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Datasheet

Meloidogyne chitwoodi
(columbia root-knot nematode)

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Datasheet

Meloidogyne chitwoodi (columbia root-knot nematode)

Summary

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

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Pictures

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PictureTitleCaptionCopyright
Scanning electron micrograph of the anterior end of a male of M. chitwoodi.
TitleMale
CaptionScanning electron micrograph of the anterior end of a male of M. chitwoodi.
Copyright©J.D. Eisenback/Nemapix Vol. 1
Scanning electron micrograph of the anterior end of a male of M. chitwoodi.
MaleScanning electron micrograph of the anterior end of a male of M. chitwoodi.©J.D. Eisenback/Nemapix Vol. 1

Identity

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Preferred Scientific Name

  • Meloidogyne chitwoodi Golden, O'Bannon, Santo & Finley, 1980

Preferred Common Name

  • columbia root-knot nematode

International Common Names

  • French: nématode cécidogène de columbia

EPPO code

  • MELGCH (Meloidogyne chitwoodi)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Nematoda
  •             Family: Meloidogynidae
  •                 Genus: Meloidogyne
  •                     Species: Meloidogyne chitwoodi

Notes on Taxonomy and Nomenclature

Top of page References to Meloidogyne chitwoodi B-type in van Meggelen et al. (1994) refer to M. fallax (Karssen, 1996).

Description

Top of page Adult males and the second-stage juveniles are vermiform, motile animals, similar in general appearance to free-living soil nematodes. Females are characteristically pear-shaped, pearly-white and sedentary. The male is 887-1268 µm in length and 22-37 µm in width with a slight taper at each end. The tail is short, 4.7-9.0 µm and rounded. Cuticular annules are distinct and are more prominent near each end. The female is 430-740 µm in length and 344-518 µm in width. Second-stage juveniles are 336-417 µm in length and 12.5-15.5 µm in width, tail short, 39-47 µm, scarcely tapered and hyaline. Eggs are 79-92 µm in length and 40-46 µm in width.

Distribution

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M. chitwoodi was first described from the Pacific Northwest of the USA in 1980, its common name deriving from the Columbia River between Oregon and Washington states. It is not clear whether this is its area of origin. It was first detected in the EPPO region in the 1980s, in the Netherlands, but a review of old illustrations and old specimens of Meloidogyne suggests that it may have occurred earlier (in the 1930s) and may have been present throughout the intervening period (OEPP/EPPO, 1991). It is possible that M. chitwoodi has a wider distribution, undetected, in Europe than is currently known; the question is now actively under investigation.

A record for Wyoming, USA, was incorrectly included in CABI/EPPO (2000). The record for Virginia, USA, has been changed to 'Absent, confirmed by survey' following surveys conducted by Virginia Department of Agriculture and Consumer Services (2007-2010) and the Department of Plant Pathology, Physiology and Weed Science, Virginia Tech.

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

TurkeyRestricted distributionCABI/EPPO, 2012; EPPO, 2014

Africa

MozambiquePresentEPPO, 2014
South AfricaRestricted distributionFourie et al., 1998; CABI/EPPO, 2012; EPPO, 2014
TunisiaAbsent, unreliable recordEPPO, 2014

North America

CanadaAbsent, confirmed by surveyEPPO, 2014
-AlbertaAbsent, confirmed by surveyEPPO, 2014
-British ColumbiaAbsent, confirmed by surveyEPPO, 2014
MexicoPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
USARestricted distributionCABI/EPPO, 2012; EPPO, 2014
-CaliforniaPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-ColoradoPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-IdahoPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-NevadaPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-New MexicoPresentThomas et al., 2001; CABI/EPPO, 2012; EPPO, 2014
-OregonPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-TexasPresentSzalanski et al., 2001; CABI/EPPO, 2012; EPPO, 2014
-UtahPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-VirginiaAbsent, confirmed by surveyVirginia Department of Agriculture & Consumer Services surveys 2007-2010; Eisenback et al., 1986; CABI/EPPO, 2012; EPPO, 2014
-WashingtonPresent, 1994; CABI/EPPO, 2012; EPPO, 2014
-WyomingAbsent, invalid recordCABI/EPPO, 2012; EPPO, 2014

