RESEARCH/INVESTIGACIÓN

PLANT PARASITIC ASSOCIATED WITH BANANA AND PLANTAIN IN EASTERN AND WESTERN DEMOCRATIC REPUBLIC OF CONGO M. Kamira1, 3, S. Hauser2, P. van Asten1,2, D. Coyne2, and H. L. Talwana3 1Consortium for Improving Agricultural-based Livelihoods in Central Africa (CIALCA) Project, Bukavu, Democratic Republic of Congo; 2International Institute of Tropical Agriculture (IITA); 3School of Agricultural Sciences Makerere University, Kampala, Uganda; Corresponding author [email protected]

ABSTRACT Kamira M., S. Hauser, P. Van Asten, D. Coyne, and H. L. Talwana. 2013. Plant parasitic nematodes associated with banana and plantain in eastern and western Democratic Republic of Congo. Nematropica 43:216-225. Plant-parasitic incidence, population densities and associated damage were determined from 153 smallholder banana and plantain gardens in Bas Congo (9 – 646 meters above sea level, m.a.s.l) and South Kivu (1043 – 2005 m.a.s.l), Democratic Republic of Congo, during 2010. Based on the frequency of total nematode soil and root extraction, Helicotylenchus multicinctus (89%), Meloidogyne spp. (54%) and (30%) were the most widespread, while goodeyi (18%) Helicotylenchus dihystera (18%), Rotylenchulus reniformis (14%), and Pratylenchus spp. (6%) were localized in occurrence. The occurrence and abundance of the nematode species was influenced by altitude:R. similis declined at elevations above 1300 m; P. goodeyi declined at elevations below 1200 m; H. multicinctus and Meloidogyne spp. were found everywhere with higher but non-dominant densities at lower altitudes; Pratylenchus spp. was restricted to lower altitudes; while H. dihystera and R. reniformis were scattered at both low and high altitudes. Helicotylenchus multicinctus occurred in higher population densities than other nematode species, especially at the lower elevations. Damage to banana roots (percentage necrotic root tissue) was moderate to high, irrespective of altitude and banana type. To facilitate development of pragmatic plant parasitic nematode management, such baseline studies provide information upon which to enable informed and suitable strategies, such as cultivar selection and distribution. Key words: D.R. Congo, Helicotylenchus multicinctus, Meloidogyne spp., Musa spp. , Radopholus similis, root necrosis.

RESUMEN Kamira M., S. Hauser, P. Van Asten, D. Coyne, and H. L. Talwana. 2013. Nematodos fitoparásitos asociados a los cultivos de banano y plátano en el este y oeste de la República Democrática del Congo.Nematropica 43:216- 225.

Se determinó la incidencia, densidad de población y el daño asociado a nematodos fitoparásitos en 153 cultivos de plátano y banano de agricultores pequeños en Bas Congo (9 – 646 metros sobre el nivel del mar) y South Kivu (1043 – 2005 m.s.n.m.) en la República Democrática del Congo, en 2010. Basados en frecuencia hallada en extracciones de suelo y raíces, Helicotylenchus multicinctus (89%), Meloidogyne spp. (54%) y Radopholus similis (30%) fueron los nematodos más comunes, mientras que Pratylenchus goodeyi (18%) Helicotylenchus dihystera (18%), Rotylenchulus reniformis (14%), y Pratylenchus spp. (6%) se encontraron de manera localizada. La altitud influyó sobre la incidencia y abundancia de las especies: R. similis declinó a elevaciones de más de 1300 m; P. goodeyi declinó a elevaciones de menos de 1200 m; H. multicinctus y Meloidogyne spp. se encontraron en todas las altitudes, pero las densidades fueron más altas a baja altitud; Pratylenchus spp. sólo se encontró en las altitudes bajas; mientras que H. dihystera y R. reniformis se econtraron tanto a altitudes bajas como altas. Helicotylenchus multicinctus se observó en mayor densidad de población que otras especies, especialmente en las altitudes más bajas. El daño a las raíces de banano (porcentaje de raíces necróticas) fue de moderado a alto, sin importar la altitud o el tipo de banano. Con el fin de facilitar el desarrollo de planes de manejo de nematodos fitoparásitos, estos estudios brindan la información necesaria para escoger estrategias adecuadas tales como la selección y distribución de cultivares. Palabras clave: Congo, Helicotylenchus multicinctus, Meloidogyne spp., Musa spp., necrosis radical, 216 Pratylenchus goodeyi, Radopholus similis. Plant parasitic nematodes of banana and plantain in DR Congo:Kamira et al. 217

