Pratylenchus goodeyi (banana lesion nematode)
- Taxonomic Tree
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Notes on Natural Enemies
- Plant Trade
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Pratylenchus goodeyi Sher & Allen, 1953
Preferred Common Name
- banana lesion nematode
Other Scientific Names
- Anguillulina musicola Apud Goodey, 1932
- Tylenchus musicola Apud Goodey, 1928
- PRATGO (Pratylenchus goodeyi)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Nematoda
- Family: Pratylenchidae
- Genus: Pratylenchus
- Species: Pratylenchus goodeyi
DescriptionTop of page (After Sher and Allen, 1953; de Guiran and Vilardebó, 1962). Female: body slender, after killing by heat almost straight but slightly curved ventrally in the posterior region. Lateral field with four inconspicuous incisures. Lip region with four annules. Stylet 16-18 µm with pronounced knobs flattened anteriorly. Vulva anterior, 71-78%. Genital tract mono-prodelphic. Post-vulval sac about a vulval body width long. Spermatheca large, sub-rectangular in shape and filled with sperm. Tail conoid, ventrally concave with the dorsal contour sinuate just prior to the tail tip. Tail usually with 22-24 ventral annules (range 19-27). Phasmid usually conspicuous, 10-14 annules from the tail tip.
Measurements (after Sher and Allen, 1953). Female: L=0.64-0.68 mm; a=27-37; b=5.5-6.1; c=16-18; V=(30-41)73-75(3.4-4.5). Male: L=0.55-0.57 mm; a=26; b=5.4-5.8; c=17-18.
DistributionTop of page The distribution of P. goodeyi is very closely linked to altitude and temperature; it is found almost exclusively in the upland areas known as the afro-montane highlands of Africa (Bridge, 1988; Sarah, 1989; Bridge et al., 1995; Price and Bridge, 1995). In Uganda, it is the only banana nematode root pest found over 1600 m above sea level (Kashaija et al., 1994). In Africa, it is a pest of banana, plantain and ensete grown in cooler climates outside the tropical lowlands, and has a lower temperature preference than the other major nematode pest of bananas, Radopholus similis. P. goodeyi had been considered a species indigenous to East Africa, where it is recognized as an important pest of highland bananas in Uganda, Tanzania, Kenya, Rwanda and Burundi, and of ensete in Ethiopia (Gichure and Ondieki, 1977; Bridge, 1988; Sarah, 1989; Peregrine and Bridge, 1992; Kashaija et al., 1994). However, it is also found in the highlands of Cameroon, West Africa (Bridge et al., 1995). It is the main nematode pest of bananas in the Canary Islands (de Guiran and Vilardebó, 1962), and the species has also been reliably identified from the Nile delta region, from Crete (Oteifa, 1962; Vovlas et al., 1994), and from one locality in Australia (New South Wales) (Bridge et al., 1997).
In East and West Africa, P. goodeyi is exclusively a pest of smallholder cultivation and is generally absent from the commercial banana plantations of the lowland areas. This would explain why the species has not been disseminated worldwide, as have other nematode pests on commercial bananas (Bridge et al., 1995).
Because of the close association of P. goodeyi with the highland or afro-montane areas in Africa, knowledge of these areas enables predictions to be made as to the possible distribution of P. goodeyi. Price and Bridge (1995) suggest that the nematode probably also occurs at altitude on the island of Fernando Po (Equatorial Guinea), just off the Cameroon coast, and could also occur in highland areas in Malawi, Mozambique and southern Africa. Additionally, there are strong similarities between the afro-montane flora of Cameroon and those of the Nimba Highlands of the Guinea/Sierra Leone/Côte d'Ivoire region, and also with the Western Angolan Escarpment, and it is possible that P. goodeyi will also be found in both these regions.
