 2005
SpainPresentPinochet et al., 1989; CABI/EPPO, 2002
SwitzerlandPresentVallotton, 1981; CABI/EPPO, 2002
UKPresentSouthey, 1974; CABI/EPPO, 2002
UkrainePresentZinovev & Volodchenko, 1984; CABI/EPPO, 2002
Yugoslavia (former)PresentGrujicic and Paunovic, 1971
Yugoslavia (Serbia and Montenegro)PresentCABI/EPPO, 2002


AustraliaWidespreadColbran, 1958; CABI/EPPO, 2002
-New South WalesPresentCABI/EPPO, 2002
-QueenslandPresentCABI/EPPO, 2002
-South AustraliaPresentCABI/EPPO, 2002
-TasmaniaPresentPethybridge et al., 2008
New ZealandPresentDale, 1973; CABI/EPPO, 2002
Norfolk IslandPresentBridge, 1988; CABI/EPPO, 2002
Papua New GuineaWidespreadBridge, 1988; CABI/EPPO, 2002

Risk of Introduction

Top of page M. hapla represents a severe risk to agricultural areas where it is not currently found, but as the species is virtually cosmopolitan the actual phytosanitary risk is probably low.


Top of page The infective second stage juveniles are found in the soil where they hatch from the eggs. The second stage penetrates a suitable root and all subsequent stages are located within the root tissue of the host where they remain as sedentary endoparasites with the exception of the adult vermiform male which may escape from the root into the soil.

Hosts/Species Affected

Top of page M. hapla is extremely polyphagous, attacking a wide variety of crops and weeds. Goodey et al. (1965) listed over 550 hosts and many more have been added since then. The species has recorded hosts in most of the higher plant families and attacks both herbaceous and woody plants. However, many grasses and cereals appear to be non-hosts. Carter (1985) provided a review of the recorded hosts.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Actinidia chinensis (Chinese gooseberry)ActinidiaceaeMain
Ageratina adenophora (Croftonweed)AsteraceaeOther
Allium cepa (onion)LiliaceaeOther
Anemone (windflower)RanunculaceaeOther
Arachis hypogaea (groundnut)FabaceaeMain
Beta vulgaris var. saccharifera (sugarbeet)ChenopodiaceaeMain
Brassica napus var. napus (rape)BrassicaceaeOther
Brassica oleracea var. capitata (cabbage)BrassicaceaeOther
Cajanus cajan (pigeon pea)FabaceaeOther
Camellia sinensis (tea)TheaceaeOther
Capsicum annuum (bell pepper)SolanaceaeOther
Chenopodium album (fat hen)ChenopodiaceaeWild host
Chenopodium quinoa (quinoa)ChenopodiaceaeOther
Chrysanthemum (daisy)AsteraceaeOther
Cichorium intybus (chicory)AsteraceaeMain
Coffea (coffee)RubiaceaeOther
Convolvulus arvensis (bindweed)ConvolvulaceaeWild host
Cucumis (melons, cucuimbers, gerkins)CucurbitaceaeOther
Daucus carota (carrot)ApiaceaeMain
Dianthus caryophyllus (carnation)CaryophyllaceaeOther
Dioscorea batatas (Chinese yam)DioscoreaceaeOther
Eustoma grandiflorum (Lisianthus (cut flower crop))GentianaceaeOther
Fabaceae (leguminous plants)FabaceaeOther
Fragaria ananassa (strawberry)RosaceaeMain
Glycine max (soyabean)FabaceaeMain
Lactuca sativa (lettuce)AsteraceaeOther
Linum usitatissimum (flax)Other
Medicago sativa (lucerne)FabaceaeMain
Mentha (mints)LamiaceaeOther
Nicotiana tabacum (tobacco)SolanaceaeOther
Olea europaea subsp. europaea (European olive)OleaceaeOther
Pelargonium (pelargoniums)GeraniaceaeMain
Phaseolus (beans)FabaceaeOther
Raphanus sativus (radish)BrassicaceaeOther
Rosa (roses)RosaceaeMain
Rubus (blackberry, raspberry)RosaceaeOther
Sinapis alba (white mustard)BrassicaceaeOther
Solanum (nightshade)SolanaceaeOther
Solanum lycopersicum (tomato)SolanaceaeMain
Solanum nigrum (black nightshade)SolanaceaeWild host
Solanum tuberosum (potato)SolanaceaeMain
Tanacetum cinerariifolium (Pyrethrum)Main
Trifolium (clovers)FabaceaeMain
Vicia (vetch)FabaceaeOther
Vitis vinifera (grapevine)VitaceaeOther