South America

ArgentinaPresentTiilikkala et al., 1995; CABI/EPPO, 2012; EPPO, 2014

Europe

BelgiumPresentCABI/EPPO, 2012; EPPO, 2014
BulgariaAbsent, confirmed by surveyEPPO, 2014
DenmarkAbsent, no pest recordIPPC, 2013; EPPO, 2014
FranceRestricted distributionEPPO, 2011; CABI/EPPO, 2012; EPPO, 2014
-France (mainland)Present, few occurrencesCABI/EPPO, 2012
GermanyTransient: actionable, under eradicationHeinicke, 1993; Muller et al., 1996; CABI/EPPO, 2012; EPPO, 2014
ItalyPresent, few occurrencesCABI/EPPO, 2012
NetherlandsRestricted distributionBrinkman & van Riel, 1990; NPPO of the Netherlands, 2013; CABI/EPPO, 2012; EPPO, 2014
PortugalPresent, few occurrencesCABI/EPPO, 2012; EPPO, 2014
-MadeiraPresent, few occurrencesCABI/EPPO, 2012
SwedenRestricted distributionEPPO, 2018
SwitzerlandEradicatedEPPO, 2011; CABI/EPPO, 2012; EPPO, 2014
UKAbsent, confirmed by surveyEPPO, 2014
-England and WalesAbsent, confirmed by surveyEPPO, 2014

Risk of Introduction

Top of page M. chitwoodi has been added to the EC Plant Health Directive 77/93/EEC.

A comparison of the air temperatures found in Finland with those found across the present distribution of the pest indicated that the nematode could survive and produce two generations per year in the southern part of Finland. Baker and Dickens (1993) concluded from their PRA that M. chitwoodi would be likely to produce three generations in the UK. They did not feel capable of predicting the likely economic impact of the pest, as this could depend on a number of other unknown factors such as soil wetness, varietal susceptibility and quality control thresholds. A recent PRA for Germany (Braasch et al., 1996) led to the conclusion that M. chitwoodi was most likely to be damaging on potatoes in the north-west of the country, in the area adjoining the Netherlands. Countries further south in the EPPO region would provide climatic conditions suitable for as many as four generations per year.

Potato crops would be most at risk from M. chitwoodi in the EPPO region. For a number of reasons, it represents a greater threat than other Meloidogyne species already widespread in the EPPO region, in particular M. hapla with which it often forms mixed populations. M. chitwoodi is less easily controlled by nematicides, it has a wider host range, it produces more severe tuber symptoms and is tolerant of lower soil temperatures. In fact, the soil temperature requirements for population development of M. chitwoodi are similar to those of Globodera rostochiensis (Tiilikkala et al., 1995), suggesting that the former species could occupy the same geographical distribution as the latter.

Hosts/Species Affected

Top of page M. chitwoodi has a wide host range among several plant families (Santo et al., 1980; O'Bannon et al., 1982), including crop plants and common weed species. Potatoes and tomatoes are good hosts, while barley, maize, oats, sugarbeet, wheat (Triticum aestivum) and various Poaceae (grasses and weeds) will maintain the nematode.

Moderate to poor hosts occur in the Brassicaceae, Cucurbitaceae, Fabaceae, Lamiaceae, Liliaceae, Umbelliferae and Vitaceae.

Fourie et al. (1998), working with a South African population of M. chitwoodi, reported Lycopersicon esculentum, Brassica rapa, Eragrostis tef and Lolium multiflorum as supporting high populations, whereas Eragrostis curvula, Arachis hypogaea and Zea mays were poor hosts.