INTRODUCTION MATERIALS AND METHODS Bananas and plantains (Musa spp.) are important Study site and Sampling crops in the Democratic Republic of Congo (D.R. Congo) (Frison and Sharrock, 1999) produced by The study was conducted in two provinces of smallholder farmers for subsistence and household Bas-Congo (western D.R. Congo) and South Kivu income and are considered the second most important (eastern D.R. Congo) (Fig. 1). Two hundred forty staple crop after cassava (Bakelana et al., 2000). one composite banana root and soil samples were They are grown across a range of environments collected from 153 randomly selected banana fields; and produce fruits year-round, providing a source 111 fields within the 9 territories of Bas Congo: of energy when other crops are often not available. (9), (10), Luozi (13), Plantain (Musa AAB), and cooking bananas (Musa (22), Mbanza Ngungu (10), Mwanda (15), Seke AAB, Musa ABB), East African highland bananas Banza (5), (11) and (16), located (Musa AAA-EA) and dessert banana (Musa AAA, at altitudes ranging between 9 m to 646 m.a.s.l.; and Musa AA) are some of the banana types grown 42 fields within the 2 territories of Walungu (27) and in D.R. Congo (Swennen et al., 1995). At lower Kabare (15) in the province of South Kivu at altitudes altitudes (below 1200 m above sea level, for example, ranging between 1043 m to 2005 m.a.s.l. Among the in Bas Congo western D.R. Congo), plantain and fields sampled in Bas Congo, 85 fields were grown dessert banana are primarily grown, while at mid to a mixture of dessert and plantain bananas, 20 to to high altitudes (1200 to 2000 m.a.s.l.) especially plantain bananas only and 13 to dessert bananas in Eastern Congo, the East African highland banana only. In South Kivu 27 fields were a monoculture types dominate (Mobambo, 2003). The Congo basin of brewing bananas, 5 of cooking bananas while 3 is also viewed as a secondary centre of plantain were an intercrop of plantain, brewing and cooking diversity and thus important from a biodiversity bananas. Where different banana cultivars existed, perspective (Swennen et al., 1995; Daniells et al., each cultivar was sampled separately. The samples 2001). However, the area planted to banana and were collected following a procedure described by plantain, and yield per unit area have declined by 20 Speijer and De Waele (1997), a hole of 20 x 20 x - 60% (Mobambo et al., 2010) even though demand 20 cm was made adjacent to the corm of the banana remains high, raising their market value and often plant and the banana roots and soil were collected out of the reach of poor urban households (Bakelana and placed in a labeled plastic bag. Samples were et al., 2000). taken from 5 to 8 randomly selected individual The reasons for the declining yields include a plants per field, pooled to form a composite root and range of factors, which may depend on site. Pests soil sample, stored in a cool box and transported to and diseases, soil fertility, access to water, etc. are the laboratory within 48 hours of collection. among the key constraints regionally, while only limited information is available for D.R. Congo Root Necrosis Assessment, Nematode Extraction (Dowiya et al., 2009). Due to the long history of And Processing banana and plantain cultivation in the D.R. Congo pests and diseases have co-evolved with the crop Root cortical necrosis (percentage) in banana roots (Mbida et al., 2001), thus adding a more difficult was evaluated on five functional roots randomly dimension to the pest and disease problem (Jones, selected per sample. The selected roots were cut to 2000). Plant parasitic nematodes are among the about 10 cm length, each root piece sliced lengthwise most important constraints to banana and plantain and necrosis in the exposed cortical tissue scored as production, globally reported to cause yield losses percentage, each root piece contributing a maximum of between 30 – 60% (Bridge, 2000; Gowen et score of 20% and for the five root pieces selected al., 2005). Although only limited information is from the sample a total score of 100% (Speijer available, a brief study previously indicated that and De Waele, 1997). The root pieces assessed for damage by plant parasitic nematodes in many of the necrosis were washed with tap water, surface dried, banana and plantain growing regions of DR Congo chopped into ~2 cm segments and thoroughly mixed is severe (Bakelana et al., 2000). Consequently, before removing a 10 g fresh weight sample. The nematode species abundance, distribution, and 10 g sub-sample was covered with water to just underlying mechanisms (e.g. ecological factors and submerge the roots and the sample was macerated farming practices) influencing nematode distribution in a kitchen blender for 10 – 15 seconds. Nematodes and pathogenicity are unknown. This study reports from roots were extracted for 24 – 36 hours using the incidence, nematode population densities and the modified Baermann funnel technique (Coyne associated damage of plant-parasitic nematodes on et al., 2007). Nematodes were also extracted from banana and plantain in DR Congo determined during 100 cm3 of soil using a modified Baermann funnel 2010. technique. The extracted nematode suspensions were decanted into a beaker, allowed to settle for 2 hours 218 NEMATROPICA Vol. 43, No. 2, 2013