Distribution TableTop 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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|China||Present||Present based on regional distribution.|
|-Hainan||Present||Zhang et al., 2015|
|Pakistan||Present||Aatika et al., 2017|
|Burundi||Present||Bridge, 1988; CABI/EPPO, 2000|
|Cameroon||Present||Bridge et al., 1995; CABI/EPPO, 2000|
|Egypt||Present||Oteifa, 1962; CABI/EPPO, 2000|
|Ethiopia||Present||Peregrine and Bridge, 1992; CABI/EPPO, 2000; Berg and Mekete, 2010|
|Kenya||Present||Gichure and Ondieki, 1977; CABI/EPPO, 2000|
|Rwanda||Present||Sarah, 1989; CABI/EPPO, 2000|
|-Canary Islands||Present||de Guiran & Vilardeho, 1962; CABI/EPPO, 2000|
|Tanzania||Present||Bridge, 1988; CABI/EPPO, 2000|
|Uganda||Present||Machon & Hunt, 1985; CABI/EPPO, 2000|
|Greece||Present||Vovlas et al., 1994; CABI/EPPO, 2000|
|Portugal||Restricted distribution||CABI/EPPO, 2000|
|Spain||Restricted distribution||CABI/EPPO, 2000|
|Australia||Restricted distribution||Bridge et al., 1997; CABI/EPPO, 2000|
|-New South Wales||Present||Bridge et al., 1997; CABI/EPPO, 2000|
|-Queensland||Present||Pattison et al., 2002|
Risk of IntroductionTop of page Endoparasitic nematodes of banana such as P. goodeyi are disseminated on banana vegetative planting material, and the presence of P. goodeyi in the Canary Islands, Crete and Egypt is almost certainly due to their introduction on planting material. Bananas were introduced to the Canary Islands in the 19th century, in part based on Cavendish banana plants raised in greenhouses in the UK. Banana in the Palm House at the Royal Botanic Gardens, Kew, UK, is the type locality of P. goodeyi, which may therefore have been introduced to the Canaries and Egypt on infected plants from European greenhouses. Plant quarantine procedures should be followed to prevent the further dissemination of P. goodeyi to other warm, temperate banana-growing areas such as Spain or Turkey where it is not yet established (Price and Bridge, 1995).
Hosts/Species AffectedTop of page The few investigations that have been made on hosts of P. goodeyi show that the nematode has a narrow host range.
Growth StagesTop of page Flowering stage, Fruiting stage, Pre-emergence, Seedling stage, Vegetative growing stage
SymptomsTop of page The endoparasitic root feeding of P. goodeyi produces necrotic lesions and destruction of cortical tissues. The lesions are initially yellow, but change to a characteristic purple and then brown when examined internally. The root surface over necrosed tissues turns black. The ensuing reduction in nutrient and water uptake causes a stunting of growth, nutrient deficiency including chlorosis, small bunches, and weak or reduced followers in bananas and plantains. Nematode feeding on tissues can cause breaking of roots around the areas of necrotic tissue, leading to toppling or uprooting of plants and the complete loss of bunches. Stunted, severely infested bananas with relatively small bunches generally have less tendency to topple compared with taller infested plants with larger and heavier bunches.
List of Symptoms/SignsTop of page
|Roots / cortex with lesions|
|Roots / internal feeding|
|Roots / soft rot of cortex|
|Roots / stubby roots|
Biology and EcologyTop of page P. goodeyi is a migratory endoparasitic nematode feeding in the cortex of banana, plantain and ensete roots, causing brown to purplish lesions which can spread throughout a large proportion of the roots. All stages of the nematode can invade the roots; eggs are laid in the root tissues or in the soil. The species multiplies at a significantly greater rate at lower temperatures of 16-21°C, and considerably less at higher temperatures of 25°C and above, compared with other, similar endoparasitic nematodes of banana (Pinochet et al., 1995). This accounts for its distribution in the cooler highlands of Africa and its presence in the Canary Islands and the Mediterranean.
Notes on Natural EnemiesTop of page No natural enemies have been recorded for P. goodeyi.
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bulbs/Tubers/Corms/Rhizomes||adults; eggs; juveniles||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||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Seedlings/Micropropagated plants||adults; eggs; juveniles||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)|
ImpactTop of page P. goodeyi is the main nematode species and a major pest causing yield losses in East African highland bananas (Musa AAA-EA) in the highlands of Tanzania, Uganda and Kenya, and in highland bananas and plantains at high elevations in Cameroon (Gichure and Ondieki, 1977; Sarah, 1989; Sikora et al., 1989; Kashaija et al., 1994) and of ensete in Ethiopia (Peregrine and Bridge, 1992). In the Canary Islands, P. goodeyi is a widespread root pest of bananas and a cause of production losses; here, as also in Crete and Egypt, P. goodeyi has been recorded as primarily a parasite of Cavendish bananas (de Guiran and Vilardebó, 1962; Oteifa, 1962; Vovlas et al., 1994).