Korthals et al. (2000) assessed host plant suitability at two sites in the Netherlands. Population growth of M. chitwoodi was greatest in potato, rye and summer wheat while nematodes in sugarbeet, hemp, perennial ryegrass and Tagetes patula remained equivalent to those of fallow. Because of poor reproduction, they recommended perennial ryegrass as a cover crop after cereals, maize, carrots or potatoes.

Lucerne is a good host for race 2 but not for race 1 (see Biology and Ecology for information on races), whereas carrots are a non-host for race 2 but a good host for race 1. Ferris et al. (1994a,b), investigating suitable crops for rotation with potato in the presence of race 1 in the USA, recommend Amaranthus, lucerne, rape (Brassica napus var. oleifera), Raphanus sativus var. oleifera and safflower (Carthamus tinctorius). In the Netherlands, host crops recorded to be attacked by M. chitwoodi are carrots, cereals, maize, peas, Phaseolus vulgaris, potatoes, Scorzonera hispanica, sugarbeet and tomatoes (OEPP/EPPO, 1991).

Recent studies have expanded information on the host range of M. chitwoodi (e.g. Brinkman et al., 1996; Griffin and Rumbaugh, 1996). Nijs et al. (2004) presented the results of experiments performed in the Netherlands and provided information on the host status of many plants together with an assessment of their potential phytosanitary risk.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Avena sativa (oats)PoaceaeHabitat/association
Beta vulgaris var. saccharifera (sugarbeet)ChenopodiaceaeHabitat/association
Chenopodium quinoa (quinoa)ChenopodiaceaeOther
Daucus carota (carrot)ApiaceaeMain
Hordeum vulgare (barley)PoaceaeHabitat/association
Medicago sativa (lucerne)FabaceaeMain
Phaseolus vulgaris (common bean)FabaceaeOther
Pisum sativum (pea)FabaceaeOther
Poaceae (grasses)PoaceaeHabitat/association
Scorzonera hispanica (oyster plant)AsteraceaeOther
Solanum lycopersicum (tomato)SolanaceaeMain
Solanum tuberosum (potato)SolanaceaeMain
Triticum aestivum (wheat)PoaceaeHabitat/association
Zea mays (maize)PoaceaeOther

Growth Stages

Top of page Pre-emergence, Seedling stage, Vegetative growing stage

Symptoms

Top of page Symptoms of M. chitwoodi vary according to host, population density of the nematode and environmental conditions. Above-ground symptoms are often not obvious but may consist of varying degrees of stunting, lack of vigour and a tendency to wilt under moisture stress, all leading to reduced yield. The galls produced on potato tubers by M. chitwoodi differ from those caused by other species of Meloidogyne. M. hapla, for example, forms small but distinct galls (together with extensive root proliferation) while M. incognita forms large, easily noticeable galls. The symptoms caused by M. chitwoodi are often not easily detected and are more apparent in some cultivars than in others; tubers may, in some cases, be heavily infected without visible symptoms. When present, the galls appear as small raised swellings on the tuber surface above the developing nematodes. A number of galls may be concentrated on one area of the tuber or single galls may be scattered near eyes or lesions. Internal tissue below the gall is necrotic and brownish. Adult females are visible just below the surface as glistening, white, pear-shaped bodies surrounded by a brownish layer of host tissue. Potato roots may also be infected, but this is difficult to detect without a magnifying lens, as little or no galling occurs, even in heavy infestations. The spherical bodies of females may protrude from the surface of small rootlets surrounded posteriorly by a large egg-filled sac which becomes dark-brown with age.

In other crops, root galls and reduced root production decrease yields and marketability. Gall formation occurs on most cereals but is more noticeable on wheat and oats than on barley or maize. In tomatoes, M. chitwoodi produces root galls in some cultivars but not in others.