Fig. 1: Map of Bas Congo (above) and South Kivu (below) showing the sampling points for the survey. Plant parasitic nematodes of banana and plantain in DR Congo:Kamira et al. 219 and reduced to 25 ml by gently decanting off the in South Kivu infested (Table 1). Pratylenchus excess water. Vermiform nematodes (male, females goodeyi was found exclusively at higher elevations, and juveniles) were identified to species level using i.e. Kabare and Walungu in South Kivu. Overall, the morphometric parameters and individuals counted frequency of nematode species occurrence in roots from 2 ml aliquots drawn from 25 ml suspension with and soil samples ranged from 6% (Pratylenchus a compound microscope with under-stage lighting. spp.) to 89.4% (Helicotylenchus multicinctus) (Table Where no adults existed, nematodes were identified 1). to genus level only. For sedentary endoparasitic Helicotylenchus multicinctus also occurred in Meloidogyne spp. and Rotylenchulus spp. only higher population densities than other nematode vermiform males and second-stage juveniles were species in Bas Congo. Particularly high population recorded. After counting, the nematodes were heat densities of H. multicinctus were recovered in relaxed, fixed in formalin (Coyne et al., 2007) and Lukula, Luozi, Madimba Mbanza Ngungu and sent to the Nematology unit of the South Africa Plant Mwanda. Meanwhile, high Meloidogyne spp. Protection Research Institute Biosystematics division population densities were found in Madimba and for confirmation of our preliminary identification. their lowest density recorded in Songololo (Tables 2 - 3). High R. similis population densities were found Data Analysis in Kasangulu, Luozi, Madimba and Mwanda. Of particular note is the relatively high population density Nematode densities were calculated per 100 g of of R. reniformis in Luozi territory, and Pratylenchus root fresh weight or 100 cm3 of soil. The frequency spp. in Madimba territory. Pratylenchus goodeyi of occurrence for each species identified was was abundant and the predominant nematode calculated for each of the 11 territories. A species species in South Kivu (Tables 2 - 3). Generally, the was considered widespread when it appeared in dominant nematode species varied with altitude but more than 30% of the fields (Adiko, 1988). Mean not with banana type (i.e. brewing, cooking, dessert population densities were calculated for each species or plantain). Radopholus similis was restricted to observed, and the maximum density recorded lower altitudes (<1,200 m), highest densities being for each banana type within a territory. A species recovered at lower altitudes. Pratylenchus goodeyi with a mean density of over 10 individuals per was the dominant species at higher altitudes (>1,200 100 g of roots was considered abundant (Adiko, m). Helicotylenchus multicinctus and Meloidogyne 1988). As the occurrence of banana and plantain spp. were found at all altitudes with higher but not cultivar types generally depended on geographical dominant densities at lower altitudes (Table 4). location, no qualitative or quantitative comparisons Necrosis observed in roots was moderate to high, were possible between Bas Congo and South Kivu for example, the lowest and highest percentage score territories. A Chi-squared test was conducted on on plantain was 18% and 59% at Mbanza Ngungu nematode frequency data within each respective and Kabare, respectively (Table 5), but it was not province to determine significance of differences significantly different among banana types. Root among banana types. Necrosis damage observations, necrosis was positively and significantly correlated expressed as percentages, were normalized using to population densities of H. multicinctus (r = 0.23, angular transformation prior to analysis of variance. P ≤ 0.001), P. goodeyi (r = 0.41, P < 0.002) and R. Correlations were calculated among nematode similis (r = 0.14, P = 0.017). densities, root necrosis damage and altitude. DISCUSSION RESULTS All the seven plant parasitic nematode species Seven plant parasitic nematode species: recovered from banana and plantain in this study are Helicotylenchus dihystera, H. multicinctus, potentially damaging. The abundance and prevalence Meloidogyne spp., Pratylenchus goodeyi, of, and damage associated with H. multicintus on Pratylenchus spp., Radopholus similis and banana in the current study is contrary to previous Rotylenchulus reniformis were recovered from roots studies that indicate R. similis and P. goodeyi as the and soil of bananas and plantains in eastern and most prevalent and damaging nematodes on banana western D.R. Congo. The most frequently occurring in the African great lakes region and only regard H. plant parasitic nematodes were H. multicinctus and multicinctus as a constraint where environmental Meloidogyne spp. Helicotylenchus multicinctus conditions, i.e., altitude, temperature and rainfall are was found in 89.4% of all sampled fields, but the sub-optimal for both the banana crop and R. similis frequency of occurrence was higher in fields at the (Gichure and Ondieki, 1977; Bridge, 1988; Sikora et lower altitudes (98.5% in Bas Congo) than in fields at al., 1988; Sarah, 1989; Kashaija et al., 1994; Elsen higher altitudes (48% in South Kivu). Meloidogyne et al., 2000; Gaidashova et al., 2009). However, it spp. were recorded in 54% of all the sampled fields supports an observation by Speijer and Fogain (1999) with 48% of fields in Bas Congo and 61% of fields that H. multicintus is the most common nematode 220 NEMATROPICA Vol. 43, No. 2, 2013 5.9 1.8 0.0 2.9 0.0 21.4 26.5 26.1 20.0 21.1 22.6 13.5 reniformis Rotylenchulus - - 71.4 41.2 10.9 53.8 32.4 21.4 57.1 26.3 12.9 29.8 similis Radopholus