Growth Stages

Top of page Seedling stage, Vegetative growing stage


Top of page Typical symptoms of attack include a galling of the root system, the galls being relatively small and subspherical, often with a marked proliferation of small roots at the site of the gall (this is in contrast to the symptoms caused by other common species of Meloidogyne). In potato tubers, brown spots appearing in the tubers after the females commence egg production may identify infection sites. Severe attack by M. hapla results in impaired root function and concomitant stunting of the above ground parts leading to a reduction in yield.

List of Symptoms/Signs

Top of page
SignLife StagesType
Leaves / abnormal colours
Roots / galls along length
Roots / reduced root system
Whole plant / dwarfing
Whole plant / early senescence

Biology and Ecology

Top of page M. hapla is an obligate sedentary endoparasite of plant roots and tubers. The second stage infective juvenile penetrates the root and settles down within the cortex. As with all root-knot nematodes, a giant cell system of trophic cells is formed by the plant in response to secretions from the nematode. With each moult the nematode becomes more obese, although males become vermiform at the last moult and then emerge into the soil. The obese female swells enormously and produces numerous eggs (typically about 500) in a protective gelatinous matrix.

Unlike many root knot nematodes, M. hapla can withstand cold, eggs and juveniles surviving field temperatures below 0°C. However, it seems to be less tolerant of high temperatures than Meloidogyne incognita, for example. The optimum temperature for invasion and growth of M. hapla is in the range 20-25°C, a mean temperature of 27°C being inimical to development.

The nematode may be associated with other pathogens, including bacteria (such as Pseudomonas caryophylli) and fungi (such as Fusarium oxysporum, Rhizoctonia solani and Verticillium dahliae).

For further information see Orton Williams (1974).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Arthrobotrys oligospora Predator
Glomus fasciculatum Antagonist
Myrothecium verrucaria Pathogen
Pasteuria penetrans Pathogen

Notes on Natural Enemies

Top of page Pasteuria penetrans is known to parasitize root knot nematodes. The soil-dwelling second stage juveniles may be infected during the soil phase (Den Belder and Jansen, 1994). Arthrobotrys is also known to entrap the juveniles (Oostendorp et al., 1990; Watson et al., 1990).

Seedborne Aspects

Top of page M. hapla is not seedborne in the usual interpretation of the term, although it may be transmitted by infected planting material such as seed potatoes.

Pathway Vectors

Top of page
VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsEggs and juveniles in soil. Yes
Containers and packaging - woodEggs and juveniles in soil. Yes
Land vehiclesEggs and juveniles in soil. Yes
MailEggs and juveniles in soil. Yes
Soil, sand and gravelEggs and juveniles in soil. Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes adults; eggs; juveniles Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Growing medium accompanying plants adults; eggs; juveniles Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Roots adults; eggs; juveniles Yes Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Seedlings/Micropropagated plants adults; eggs; juveniles Yes Yes Pest 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
Fruits (inc. pods)
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)


Top of page M. hapla attacks nearly all temperate vegetables of economic importance and is well known as being capable of causing considerable reductions in yield, even to the point of total crop loss. In the field, crops including lucerne, groundnut, potato, carrot, sugarbeet, strawberry, pyrethrum and onion may be severely affected. For further quantitative information, see Luc et al. (1990) and Evans et al. (1993).

Detection and Inspection

Top of page M. hapla may be detected within the galled roots and tubers of the host by careful dissection and examination under the stereomicroscope. The infective juveniles may be recovered from soil using standard extraction methodologies as may the adult vermiform males when these occur.

Similarities to Other Species/Conditions

Top of page M. hapla is broadly similar to other members of the genus and normally requires the service of an experienced nematologist to identify to species level with any certainty. A suite of morphological characters, including the form of the perineal pattern of the mature female and the length and form of the second stage juvenile tail facilitate identification. Confusion with Meloidogyne chitwoodi is possible and PCR assays have been developed to assist in the rapid differentiation of the species. The galls formed by this species are usually rather atypical of root-knot nematodes in general in that they are often rather discrete and globular and arranged along the root. This symptom may be confused with galling by Nacobbus aberrans, the false root-knot nematode, and care should be taken to examine the nematodes inside the gall to confirm their identity.