List of Symptoms/Signs

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SignLife StagesType
Leaves / wilting
Leaves / yellowed or dead
Roots / galls along length
Roots / hairy root
Roots / swollen roots
Whole plant / dwarfing

Biology and Ecology

Top of page The life cycle of M. chitwoodi takes approximately 3-4 weeks under favourable conditions. Juveniles hatch from eggs in soil or on the root surface. Second-stage juveniles (the infective stage) penetrate root tips through unsuberized epidermal cells or wounds and move into the cortical region. Soon after entry, nematodes stimulate giant cell and gall formation in the host tissue. Necrotic lesions occur in the cortex. Juveniles then swell to become sausage-shaped, stop feeding and undergo three rapid successive moults to become adult males or females. Adult males have slender, vermiform bodies; they leave the root and are found free in the rhizosphere or near the protruding body of the female. However, as in the case of other Meloidogyne spp., it is probable that males are largely functionless and reproduction is nearly always parthenogenetic. Adult females have characteristically pear-shaped, pearly-white bodies and they are found embedded in host tissue. Eggs are laid by the female in a gelatinous sac near the root surface. In potato tubers, modified host cells form a protective layer or 'basket' around the egg mass and the juveniles as they hatch. The egg shell itself is hyaline with no visible markings.

M. chitwoodi passes the winter as eggs or juveniles and can survive extended periods of sub-zero temperatures. M. chitwoodi needs a temperature of 4°C for hatching and penetrating roots, and 6°C for development. Charchar and Santo (2001) reported on the effect of temperature on embryonic development and hatching of M. chitwoodi in Brazil. M. chitwoodi requires 600-800 degree-days to complete the first generation, whilst subsequent generations require 500-600 degree-days. Studies on potato in Oregon, USA, during 1992-1994 indicated that rapid population increase occurred after 1200 degree-days (Ingham and Rykbost, 1995). M. hapla requires a similar number of degree-days for development but does not begin development until temperatures rise above 10°C. Population dynamics in relation to degree-day accumulation have been considered by Pinkerton et al. (1991). M. chitwoodi was found to be more pathogenic at 25°C than at 30°C in glasshouse tests performed on yellow sweet clover (Griffin and Jensen, 1997). Brommer and Molendijk (2001) reported that there were at least two generations per year in fields naturally infected with M. chitwoodi in the Netherlands.

Several races of M. chitwoodi have been described, distinguished by slight differences in host range. The first two, known as race 1 and race 2, were in particular distinguished with regard to carrot and lucerne (Santo and Pinkerton, 1985; Mojtahedi et al., 1988). Race 3 has recently been described in California, USA (Mojtahedi et al., 1994). The nematode found in the Netherlands previously named M. chitwoodi B-type has been redescribed as a new species (M. fallax) (Karssen, 1996). Current research is attempting to clarify the position regarding races, Beek et al. (1999) proposing a pathotype scheme to describe intraspecific variation in pathogenicity.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Hypoaspis aculeifer Predator Eggs
Myrothecium verrucaria Pathogen

Notes on Natural Enemies

Top of page Predatory mites (Hypoaspis aculeifer) were observed feeding on egg-masses of this nematode on tomato roots in Utah, USA (Inserra and Davis, 1983).

M. chitwoodi was relatively susceptible to the nematode-trapping fungi Monacrosporium ellipsosporum and M. cionopagum, compared with Heterodera schachtii, in laboratory tests (Jaffee and Muldoon, 1995). Jacobsen (2002) reviewed biological control of potato pathogens. Wishart et al. (2004) reported attachment of the spores of Pasteuria penetrans and P. nishizawae to juveniles of M. chitwoodi, M. fallax and M. hapla.

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsEggs in soil. Yes
Containers and packaging - woodEggs in soil. Yes
Land vehiclesEggs in soil. Yes
Soil, sand and gravelEggs 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
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
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Leaves
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Impact

Top of page M. chitwoodi reduces the market value of potatoes as a result of internal necrosis and external galling. Necrotic spots in the flesh of tubers of as little as 5% of a crop make it commercially unacceptable. Overall yields of tubers are also reduced. This species is considered to be the major nematode pest of potatoes in the Pacific Northwest states of the USA and the annual predicted loss there would be ca $40 million if control measures were not applied (Santo, 1994). No information is available on the economic impact of this pest in European countries.