- - 7.1 0.0 0.0 7.5 0.0 1.8 1.6 6.0 spp. 14.3 34.2 Pratylenchus ------18.2 100.0 100.0 goodeyi Pratylenchus

spp. 53.6 67.6 37.0 51.3 61.8 48.2 57.1 13.2 77.4 47.1 75.0 53.6 Meloidogyne 95.7 92.9 98.4 41.2 55.0 89.4 100.0 100.0 100.0 100.0 100.0 100.0 H. multicinctus 0.0 9.7 35.7 32.4 15.2 16.3 23.5 14.3 15.8 20.6 16.7 18.2 dihystera Helicotylenchus range 9 – 272 Altitude (m.a.s.l.) 512 – 631 107 – 199 196 – 409 563 – 646 564 – 643 370 – 444 178 – 454 144 – 330 1485 – 1705 1043 – 2005 9 5 11 10 13 22 10 15 16 15 27 153 of fields Number sampled Table 1. Frequency of occurrence (percentage) plant parasitic nematodes in roots and soil bananas two provinces Democratic Republic Congo. Table Province/ Territory Bas Congo Kasangulu Lukula Luozi Madimba Mbanza Ngungu Mwanda Seke Banza Songololo Tshela South Kivu Kabare Walungu Overall Plant parasitic nematodes of banana and plantain in DR Congo:Kamira et al. 221 ------25 (3) 70 (5) 35 (2) 12 (1) 49 (5) 32 (5) 17 (2) 20 (1) 38 (2) 391 (33) 244 (22) 190 (15) reniformis reniformis 3,120 (205) Rotylenchulus Rotylenchulus 15,517 (1,710) 41,125 (2,100) 13,934 (1,281) - - - - 16 (1) 17 (6) 75 (9) 131 (9) 174 (42) 348 (52) 106 (18) 105 (16) 120 (10) 1,875 (559) 4,077 (346) 8,195 (1,358) 11,482 (1,041) 11,482 44,250 (2,609) 43,625 (5,247) 36,875 (3,566) 14,625 (1,044) 69,000 (10,866) Radopholus similis Radopholus similis spp. spp. ------6 (1) 25 (2) 12 (1) 117 (5) 117 163 (7) 250 (36) 459 (80) 24,375 (1,741) 43,000 (1,075) 21,216 (1,800) Pratylenchus Pratylenchus ------goodeyi goodeyi 300 (95) 866 (244) Pratylenchus Pratylenchus 149,700 (63,246) 270,380 (26,218) spp. spp. soil of bananas in two provinces Democratic Republic Congo. 3 44 (15) 34 (12) 129 (7) 600 (46) 124 (39) 172 (30) 128 (28) 180 (20) 199 (41) 365 (121) 781 (141) 401 (102) 1,250 (411) 6,875 (625) 4,000 (765) 4,950 (966) 4,400 (636) 3,750 (414) 4,750 (1,429) 11,814 (2,845) 11,814 52,500 (3,971) 258,250 (6,506) Meloidogyne Meloidogyne 121 (17) 120 (16) 575 (219) 526 (184) 969 (199) 942 (223) 884 (252) 886 (129) 644 (121) 4,410 (660) 2,426 (234) 2,686 (456) 8,880 (1366) 25,832 (4,966) 81,328 (8,220) H. multicinctus H. multicinctus 59,813 (20,873) 76,875 (19,843) 119,643 (35,587) 119,643 156,750 (28,992) 170,363 (21,308) 153,365 (19,355) 263,500 (56,484) - - 11 (1) 11 77 (5) 48 (8) 85 (9) 47 (4) 33 (2) 54 (7) 29 (4) 12 (1) 13 (2) 20 (2) 93 (13) 207 (19) 381 (26) 207 (21) 210 (15) 820 (81) 120 (10) dihystera dihystera 572 (108) 1,009 (169) Helicotylenchus Helicotylenchus Table 2. Maximum (and mean) nematode population densities per 100 g root fresh weight of bananas in two provinces Democratic Republic Congo. Table Territory Province/ Bas Congo Kasangulu Lukula Luozi Madimba Mbanza Ngungu Mwanda Seke Banza Songololo Tshela South Kivu Kabare Walungu 3. Maximum (and mean) nematode population densities in 100 cm Table Territory Province/ Bas Congo Kasangulu Lukula Luozi Madimba Mbanza Ngungu Mwanda Seke Banza Songololo Tshela South Kivu Kabare Walungu 222 NEMATROPICA Vol. 43, No. 2, 2013 ------7 7 8 2 4 1 73 324 424 0 4 0 0 7 1,694 4,381 2,196 1,724 1,087 reniformis reniformis Rotylenchulus Rotylenchulus ------