In the main highland banana-growing area of Tanzania, P. goodeyi was found in 94.5% of farms sampled, and was directly associated with toppling and other symptoms of crop damage in 84% of these farms (Bridge, 1988).
DiagnosisTop of page P. goodeyi can be extracted from both soil and roots using standard nematological techniques (Southey, 1986; Hooper, 1990), but root extractions will give more meaningful results in relation to root and crop damage. The methods used for diagnosis are similar to those described for Pratylenchus coffeae and Radopholus similis. The nematodes can be detected by inspecting and extracting nematodes from roots and bases of pseudostems (corms) in banana, plantain and ensete.
For bananas and plantains, root samples are collected by digging a rectangular trench 20 cm wide by 20 cm deep, 25 cm from the corm of the mother (oldest) stem. Roots are cut near the corm and at the end of the trench; surface roots and those that are dried and shrivelled are discarded. The plant material is carefully washed free of soil and/or debris before extracting. A representative root sample taken from one or a number of plants is chopped in 0.5 cm lengths, mixed thoroughly, and a 5-10 g subsample taken for processing. A 24-h period of incubation is sufficient for macerated root samples. Chopped roots should be incubated for 2-4 days and mist extractions may be run for up to 14 days in some laboratories. Nematode populations are normally shown as per 100 g of fresh roots, although this amount is not used for extraction.
Chopped plant material: most endoparasitic nematodes will move out of plant material if it is first chopped into 0.5-1 cm lengths or portions. These are placed on the tissue of a Baermann funnel or on a fine sieve, etc. and kept in a Petri dish or similar container. This is left for up to 72 h, but can be examined earlier.
Macerated plant material: roots are cut into short lengths and placed in an electric mixer with a quantity of water. Maceration for 5 s at half speed and then 10 s at full speed is normally sufficient to break open the plant material without damaging the nematodes. The macerated material is poured onto the tissue of a Baermann funnel or a fine sieve in a Petri dish, and left as above.
Mistifier technique: this requires more elaborate apparatus. A continuous fine mist of water is sprayed onto chopped plant material suspended on a sieve over a funnel. The emerging nematodes are washed into the funnel and collected in a container, where they settle to the bottom and excess water overflows.
Hydrogen peroxide technique (Gowen and Edmunds, 1973): this method, which has been shown to be most effective, consists of maceration of roots, washing through a bank of sieves and then incubation of the remaining root material in dilute (1%) hydrogen peroxide and some liquid detergent (1 ml/l) for about 2 days. The recovery of nematodes from chopped rather than macerated roots is reduced, possibly because of oxygen starvation or exhaustion of the nematodes. Washing after maceration removes inactive and damaged nematodes that would not be recovered by incubation; it also washes away any phenolic compounds released by the roots (for example, bananas) which may inactivate nematodes during incubation. For the incubation period hydrogen peroxide is used both to provide oxygen for the nematodes and to oxidize any further phenols that may be released by the roots. Liquid detergent ensures that all surfaces of the macerated root tissue are thoroughly wetted.
Direct recovery techniques using maceration and sieving (Vilardebó et al., 1972), or maceration and flocculation/flotation (Escobar and Rodriguez-Kabana, 1980; Hooper, 1990) can also be used.
Where nematologists or laboratory facilities are unavailable, nematode damage is sometimes assessed by recording the incidence of toppling/uprooting per hectare per month (Tarté and Pinochet, 1981). This may also be correlated with assessments of necrosis on primary root and on rhizomes taken from randomly selected plants from a plantation (Stover, 1972; Tarté and Pinochet, 1981; Bridge, 1988; Sikora et al., 1989).
Assessment of root necrosis caused by P. goodeyi is a useful and practical means of determining damage in the field. As for extraction, root samples are collected by digging a rectangular trench 20 cm wide by 20 cm deep, 25 cm from the corm of the mother (oldest) stem. Five or six roots approximately 10-15 cm long are used for assessment from each plant. The roots are cut longitudinally to expose the cortex and stele (central vascular cylinder) and laid side by side. The severity of root damage is assessed by estimating the amount of cortical tissue necrosed as a percentage of total cortical tissues exposed in the six roots (Broadley, 1979; Pinochet, 1988; Bridge, 1988; Bridge and Gowen, 1993).