Prevention and Control

Top of page

Various methods have been used to cleanse planting material, including hot water treatment (McDonald and Misari, 1976; Zunke, 1981) and a range of nematicidal drenches. Treatment in the field also relies upon the application of nematicides although frequent rotation with cereals or other graminaceous non-host crops may also be efficacious. Glasshouse soils may be fumigated to eradicate the pest. Many crops have potential for development of resistant or tolerant varieties.

For general information on the use of nematicides see Luc et al. (1990) and Evans et al. (1993). Both granular and liquid formulations have been used to control this nematode. More recent papers on this subject include LaMondia (1994), Johnston et al. (1995, 1996), Phipps et al. (1995) and Phipps and Eisenback (1996).


Top of page

?irca S; Urek G, 2005. Root-knot nematodes Meloidogyne spp. in Slovenia. (Ogorcice koreninskih ?i?k Meloidogyne spp. v Sloveniji.) In: Lectures and papers presented at the 7th Slovenian Conference on Plant Protection, Zrece, Slovenia, 8-10 March 2005. Ljubljana, Slovenia: Dru?tvo za varstvo rastlin Slovenije, 353-355.

Ambrogioni L, 1969. Two cases of mixed infections by nematodes of the genera Heterodera and Meloidogyne. Redia, 51:159-168

Arutyunov AV, 1992. The northern gall nematode Meloidogyne hapla Chitwood, 1949 - parasite of wild medicinal plants of Turkmenistan. Izvestiya Akademii Nauk Turkmenskoi SSR. Seriya Biologicheskikh Nauk, No. 2:24-29; 15 ref.

Belder E den; Jansen E, 1994. The influence of temperature, nutrition, light and the growth time of the mycelium on capture and infection of Meloidogyne hapla and Arthrobotrys oligospora. Fundamental and Applied Nematology, 17(1):57-66

Berbec E, 1972. Badania nad wystepowaniem i szkodliwoscia matwika polnocnego (Meloidogyne hapla Chitwood) na marchwi. Prace Wydzialu Nauk Przyrodniczych Bydgoskiego Towarzystwa Naukowego Ser. B, 15:3-32.

Berge JB; Dalmasso A; Ritter M, 1972. Studies on Meloidogyne hapla found in France. International Symposium of Nematology (11th), European Society of Nematologists, Reading, UK, 3-8 September, 1972. 2-3.

Bridge J, 1988. Plant-parasitic nematode problems in the Pacific Islands. Journal of Nematology, 20(2):173-183.

Brinkman H, 1975. Nematological observations in 1973 and 1974. Gewasbescherming, 6(4):57-64

Budai C, 1979. Spread of, and damage caused by the root knot nematode, Meloidogyne hapla Chitwood in the red pepper growing area of Szeged. Acta Phytopathologica Academiae Scientiarum Hungaricp, 14(3/4):543-548

CABI/EPPO, 2002. Meloidogyne hapla. Distribution Maps of Plant Diseases, No. 853. Wallingford, UK: CAB International.

Carter CC, 1985. Literature search: Host range of Meloidogyne hapla. International Nematology Network Newsletter, 2:16-24.

Chaves E; Torres M, 1993. Parasitic nematodes of potatoes in the south east of Buenos Aires. Boletin Tecnico, Estacion Experimental Agropecuaria, Balcarce, 115.

Chitwood BG, 1949. 'Root-knot nematodes'. Part 1. A revision of the genus Meloidogyne Goeldi, 1887. Proceedings of the Helminthological Society of Washington, 16:90-114.

Choi YE, 1981. The root-knot nematodes, Meloidogyne spp., in Korea. Proceedings of the 3rd Research Planning Conference on root-knot nematodes, Meloidogyne spp., Region VI, 20-24 July 1981, Jakarta, Indonesia. Raleigh, NC USA: North Carolina State University, 20-30

Colbran RC, 1958. Studies of plant and soil nematodes. 2. Queensland host records of root-knot nematodes (Meloidogyne species). Queensland Journal Agricultural Science, 15:101-136.

Coolen WA; Hendrickx GJ, 1972. Monograph on the nematological situation in Belgian rose culture. Publikatie nr. W 10, Rijksstation voor Nematologie en Entomologie, Merelbeke, 29 pp.

Dabaj KH; Jenser G, 1987. List of plants infected by root-knot nematodes in Libya. International Nematology Network Newsletter, 4(3):28-33

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