625 217 344 499 500 500 838 510 462 2,324 4,909 2,109 7,016 1,285 7,804 9,953 1,695 13,929 similis Radopholus 0 0 0 0 5,262 1,346

Radopholus similis ------

63 809 spp. 3,482 1,792 2,692 Pratylenchus 0 0 0 52,611 45,775 49,260 goodeyi ------

Pratylenchus 2,730 1,472 58,955 77,865 68,240 22,529 29,152 goodeyi 132,750 Pratylenchus 0 0 0 586 862 288 spp.

30 60 114 339 807 531 292 831 482 458 710 972 724 125 515 785 424 656 spp. 3,307 2,979 2,435 4,821 2,281 1,092 1,303 10,370 Pratylenchus Meloidogyne spp. - - -

275 883 150 766 2,112 3,300 5,915 3,315 6,465 0 17,374 29,306 26,722 22,937 37,400 55,394 13,634 24,372 41,171 31,073 18,864 51,419 24,499 10,381 430 488 749 3,568 1,773 multicinctus Helicotylenchus Meloidogyne Nematode population density (per 100 g root fresh weight) ------3 8 2 63 59 40 26 42 31 68 25 28 94 16 10 154 267 dihystera Helicotylenchus 842 246 1,132 1,579 20,830 26,509 multicinctus Helicotylenchus

42.0 + 8.1 38.9 + 7.6 31.2 + 6.4 20.1 + 4.4 26.6 + 7.6 23.8 + 6.2 19.8 + 7.1 21.1 + 5.5 26.4 + 8.1 37.1 + 7.1 34.6 + 6.2 38.1 + 5.3 17.8 + 7.1 34.0 + 5.7 35.5 + 6.8 22.2 + 5.3 47.5 + 6.4 39.9 + 5.3 36.4 + 9.6 24.6 + 9.6 21.3 + 12.3 30.0 + 10.7 53.5 + 15.1 23.0 + 15.1 59.0 + 15.1 35.8 + 10.7 [Mean + s.e] Root necrosis 2 1 39 14 53 862 dihystera