The other symptoms of damage related to banana root destruction by nematodes are toppling of plants and stunted growth, poor followers, and reduced bunch size. Toppling caused by nematodes is characterized by uprooting of the corm with the remains of necrosed roots still attached. Assessing the amount of toppling/uprooting in a plantation can help in determining the extent of nematode damage.
Detection and InspectionTop of page P. goodeyi can be detected in the roots and corms of banana, plantain and ensete by the characteristic purplish necrosis that occurs in nematode lesions throughout the cortex. This necrosis does not extend into the stele, which remains white in growing roots. The symptoms can be most easily observed if roots are cut longitudinally. Two similar nematodes (Pratylenchus coffeae and Radopholus similis) also cause the same symptoms (see Similarities to Other Pests), and in order to distinguish them they have to be extracted from roots and examined microscopically.
Similarities to Other Species/ConditionsTop of page Distinguishing between species of the genus Pratylenchus can be difficult without expert taxonomic assistance. The other nematodes found in roots of bananas and plantains, with which P. goodeyi may be confused, include another species of Pratylenchus, P. coffeae, and the banana burrowing nematode Radopholus similis. All three species produce the same and indistinguishable symptoms of root and crop damage.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.Introduction
The different practices used for controlling P. goodeyi are the same as for other lesion nematodes of banana, Pratylenchus coffeae and Radopholus similis. In permanent cultivation, the opportunities for control are limited to regular nematicide treatment; however, in subsistence cultivation the only realistic or economically justifiable techniques for preserving losses due to nematodes may be by applying large quantities of mulch to stimulate root growth, and by propping fruit stems. Established practices for decreasing nematode populations in different banana-growing systems are described here (after Gowen and Quénéhervé, 1990; Quénéhervé, 1993).
1. Rotation or break with non-host crops, e.g. cassava.
2. Flooding for 8 weeks after having destroyed previous banana crop.
3. Fallow in absence of banana volunteers for 10-12 months.
4. Use of disease-free suckers (from nurseries or clean land) as planting material.
5. Use of in vitro-produced plants.
6. Paring diseased tissue from corms.
7. Paring and leaving large corms in the sun for 14 days.
8. Heat treatment (hot water, solarization) of corms before planting.
9. Coating corms with nematicide in mud.
10. Applying nematicide to planting hole and in-fill soil.
11. Regular spot applications with granular or liquid nematicide formulations.
1. Regular spot applications with granular or liquid nematicide formulations.
2. Growing resistant or tolerant cultivars.
3. Heavy mulches with organic wastes.
4. Propping fruiting stems with poles or with string guy ropes to prevent plants uprooting.
ReferencesTop of page
Aatika S; Nasira K; Shahina F, 2017. Description of Filenchus maqbooli n. sp., and redescriptions of five new records of plant parasitic nematodes of maize crops from Punjab, Pakistan. Pakistan Journal of Nematology, 35(1):47-64.
Bridge J, 1988. Plant nematode pests of banana in East Africa with particular reference to Tanzania. In: INIBAP 1988. Nematodes and the borer weevil in bananas: present status of research and outlook. Proceedings of a workshop held in Bujumbura, Burundi, 7-11 December, 1987, 35-39.
Bridge J, 1993. Worldwide distribution of the major nematode parasites of bananas and plantains. Biological and integrated control of highland banana and plantain pests and diseases: Proceedings of a Research Coordination Meeting, Cotonou, Benin 12-14 November 1991, November:185-198; 77 ref.
Bridge J; Fogain R; Speijer P, 1997. Les nématodes parasites des bananiers. Pratylenchus coffeae (Zimmermann, 1898) Filip.& Schu. Stek, 1941, Pratylenchus goodeyi Sher & Allen, 1953. Parasites et ravageurs des Musa: fiche technique no 2. Montpellier, France: INIBAP.
Bridge J; Gowen SR, 1993. Visual assessment of plant parasitic nematode and weevil damage on bananas and plantains. In: Gold CS, Gemmill B, eds. Biological and Integrated Control of Highland Banana and Plantain Pests and Diseases. Proceedings of a Research Coordination Meeting, Cotonou, Benin, 12-14 November, 1991, 147-154.
