Preferred name: Meloidogyne fallax
Authority: Karssen
Taxonomic position: Animalia: Nematoda: Chromadorea: Rhabditida: Meloidogynidae
Common names in English: false Columbia root-knot nematode
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Notes on taxonomy and nomenclature EPPO Categorization: A2 list
EU Categorization: A2 Quarantine pest (Annex II B)
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EPPO Code: MELGFA

M. fallax was detected on and described from potato (Solanum tuberosum). Host-range includes a wide range of dicotyledonous and monocotyledons plants, including weeds, ornamentals, and economically important crops such as carrot (Daucus carota), black salsify (Scorzonera hispanica) and tomato (Solanum lycopersicum). The experimental host range of M. fallax mostly overlaps that of M. chitwoodi, but differential hosts have been found. For example dwarf beans (Phaseolus vulgaris), valerian (Valeriana officinalis), maize (Zea mays), Erica cinerea and Potentilla fruticosa are good hosts for M. chitwoodi and not for M. fallax, while the reverse is the case for Oenothera glazioviana, Phacelia tanacetifolia, Hemerocallis cv. Rajah and Dicentra spectabilis (Brinkman et al., 1996). It is expected that many more plant species will be hosts of M. fallax than currently known, since this is the case also with other, closely related root knot nematodes.

Host list: Allium porrum, Asparagus officinalis, Avena strigosa, Beta vulgaris, Cichorium intybus, Cynara scolymus, Daucus carota subsp. sativus, Fragaria x ananassa, Hemerocallis sp., Hordeum vulgare, Lactuca sativa, Lamprocapnos spectabilis, Leptinella sp., Lolium multiflorum, Medicago sativa, Oenothera glazioviana, Phacelia tanacetifolia, Scorzonera hispanica, Solanum lycopersicum, Solanum nigrum, Solanum physalifolium, Solanum tuberosum, Trifolium repens, Triticum aestivum

After the first record near Baexem (NL) in 1992, M. fallax was recorded on potato at several locations in the southern and south-eastern part of the Netherlands (Karssen, 1996), close to the German and Belgium borders. Within the EPPO region it was detected locally in Belgium, France, Germany, Sweden, Switzerland and the United Kingdom (England). In addition, Topalović et al. (2017) revealed that a Meloidogyne species detected in Ireland in 1965 belongs to M. fallax. M. fallax has never been reported from the natural environment in Europe. Outside Europe it has been reported from Australia, Chile, New Zealand and South Africa. New Zealand is the only known country where M. fallax is widely distributed (North and South Island) and detected in cropping and pasture fields (Rohan et al., 2016), strongly suggesting that it could be the place of origin of this pest. 

EPPO Region: Belgium, France (mainland), Germany, Netherlands, Sweden, Switzerland, United Kingdom (England)
Africa: South Africa
Asia: Indonesia (Java)
South America: Chile
Oceania: Australia (Queensland, South Australia, Tasmania, Victoria, Western Australia), New Zealand

The life cycles of M. fallax and M. chitwoodi are, in general, the same with respect to root penetration, gall induction, symptomatology, number of moults, parthenogenetic reproduction and chromosome number: Both M. chitwoodi and M. fallax usually reproduce by parthenogenesis. They can have one to three generations per year in the Netherlands and produce several hundreds of eggs per female, deposited in an egg sac. These egg sacs allow the eggs to survive under unfavorable conditions (EFSA, 2019). Initial results by van der Beek (1997) indicated that M. fallax had a shorter life cycle than M. chitwoodi in a virulence study on potato.

Host races, as described for M. chitwoodi, have not been detected for M. fallax so far. Successful hybridization was not obtained when M. fallax and M. chitwoodi were crossed in two different experiments; the F1 was viable, but the F2 second-stage juveniles were not viable and showed morphological distortions (van der Beek & Karssen, 1997).

In addition, differences in terms of hatching responses to root diffusate and host age between M. chitwoodi and M. fallax suggests different survival strategies between these two species (Wesemael et al., 2006).