Effects on other crops are not as marked nor as well documented, but yields of cereals (wheat, barley, oats and maize) have been shown to be significantly reduced by infestation (Santo and O'Bannon, 1981).

In the Netherlands, M. chitwoodi has recently been found to have damaged potato crops (and certain vegetables) in a limited area in the east of the country. This damage is apparently associated with sandy soils and a succession of warm summers. It is possible that M. chitwoodi has been present in the area for many years, undetected because it caused no significant damage (see Geographical Distribution).

M. chitwoodi has also caused damage to potatoes in Germany (Muller et al., 1996). The multiplication rate and damage probability of M. chitwoodi seems to depend mainly on weather conditions in spring and temperature accumulation during the vegetative period (Braasch et al., 1996).

Chaves and Torres (2000) reported the presence of M. chitwoodi on golf courses in Buenos Aires province, Argentina. Chaves and Torres (2001) reported the presence of M. chitwoodi in potato seed producing areas in Argentina. The nematode occurred in 12.5% of the samples taken.

Diagnosis

Top of page M. chitwoodi has been distinguished from M. fallax by esterase and malate dehydrogenase patterns (Karssen, 1996) and DNA methods (Petersen and Vrain, 1996; Williamson et al., 1997; Zijlstra et al., 1997). A number of molecular based methodologies have been developed to characterize the species and to help differentiate it from related root-knot nematodes (Castagnone-Sereno et al., 1998, 1999; Castagnone-Sereno, 2000; Zijlstra, 2000; Fourie et al., 2001; Wishart et al., 2002; Fargette et al., 2005; Skantar and Carta, 2005).

Detection and Inspection

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The presence of M. chitwoodi in infested soil can be determined by sampling and extraction of the second-stage juveniles, using a standard nematode extraction procedure for free-living nematodes of this size. External symptoms on tubers are obvious in the case of heavy infestations but, where nematode numbers are low or in the early stages of infection, such symptoms are not obvious. Clearing and staining of the tissues can show the presence of nematodes (Hooper, 1986) but this can be a laborious procedure. Storage of lightly infested tubers may lead to the development of obvious external symptoms.

Means of Movement and Dispersal

M. chitwoodi has very limited potential for natural movement; only second-stage juveniles can move in the soil and, at most, only a few tens of centimetres. The most likely method of introducing M. chitwoodi into a new area is through the movement of infected or contaminated planting material. Infected host plants or host products such as bulbs or tubers can easily transport the nematode. The movement of non-host seedling transplants, nursery stock, machinery or other products which are contaminated with soil infested with M. chitwoodi could also result in spread. Infective larvae of this genus have been known to persist for more than one year in the absence of host plants. Nematode movement can also be facilitated by contaminated irrigation water.

Detection based on host plant symptoms, and identification by morphological and molecular methods are detailed in OEPP/EPPO (2009).

Similarities to Other Species/Conditions

Top of page M. chitwoodi closely resembles M. hapla; it can be distinguished from this and other species by the perineal pattern, the appearance of vesicles or vesicle-like structures in the median bulb of the adult female and by the short, blunt, larval tail which has a hyaline, rounded (not tapered) terminus (Golden et al., 1980; Nyczepir et al., 1982; Jepson, 1985).

M. chitwoodi differs from M. fallax by having smaller female and male stylet length, presence of small, irregular outlined male and female stylet knobs, male labial disk not elevated, shorter juvenile body-, tail-, and hyaline tail length, different hyaline tail shape, hemizonid position, esterase and malate dehydrogenase patterns (Karssen, 1996).