Broadley RA, 1979. A simple method for estimating banana root rot. Australian Plant Pathology, 8:24-25.
Gowen S; Queneherve P, 1990. Nematode parasites of bananas, plantains and abaca. In: Luc M, Sikora RA, Bridge J, eds. Plant Parasitic Nematodes in Subtropical and Tropical Agriculture. Wallingford, UK: CAB International, 431-460.
Gowen SR; Edmunds JE, 1973. An evaluation of some simple extraction techniques and the use of hydrogen peroxide for estimating nematode populations in banana roots. Plant Disease Reporter, 57(8):678-681
Guiran de G; Vilardebo A, 1962. Le bananier aux îles Canaries. IV. Les nématodes parasites du bananier. Fruits, 17:263-277.
Hooper DJ, 1990. Extraction and processing of plant and soil nematodes. In: Luc M, Sikora RA, Bridge J, eds. Plant Parasitic Nematodes in Subtropical and Tropical Agriculture. Wallingford, UK: CAB International, 45-68.
Kashaija IN; Speijer PR; Gold CS; Gowen SR, 1994. Occurrence, distribution and abundance of plant parasitic nematodes of bananas in Uganda. African Crop Science Journal, 2:99-104.
Mbwana ASS; Waudo SW; Seshu-Reddy KV, 1995. Host-range of the lesion nematode, Pratylenchus goodeyi, commonly found in highland bananas of East Africa. International Journal of Pest Management, 41(1):46-49; 34 ref.
Oteifa BA, 1962. Species of root-lesion nematodes commonly associated with economic crops in the Delta of the U.A.R. Plant Disease Reporter, 46:712-714.
Pattison AB; Stanton JM; Cobon JA; Doogan VJ, 2002. Population dynamics and economic threshold of the nematodes Radopholus similis and Pratylenchus goodeyi on banana in Australia. International Journal of Pest Management, 48(2):107-111.
Pinochet J, 1988. A method for screening bananas and plantains to lesion forming nematodes. Nematodes and the borer weevil in bananas. Present status of research and outlook. Proceedings of a workshop held in Bujumbura, Burundi, 7-11 December, 1987. International Network for the Improvement of Banana and Plantain, 62-65.
Quénéhervé P, 1993. Nematode management in intensive banana agrosystems: comments and outlook from the Côte d'Ivoire experience. Crop Protection, 12:164-172.
Sarah JL, 1989. Banana nematodes and their control in Africa. Nematropica, 19:199-215.
Sikora RA; Bafokuzara ND; Mbwana ASS; Oloo GW; Uronu B; Reddy KVS, 1989. Interrelationship between banana weevil, root lesion nematode and agronomic practices, and their importance for banana decline in the United Republic of Tanzania. FAO Plant Protection Bulletin, 37(4):151-157
Speijer PR; Gold CS; Karamura EB; Kashaija IN, 1994. Banana weevil and nematode distribution patterns in highland banana systems in Uganda: Preliminary results from a diagnostic survey. African Crop Science Conference Proceedings, 1:285-289.
Speijer PR; Sikora RA, 1993. Influence of a complex disease involving Pratylenchus goodeyi and a non-pathogenic strain of Fusarium oxysporum on banana root health. Biological and integrated control of highland banana and plantain pests and diseases: Proceedings of a Research Coordination Meeting, Cotonou, Benin 12-14 November 1991, November:231-239; 23 ref.
Tarté R; Pinochet J, 1981. Nematological problems of banana. Recent findings and control methods. Problemas nematologicos del banano. Contribuciones recientes a su conocimiento y combate. Union de Paies Exportadores de Banano (UPEB) Panama, 32 pp.
Vilardebó A; Aubert A; Beugnon M; Melin Ph; Guerout R; LassoudiFre A; Martin P; Pinon A, 1972. Techniques experimentales et progres dans la lutte contre les nematodes en culture bananiere. Phytiatrie-Phytopharmacie, 21:129-139.
Zhang F; Yan S; Zhou Y; Guo G; Guo S; Jin Z; Zeng H; Peng D; Ruan L; Sun M, 2015. First report of Pratylenchus goodeyi on banana in Hainan Province, China. Plant Disease, 99(5):731-732. http://apsjournals.apsnet.org/loi/pdis
Distribution MapsTop of page
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