M. fallax, M. chitwoodi and M. minor Karssen are closely related, morphologically and at DNA level. Phylogenetically they appear to be in the same distinct clade within the genus Meloidogyne (Holterman et al., 2009; Elling, 2013).

Symptoms

Above-ground symptoms of heavily infested plants include stunting and yellowing, while below ground galling is typical. The root galls produced by M. chitwoodi and M. fallax are comparable to those produced by several other root-knot species, i.e. relatively small galls in general without secondary roots emerging from them (these secondary roots are seen in M. hapla). On potato tubers, M. chitwoodi and M. fallax cause numerous small pimple-like raised areas on the surface (in M. hapla these swellings are not evident). Some potato cultivars, although heavily infested, may be free from visible external symptoms, while the internal potato tissue is necrotic and brownish, just below the peel (EPPO, 2016a).

Morphology

Sedentary females are annulated, pearly white and globular to pear-shaped, 400-720 µm long and 250-460 µm wide. The stylet is dorsally curved, 13.9-15.2 µm long, with rounded to ovoid stylet knobs, slightly sloping posteriorly. The non-sedentary males are vermiform, annulated, slightly tapering anteriorly, bluntly rounded posteriorly, 735-1520 µm long and 27-44 µm wide. The stylet is 18.9-20.9 µm long, with large rounded knobs, set off from the shaft. The non-sedentary second-stage juveniles are vermiform, annulated, tapering at both ends, 380-435 µm long, 13.3-16.4 µm wide, with a tail length of 46-56 µm and a hyaline tail part 12.2-15.8 µm in length (Karssen, 1996).

M. fallax is closely related morphologically to M. chitwoodi, and this misleading resemblance was the reason for giving the species its name. The most striking differences for males and females are stylet length (longer for M. fallax) and stylet knob shape (M. fallax: prominent and rounded; M. chitwoodi: small and irregular). The second-stage juveniles differ in mean body length, tail length and hyaline tail length (all longer for M. fallax). With the scanning electron microscope, it can be observed that the male head of M. fallax has an elevated labial disk. Differences exist in the female perineal pattern (M. fallax has a relatively higher dorsal arch and thicker striae) (EPPO, 2016a; see also Karssen, 2002). 

The species can be reliably distinguished by morphological observation of females, males, and second-stage juveniles in combination with biochemical (isozyme electrophoresis) or molecular (PCR) methods; see EPPO, 2016. To predict the amount of potato tuber damage caused by M. fallax, a quantitative DNA-test was developed for soil (Hay et al., 2016).

Detection and inspection methods

Specific guidance on the sampling of soil and potato tubers is given in the EPPO Standards PM 9/17 (EPPO, 2013b), PM 3/71 (EPPO, 2007) and PM 3/69 (EPPO, 2019a). Populations in the soil rapidly decline in the absence of a host and nematodes reproduce better on a good host. Therefore, detection of the nematodes through field inspection and soil sampling is more sensitive if done as close as possible to the time of harvest of a host crop, targeting particularly susceptible plants (EPPO, 2013b). In each lot of potato (typically 25 tonnes), 200 tubers are randomly sampled and processed (EPPO, 2019a). 

Nematode extraction and identification should be carried out according to EPPO Standards PM 7/119 (EPPO, 2013a) and PM 7/41 Diagnostic protocol for Meloidogyne chitwoodi and M. fallax (EPPO, 2016a).