Helicotylenchus Dessert Dessert Dessert Plantain Plantain Plantain Brewing Brewing Cooking Cooking Banana type Bas Congo Table 4. Influence of altitude on nematode species occurrence and population density banana in Democratic Republic Congo. Table Altitude ranges (m.a.s.l.) > 2000 1601 - 2000 1201 - 1600 801 - 1200 401 - 800 0 - 400 Republic of Democratic in two provinces sampled cultivars of banana (per 100 g root fresh weight) densities population (root necrosis) and nematode damage 5. Nematode Table of Congo. Province/ Territory Kasangulu Lukula Luozi Madimba Mbanza Ngungu Mwanda Seke Banza Songololo Tshela Kasangulu Lukula Luozi Madimba Mbanza Ngungu Mwanda Seke Banza Songololo Tshela South Kivu Kabare Walungu Plant parasitic nematodes of banana and plantain in DR Congo:Kamira et al. 223 species in roots of Musa or Ensete in Africa. The (Speijer and Fogain, 1999; Sarah, 1989; Bridge et magnitude of necrosis associated to H. multicintus al., 1995; Coyne, 2009). This nematode is highly observed in the current study may be a result of pathogenic and a potentially serious threat to banana either the high populations recovered or a shift in the and plantain production (Bridge et al., 1997; Speijer “known” reproductive fitness and pathogenicity of et al., 2001; Brentu et al., 2004; Coyne and Dubois, H. multicinctus, since histopathological observations 2011). of H. multicinctus-infected roots showed superficial The observed high root necrosis together with the lesions that do not extend into the stele of banana low nematode densities recovered in soil samples roots (Zuckerman and Strich-Harari, 1963). It indicates that farmers are suffering substantial losses is apparent that this damage could translate into due to nematodes, which are undoubtedly being heavy yield losses, because elsewhere where H. spread using contaminated suckers for planting multicinctus was found as the dominant species on material, a practice common in many smallholder banana, it caused significant crop losses, for example banana growing areas (Tenkouano et al., 2006). This on plantain in western Africa (Badra and Caveness, also shows the relatively high susceptibility and 1983; Sarah, 1985). sensitivity to plant parasitic nematodes of the banana The prevalence and abundance of the other cultivars grown by the farmers in D.R. Congo. nematode species; namely, Meloidogyne spp., R. In conclusion, the results of this survey revealed similis and P. goodeyi was typical of what is reported the significant prevalence of Helicotylenchus in the region. Meloidogyne spp. was encountered in multicinctus, Meloidogyne spp., Pratylenchus 54% of the fields sampled distributed throughout the goodeyi and Radopholus similis, the four most two provinces, with higher populations recovered important nematodes on banana. These species are at lower to mid altitudes, for example, in Madimba. known to cause severe damage and yield losses Radopholus similis was encountered in 30% of the throughout most banana growing areas in the fields with its highest density recovered at mid- tropical and subtropical regions of the world and altitudes, in Kasangulu. This distribution of R. their distribution is very closely linked to altitude similis reflects its tropical (thermophilic) nature, and temperature. However, the current study is not and supports other reports, such as Bridge (1988) exhaustive but provides baseline information upon in Tanzania and Kashaija et al. (1994) in Uganda. which further studies can extend the investigations, Pratylenchus goodeyi was abundant at elevations for example, partitioning and quantifying damage above 1200 m.a.s.l. in South Kivu, confirming caused to banana by individual nematode species similar observations by Sarah (1989), Kashaija et al. and, prioritizing and guiding development of (1994), Price and Bridge (1995), Bridge et al. (1997), management strategies. Fogain (1998), Price (2000), and Gaidashova et al. (2009). The high P. goodeyi population densities ACKNOWLEDGEMENT and frequencies in the territories of Walungu and Kabare are most likely enhanced by the lengthy We gratefully acknowledge the financial support banana monoculture (Speijer and Fogain 1999), but from the European Union through the International their distribution and density is related to altitude and Institute of Tropical Agriculture (IITA) to the are similar to reports elsewhere (Elsen et al., 2000; Agriculture and Forestry Research (REAFOR) Fogain, 2001; Talwana et al., 2003; Gaidashova et project in D.R. Congo of which this study was part. al., 2009). Helicotylenchus dihystera and R. reniformis LITERATURE CITED appeared to have a scattered distribution at both low and high altitudes. Pratylenchus spp., however, had Adiko, A. 1988. Plant parasitic nematodes associated a clustered but erratic distribution at lower altitudes with plantain, Musa paradisiaca (AAB) in Ivory (9 m to 646 m.a.s.l.) within Bas Congo province, Coast. 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Received: Accepted for publication: 2/XI/2012 29/V/2013 Recibido: Aceptado para publicación: