Fundam. appl. NemaLOI., 1995, 18 (5), 435-444

Ecology and pathogenicity of the (Nemata) from the sahelian zone of West Mrica. 6. pararobustus (Schuurmans Stekhoven & Teunissen, 1938) Sher, 1963 and comparison with Luc, 1958

Pierre BAuJARD* and Bernard MARTINY ORSTOM, Laboratoire de Nématologie, B.P. 1386, Dakar, Sénégal. Accepted for publication 10 August 1994.

Summary- The geographical distribution, population dynamics and vertical distribution in the soil were srudied for the Hoplolaimus pararobuslUS. The factors influencing the multiplication and survival rates of H. pararobus/Us and H. seinhorSli were srudied in the laboratory. Both species appeared adapted to the ecological conditions of the semi-arid tropics of West Africa. The absence of H. sein1wmi from these ecological zones remains unexplained; the possibiliry of the loss of reproduction capaciry after anhydrobiosis is suggested. A the inoculum levels tested, these IWO species did not have any pathogenic effects on the plants with the exception of H. pararobuslUs which induced a reduction of root fresh weight for cowpea.

Résumé - Éco/ogW et nocuité des Hoplolaimidae (Nemata) de la zone sahélienne de l'Afrique de l'Ouest. 6. Hoplolai­ mus pararobustus (Schuurmans Stekhoven & Teunissen, 1938) Sher, 1963 et comparaison avec Hoplolaimus sein­ horsti Luc, 1958 - La répartition géographique, la dynamique et la répartition verticale des populations sont érudiées pour le nématode Hoplolaimus pararobusluS. Les facteurs influençant les taux de multiplication et de survie sont éudiés au laboratoire pour H. pararobuslUs et H. sein1wrsli. Ces travaux montrent que ces deux espèces sont parfaitement adaptées aux conditions climatiques régnant dans la zone sahélienne ouest africaine. L'absence d'H. seinhorsli de ces biotopes reste donc inexpliquée; l'évenrualité d'une pene de capacité reproductrice après l'entrée en anhydrobiose est évoquée. Aux taux d'inoculum testés, ces deux espèces n'ont pas d'effets sur la croissance des plantes à l'exception d'H. pararobuS!us qui induit une diminution du poids frais des racines pour le niébé. Key-words : Hoplolaimus pararobus/Us, Hoplolaimus seinhorsli, nematode, West Africa, geographical distribution, population dy­ namics, venical distribution, soil temperarure, soil moisrure, host plant, multiplication rate, anhydrobiosis pathogeniciry.

Species of the genus Hoplolaimus are suspected or Material and methods known to be parasites of numerous tropical crops in the world (Luc el al., 1990). In Africa, rwo species of this Studies on geographical distribution, field population genus, Hoplolaimus pararobusLUs (Schuurmans Stekho­ dynamics and vertical distribution have been conducted ven & Teunissen, 1938) Sher, 1963 and H. seinhorsli as previously described (Baujard & Martiny, 1995 c). Luc, 1958, are widely distributed (Siddiqi, 1974; Van Unless otherwise stated, nematode extractions, nema­ den Berg, 1976). H. pararobusluS is identified in the soils tode cultures and technics, host plants and cultivars of semi-arid tropics of West Africa in Mauritania (Bau­ (peanut [Arachis hypogea L. cv. 55 437], millet [Pennise­ jard & Martiny, 1995 a), Senegal (Baujard & Martiny, lum lyphoides Rich. cv. Souna III], sorghtun [Sorghum 1995 b), Mali (Baujard & Martiny, 1994), Burkina Faso vulgare L. cv. 51 69], cowpea [Vigna unguiculata (L.) (Sharma, 1989), Niger (Sharma el al., 1988, 1990, Walp.] cv. N 58 57 used for laboratory studies are those 1992), where H. seinhorsli has never been recorded. described by Baujard (1995).

This sixth paper on the ecology and pathogenicity of ORlGIN OF AND LABORATORY STOCK the Hoplolaimidae (Baujard & Martiny, 1995 c, d, e,1, CULTURE CONDITIONS g), presents the results offield and laboratory studies on H. pararobuslUs: nematodes originated from samples H. pararobusluS and H. seinhorSli. taken at Dombe, km 5 Diourbel to Thies road, Senegal,

* Present address: Muséum Nalional d'Hisloire Nalurelle, Laboraloire de Biologie Parasitaire, ProlisLOlogie, Helminthologie, 61, rue Buffon, 75005 Paris, Frame. fSSN 1164-5571/95/05 S 4.00/ © Gaulhier- Villars - ORSTOM 435 P. Baujard & B. Marllny

in a field previously cropped with millet in June 1981 H. seinhorsti: tubes were inoculated with 152 ± 12 during the dry season. The nematodes were extracted in nematodes (mixture of aU stages) originating from soil August 1983 and reared on millet in the laborarory. and sorghum roots from the previous experiment on soil From August 1983 to May 1989, the nematodes were temperature, planted with sorghum, and maintained at reared without control of soil temperature or moisture; four constant soil moisture levels (S, 7, 9 or Il %) at from May 1989 until May 1992, the soil temperature 32 oC constant soil temperature for 60 days in a green­ and moisture were kept constant respectively at 34 oC house; the four treatments were replicated ten rimes in a and 10 %. completely randomized design. H. seinhorsli: nematodes originated from a yam tuber from Maninique, French West Indies, sent to us by Dr. HOST PLANTS AND TEST FOR ANHYDROBIOTIC SlJR­ P. Cadet in May 1989. Nematodes were extracted in a VIVAL mist chamber and reinoculated on Zea mays cv. HVB1, H. pararobuslUs: tubes were inoculated with 143 ± 14 kenaf (Hibiscus cannabinus L.), sorghum and millet, and nematodes (mixture of aU stages) originating from a maintained at 32 oC constant soil temperature and 10 % 100-day-old stock culture on millet, planted with one of constant soil moisture. the four host plants (cowpea, millet, peanut or sor­ SaIL TEMPERATURE ghum), and maintained at 34 oC constant soil temper­ H. pararobustus : in a fust experiment, tubes were in­ ature and 10 % constant soil moisture in a greenhouse. oculated with 25 hand-picked nemarodes (mixture of ail The four treatments were replicated 20 times in a com­ stages) originating from 40-day-old stock cultures, pletely randomized design. After 60 days, nemarodes planted with millet, and maintained at four constant soil were extracted from ten replications to obtain final temperature levels (30, 32, 34 or 36 oC) with seven population counts. At this rime, watering was stopped replications for each temperature level, at 7 % constant for the ten remaining repJications of each treatment. soil moisture, for 75 days in a growth chamber with These tubes were kept at 34 oC constant soil temper­ artificial lighting (l6-h photoperiod). In a second ature and weighed daily to foUow the evolution of soil experiment, tubes were inoculated with 67 ± 5 nema­ desiccation. Sixry days later, nematodes were extracted todes (mixture of aU stages) originating from 100-day­ by elutriation. old stock cultures, planted with sorghum, and main­ H. seinhorsti: tubes were inoculated with 103 ± 7 ne­ tained at four constant soil temperature levels (30, 32, matodes (mixture of ail stages) originating from sor­ 34 or 36 oC) with seven replications for each temper­ ghum roots of the previous experiment on soil moisture, ature level, at 10 % constant soil moisture, for 60 days in planted with one of the four host plants (cowpea, millet, a growth chamber. peanut or sorghum), and maintained at 32 oC constant H. seinhorsti : tubes were inoculated with 91 ± 7 nema­ soil temperature and 10 % constant soil moisture in a rodes (mixture of all stages) originating from stock cul­ greenhouse. The experiment was conducted as de­ tures on kenaf, miUet and sorghum, planted with sor­ scribed for H. pararobustus. ghum, and maintained at four constant soil temperature levels (30, 32, 34 or 36 oC) with seven replications for CULTURE DURATION each temperature level, at 10% constant soil moisture, for 60 days in a growth chamber. This experiment was conducted orny with H. pararo­ bustus: 100 hand-picked nemarodes (mixture of aU SaIL MOlSTURE stages) originating from a 60-day-old stock culture on H. pararobustus : in a first experiment, tubes were in­ millet at constant soil temperature (34 OC) and soil oculated with 110 hand-picked nematodes (mixture of moisture (l0 %) were inoculated on millet at constant ail stages) originating from the first soil temperature soil temperature (34 oC) and soil moisture (10%) for experiment, planted with miUet, and maintained at four three different durations (60,75, 100 days) with seven constant soil moisture levels (5, 7, 9 or Il %) at 34 oC replications per treatrnent in a growth chamber. At the constant soil temperature for 75 days in a greenhouse end of each culture period, nematodes were extracted, with naturallighting; the four treatrnents were replicated counted and reinoculated in the same conditions (in­ ten times in a completely randomized design. In a se­ oculum level, host-plant, soil temperature and moisture, cond experiment, tubes were inoculated with 94 ± 10 durations) three times. In addition, nematodes originat­ nematodes (mixture of aU stages) originating from a ing from the (, 75 days ,} treatment were reinoculated in 75-day-old srock culture on millet, planted with sor­ the same experimental conditions for a culture dmation ghum, and maintained at four constant soil moisture of 60 days; those originating from the (' 100 days I} treat­ levels (5, 7, 9 or Il %) at 34 oC constant soil temper­ ment were reinoculated in the same experimental condi­ ature for 60 days in a greenhouse; the four treatments tions for a culture dmation of 60 and 75 days in order ro were replicated ten times in a completely randomized evaluate the effect of the previous culture duration on design. the multiplication rate.