Schemes have been described to differentiate M. chitwoodi from M. hapla, M. microtyla and M. incognita by differential host tests (Townshend et al., 1984), and also to distinguish the two races of M. chitwoodi (Mojtahedi et al., 1988). Biochemical methods have also been used. A diagnostic DNA probe which distinguishes M. chitwoodi from M. hapla has been developed by Piotte et al. (1995) and Wishart et al. (2002) used a PCR-based technique involving the ribosomal intergenic spacer. Recently, M. chitwoodi has been distinguished from M. fallax by DNA methods (Petersen and Vrain, 1996; Williamson et al., 1997; Zijlstra et al., 1997; Castagnone-Sereno et al., 1999; Castagnone-Sereno, 2000; Zijlstra, 2000; Wishart et al., 2002).

Prevention and Control

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Chemical Control

Control measures currently used against other root-knot nematodes have proved to be less effective against M. chitwoodi. In north-western USA, crop failures in several potato fields have been attributed to M. chitwoodi despite the use of spring soil fumigations.
Cultural Control and Sanitary Methods

Recent work on sugarbeet suggests that late drilling and reduced irrigation can play a part in reducing damage to crops (Heijbroek, 1996).

Crop rotation with cereals is often used to reduce populations of M. hapla. However, M. chitwoodi reproduces well on wheat, oats, barley and maize. Therefore, rotation with any of these crops will favour a build-up rather than a decrease of M. chitwoodi populations in infested soils. There are, however, other crops which will reduce populations (see Host Range) which can be used in rotations. Brinkman et al. (1996) suggested that chicory cultivars (Cichorium intybus), Dahlia (cv. Vuurvogel) and borage (Borago officinalis) supported low populations of M. chitwoodi and can offer alternatives in crop rotations. However, further studies are necessary to collect more information on crops which can be used in rotation to reduce nematode populations.

Some success has been achieved by the incorporation of green manure into the soil, which reduces population densities of M. chitwoodi (Mojtahedi et al., 1993a, b; Suloiman and Hafez, 1996).

Measures similar to those for potato cyst nematodes (EPPO/CABI, 1992) would appear relevant, i.e. that consignments of rooted plants should come from areas where the pest does not occur or from fields found free from the pest. Suitable survey and test methods have still to be established. Freedom from M. chitwoodi should be specifically assured by certification schemes for seed potatoes.

Host-Plant Resistance

Potato cultivars differ in their tolerance of M. chitwoodi (van Riel, 1993). There is interest in breeding for host resistance to M. chitwoodi, for example, in potatoes, using resistance from Solanum bulbocastanum and other sources (Brown et al., 1991; Austin et al., 1993; Janssen et al., 1997) and cereals (Jensen and Griffin, 1994). Yu (2001) and Yu and Lewellen (2004) registered nematode-resistant sugarbeet germplasm; Zoon et al. (2002) discussed durable resistance against M. chitwoodi and M. fallax whilst Brown et al. (1999, 2003, 2004) investigated resistance to M. chitwoodi in potato and evaluated several wild Solanum species as sources of resistance. Tovar-Soto et al. (1997) reported on the response of five potato genotypes to M. chitwoodi race 2 in Mexico. Berthou et al. (2003) characterized virulence in populations of M. chitwoodi by challenging with Capsicum annuum line PM217. Their results demonstrated great polymorphism in M. chitwoodi populations and the existence of a major gene in pepper controlling a specific resistance against some nematode populations.

IPM Programmes

Guidelines for integrated pest management in potatoes are still being devised in those countries affected by this M. chitwoodi (Ferris et al., 1994a,b; Santo, 1994; Hafez et al., 1997).

References

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Anon, 2005. Meloidogyne chitwoodi and Meloidogyne fallax. Bulletin OEPP, 34(2): 315-320

Austin S, Pohlman JD, Brown CR, Mojtahedi H, Santo GS, Douches DS, Helgeson JP, 1993. Interspecific somatic hybridization between Solanum tuberosum L. and S. bulbocastanum Dun. as a means of transferring nematode resistance. American Potato Journal, 70(6):485-495; 36 ref

Baker RHA, Dickens JSW, 1993. Practical problems in pest risk assessment. In: Ebbels DL, ed. Plant Health and the European Single Market. Farnham, UK: BCPC. pp 209-220

Beek JGvan der, Maas PWT, Janssen GJW, Zijlstra C, Silfhout CHvan, 1999. A pathotype system to describe intraspecific variation in pathogenicity of Meloidogyne chitwoodi. Journal of Nematology, 31(4):386-392; 30 ref

Berthou F, Palloix A, Mugniery D, 2003. Characterisation of virulence in populations of Meloidogyne chitwoodi and evidence for a resistance gene in pepper Capsicum annuum L. line PM 217. Nematology, 5(3): 383-390

Braasch H, Wittchen U, Unger JG, 1996. Establishment potential and damage probability of Meloidogyne chitwoodi in Germany. Bulletin OEPP, 26(3/4):495-509; 21 ref

Brinkman H, Goossens JJM, Riel HRvan, 1996. Comparative host suitability of selected crop plants to Meloidogyne chitwoodi Golden et al. 1980 and M. fallax Karssen 1996. Anzeiger fu^umlaut~r Scha^umlaut~dlingskunde, Pflanzenschutz, Umweltschutz, 69(6):127-129; 13 ref

Brinkman H, van Riel HR, 1990. In: Jaarboek 1989-1990. Wageningen, Netherlands: Directie Gewasbescherming, Plantenziektenkundige Dienst

Brommer E, Molendijk LPG, 2001. Population development of the root-knot nematodes Meloidogyne chitwoodi and M. fallax in potato. Gewasbescherming, 32(6):148-150; 2 ref

Brown CR, Mojtahedi H, Bamberg J, 2004. Evaluation of Solanum fendleri as a source of resistance to Meloidogyne chitwoodi. American Journal of Potato Research, 81(6): 415-419

Brown CR, Mojtahedi H, Santo GS, 1991. Resistance to Columbia root-knot nematode in Solanum ssp. and in hybrids of S. hougasii with tetraploid cultivated potato. American Potato Journal, 68(7):445-452; 17 ref

Brown CR, Mojtahedi H, Santo GS, 1999. Genetic analysis of resistance to Meloidogyne chitwoodi introgressed from Solanum hougasii into cultivated potato. Journal of Nematology, 31(3):264-271; 25 ref

Brown CR, Mojtahedi H, Santo GS, 2003. Characteristics of resistance to Columbia root-knot nematode introgressed from several Mexican and North American wild potato species. Acta Horticulturae, 619: 117-125

CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp

CABI/EPPO, 2000. Meloidogyne chitwoodi. Distribution Maps of Plant Diseases, Map No. 803. Wallingford, UK: CAB International

CABI/EPPO, 2012. Meloidogyne chitwoodi. [Distribution map]. Distribution Maps of Plant Diseases, No.April. Wallingford, UK: CABI, Map 803 (Edition 2)

Castagnone-Sereno P, 2000. Use of satellite DNA for specific diagnosis of the quarantine root-knot nematodes Meloidogyne chitwoodi and M. fallax. Bulletin OEPP, 30(3/4):581-584; 11 ref

Castagnone-Sereno P, Leroy F, Bongiovanni M, Zijlstra C, Abad P, 1999. Specific diagnosis of two root-knot nematodes, Meloidogyne chitwoodi and M. fallax, with satellite DNA probes. Phytopathology, 89(5):380-384; 21 ref

Castagnone-Sereno P, Semblat JP, Leroy F, Abad P, 1998. A new AluI satellite DNA in the root-knot nematode Meloidogyne fallax: relationships with satellites from the sympatric species M. hapla and M. chitwoodi. Molecular Biology and Evolution, 15(9):1115-1122; 37 ref

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