Historically, in order to distinguish M. fallax from other Meloidogyne spp., molecular methods have been used. Karssen et al. (1995) discriminated M. fallax and M. chitwoodi females by their esterase (EC 3.1.1.1) and malate dehydrogenase (EC 1.1.1.37) isozyme patterns, using the general method of Esbenshade & Triantaphyllou (1985) for identification of female Meloidogyne species by isozyme electrophoresis. Additionally, the isozyme glucose 6-phosphate dehydrogenase (EC 1.1.1.49) was used to differentiate the two species (van der Beek & Karssen, 1997). van der Beek et al. (1997) and Tastet et al. (1999) used mini two-dimensional gel electrophoresis to study the total soluble protein patterns of M. hapla, M. chitwoodi and M. fallax, and confirmed these species to be distinct biological groups.  An overwhelming number of molecular tests has been developed to identify M. fallax from soil, root or tubers and to separate it from related species such as M. chitwoodi, including PCR (Peterson & Vrain, 1996, Peterson et al., 1997; Wishart et al., 2002), PCR RFLP (Zijlstra et al. 1995, Zijlstra, 1997) AFLP (v.d. Beek et al., 1998; Fargette et al., 2005), SCAR (Zijlstra, 2000; Fourie et al., 2001), real time TaqMan PCR (Zijlstra & v. Hoof, 2006; de Haan et al., 2014), RAPD (Adam et al., 2007), satDNA (Castagnone-Sereno et al., 1999; Castagnone-Sereno, 2000; Mestrovich et al., 2009 & 2013), LAMP (Zhang & Gleason, 2019), HRMC (Holterman et al., 2012), barcoding (Hodgetts et al., 2016, EPPO, 2016b). A serological test was also developed to separate M. fallax from M. chitwoodi and other root-knot nematodes (Tastet et al., 2001). Only a selection of the above-mentioned tests are recommended for identification (see EPPO, 2016a).

M. fallax 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 pathway for introducing M. fallax into a new area is through the movement of infested or contaminated planting material. Infested host plants or host products such as bulbs or tubers can easily transport the nematode. The movement of non-host plants for planting (e.g. seedling transplants, nursery stock), non-host plant products (e.g. bulbs, tubers, corms and rhizomes), equipment and machinery which are contaminated with soil infested with M. fallax could also result in spread (EPPO, 2013b). Soil as such is also a possible pathway. Infective juveniles 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.

Economic impact

In trials, M. fallax caused the same symptoms on potato tubers, black salsify and carrots as M. chitwoodi, i.e. external galling and internal necrosis just below the skin (Brinkman et al., 1996; van Riel & Goossens, 1996). The reported natural outbreaks of M. fallax on potato showed these external symptoms (Karssen, 1996). Goosens (1995) reported infected Asparagus officinalis and several ornamentals with root-knots in an experimental field with an infestation of M. fallax. M. fallax sometimes occurs in mixed infestations with M. chitwoodi (Wesemael et al., 2006). 

Meloidogyne fallax mainly induces quality losses caused by cosmetic damage on black salsify, leek, carrot and potato. If the level of infection is high a complete rejection of these crops is possible. So far, this nematode has a limited damaging effect on other known host crops (EFSA, 2012).

In countries where M. fallax is regulated, traded plants and plant products infested with M. fallax may need to be destroyed.

Control

There is no direct practical experience of the control of M. fallax. Research on M. fallax in the Netherlands has focused on host suitability, damage thresholds, effect of fallow, the use of green manure crops and time of sowing. The first results indicate that fallow for one year reduced the population by more than 95%, but this reduction was not sufficient to ensure that subsequent crops met quality standards. There was less damage in sugar beet and carrot when these crops were sown later in spring. Farmers are advised to be careful when growing green-manure crops on infested fields, because some species are suitable hosts for M. fallax. Phaseolus vulgaris was the only tested crop with no reproduction of M. fallax, while maize and cereals were poor hosts (Brommer, 1996). Several green manures were tested, and no reproduction of M. fallax was found on Eruca Sativa cv Trio, Tagetus patula and Borago spp., while Raphanus sativus was a poor host. Avena strigosa turned out to be a very good host (Visser & Molendijk, 2015).

Janssen et al. (1996) tested several wild tuber-bearing Solanum species, to determine the level of resistance to M. hapla, M. chitwoodi and M. fallax. High resistance to M. chitwoodi and M. fallax was observed in genotypes of S. bulbocastanum, S. hougasii, S. cardiophyllum, S. fendleri and S. brachistotrichum. Differential resistance between M. chitwoodi and M. fallax was observed in S. chacoense, S. stoloniferum and S. gourlayi. Resistance to M. fallax was also found in Solanum sparsipilum (Kouassi et al., 2004) and the resistance gene was studied (Kouassi et al., 2006). By crossing wild tuber bearing Solanum species with Solanum tuberosum, successful introgression of resistance to M. fallax and M. chitwoodi into potato was obtained (Janssen et al., 1997).

In addition, useful resistance against M. fallax was found in sea beet (Beta maritima) (Yu et al., 1999) and used to develop a resistant sugar beet line (Beta vulgaris) (Yu, 2001).

Wesemael & Moens (2012) tested ten common bean (Phaseolus vulgaris) cultivars and noted that all these cultivars were non-hosts or poor hosts for M. fallax.

An interesting potential control method consists of using the hyper-parasitic bacterium Pasteuria spp. So far three different species of Pasteuria are reported to be able to parasitize M. fallax: P. penetrans, P. nishizawae and P. hartismeri (Wishart et al., 2004; Bishop et al., 2007).

Phytosanitary risk

Because M. fallax occurs in crops and situations which are similar to those for M. chitwoodi, and the two species are closely related and difficult to distinguish, M. fallax presents a phytosanitary risk similar to that of M. chitwoodi (EFSA, 2012). M. fallax may be able to establish in a large proportion of its host area but may only cause significant damage in certain areas and under certain conditions, causing complete crop rejection (e.g. on potato, carrot and/or black salsify). Soils with a coarse texture such as sandy and sandy-loam soils have a higher probability of being contaminated. Narrow rotation or rotation with alternative hosts facilitates a rapid build-up of population levels and therefore increases the risk for establishment of M. chitwoodi and M. fallax in a particular field. As observed since 1992, M. fallax is expected to spread slowly (EFSA, 2012).

Measures similar to those for other root-knot nematodes would appear relevant, i.e. that rooted host plants for planting (with or without soil), non-host plants for planting with soil attached and plant products with soil attached come from a pest free area, a pest free place of production or are produced under protected cultivation. Alternatively, soil from non-host plants for planting or plant products can be removed. Soil as such can originate from a pest free area or a pest free place of production. Used machineries, equipment, vehicles, and passengers’ shoes can be cleaned. Publicity would allow to enhance public awareness on M. fallax when travelling. 

Specific requirements are recommended in EPPO Standard PM 8/1 Commodity-specific phytosanitary measures for Potato (EPPO, 2017) for seed and ware potatoes to be imported from third countries. In this Standard, seed potato freedom from M. fallax can also be guaranteed by testing the seed potatoes after harvest following EPPO Standard PM 3/69 Meloidogyne chitwoodi and M. fallax: sampling potato tubers for detection (EPPO, 2019a). Ware potatoes freedom can also be guaranteed by implementing EPPO Standard PM 9/17 National regulatory control system for Meloidogyne chitwoodi and M. fallax (EPPO, 2013b). EPPO Standard PM 3/61 details conditions for establishing pest-free areas and pest-free production and distribution systems for quarantine pests of potato (EPPO, 2019b).

Measures to contain or eradicate M. chitwoodi and M. fallax are described in the national regulatory control system (EPPO, 2013b).

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Zijlstra C (2000) Reliable identification of quarantine root-knot nematodes Meloidogyne chitwoodi and M. fallax by PCR-based techniques. EPPO Bulletin 30, 575-579.

Zijlstra C & van Hoof RA (2006) A multiplex real-time polymerase chain reaction (TaqMan) assay for the simultaneous detection of Meloidogyne chitwoodi and M. fallax. Phytopathology 96, 1255-1262. 

This datasheet was extensively revised in 2020 by Karssen G. His valuable contribution is gratefully acknowledged.

EPPO (2024) Meloidogyne fallax. EPPO datasheets on pests recommended for regulation. https://gd.eppo.int (accessed 2024-11-21)

This datasheet was first published in the EPPO Bulletin in 1999 and is now maintained in an electronic format in the EPPO Global Database. The sections on 'Identity', ‘Hosts’, and 'Geographical distribution' are automatically updated from the database. For other sections, the date of last revision is indicated on the right.

EPPO (1999) EPPO Data sheets on quarantine pests - Meloidogyne fallax.  EPPO Bulletin 29(4), 493-496. https://doi.org/10.1111/j.1365-2338.1999.tb01425.x