436 Fundam. appl. NemaLOl. Ecology ofHoplolaimidae. 6

PATHOGENICITY TO COWPEA, MILLET, PEANUT AND cultures on millet were inoculated onto each host at two SORGHUM inoculum levels : onto peanut at 500 ± 50 or 1000 ± 100, onto rniUet at 220 ± 20 or 440 ± 40 in a first experiment H. pararobuslus : separate experiments were conduct­ and at 500 ± 20 or 1000 ±40 in a second experiment, ed with each crop species at 34 oC soil temperature and onto sorghum at 500 ± 40 or 1000 ± 80, onto cowpea at 10 % soil moisture in the greenhouse. Nematodes (mix­ 350 ± 40 or 700 ± 80 nematodes per tube. Nematode ture of aU stages) originating from 100-day-old stock effects were compared to control plants without nema- 400 200 lA , 0 ,1::1,"" 800 "El ",,--f,m-if'RRR ~ ~ 600 B ;------II----~

400

200

0 200 ~OO~OOOOO~_ OOOO~OOOO~%c+OOOO 0 le 0 ..J 0 (j) LI.. o 0 1000 M ::E c 800 a: LU a. 600 a: LU al 400 ::E :::l 200 Z LU C 0 0 t-

800

600

400

200

0 1984 1985 1986 1987 1988 Fig. 1. Popuialion dynamics ofHoplolaimus pararobusrus according La lhe cullUral praclices. A .' Peanul morUJcullUre; B : Peanul-millel roullion wiLhoul nernaLicidal lreaLmenL; C.' Peanw-millel rOlaLion wiLh nemaLicidal lrealmenl; D.' Mi/lel monoculLure; E.' Sorghum monoculLure; F.' Cowpea monoculLure; G.' Pennanenlfallow (Ha(ched areas : rainy seasons; squares: soil population; c1iamonds : root population) .

Vol. 18, n° 5 - 1995 437 P. Bat/jard & B. Maniny

RAlNY SEASON DRY SEASON 1984 fJ 1985 t:il 1986 Cl 1987 1 • 1984-1985 ~ 1985-1986 100 • 100 ~ 1986-1987 Cl 1987-1988 r

,

r .....0:: 80 , ~ 80 '.'t> ~ .E , " / w ,~ ~ , , 1- :.- « 60 -:: 60 cr: ~ z " w 0 " 1-« ::. ~ a: « " , () 40 ...J 40 ::::; « c.. ~ , ~ :> ...J a: ,- :l :l ~ 20 fi) 20 " " " , ~] ~ n n ~ 0 --9 - 0 L. l A BCDEFa G A BCDEF G CULTURAL PRACTICE CULTURAL PRACTICE

Fig. 2. Multiplication and sU7"vivai rates ofHoplolaimus pararobusrus according ta the cultural praclices and year ofobservations (See Fig. 1 for the legend).

todes. The three treatments were replicated ten times in bania pachycarpa De., Andropogon guayanus Hack., Ica­ a completely randomized design. After 40 days, nema­ cina senegalensis A. Juss., PilioSligma reticulaturn (De.) todes were extracted from soil and roots to determine Hochst., Acacia albida Del., Acacia SenegaL (L.) Willd., the multiplication rate, and the fresh weight of roots and Acacia niLotica (L.) Willd., Acacia IOrtilis ssp. raddiana fresh and dry weights of shoots were measured. Savi., Prosopis juLiflora De.]. Ir has never been found in H. seinhorsti: a separate experiment was conducted the rhizosphere of Euphorbia balsamijera Ait., a plant only with sorghum and cowpea at 32 oC soil temper­ commonly used as a green fence in this area. ature and la % soil moisture in the greenhouse. Nema­ FIELD STUDIES ON H. PA.RAROBUSTUS todes (mixture ofail stages) originating from roots ofthe soil moisture experiment were inoculated at two inocu­ Population dynamics lum levels (325 ± 23 or 650 ± 46 nematodes per tube) The soil population increased from the beginning to and other nematodes (mixture of ail stages) originating the end of the rainy season slowly in the rhizosphere of From soil and roots of a 49-day-old stock culture on peanut, sorghum, cowpea and fallow plants and strongly sorghum were inoculated at two inoculum levels onto in the rhizosphere ofmillet. Nematodes invaded the root cowpea (500 ± 35 or 1 000 ± 70 nematodes per tube). system only on millet where root populations never ex­ Nematode effects were compared to control plants with­ ceeded 200 nematodes per root system. At the end or out nematodes. The three treatments were replicated ten just afrer the end of the rainy season, the population times in a completely randomized design. After 40 days, densi!)' decreased rapidly and remained constant nematodes were extracted From soil to determine the throughout the dry season (Fig. 1). Multiplication rates multiplication rate, and the fresh weight of roots and the during the rainy season varied From a to 80 and survival fresh and dry weights of shoots were measured. rates during the dry season from a to 100 % according to the crop and to the year. For millet, sorghum, cowpea, Results and fallow, an alternation of rainy seasons with high and low multiplication rates occured (Fig. 2). GEOGRAPHICAL DISTRlBUTION Although millet appeared to be a good host for the AND HOSTS OF H. PARAROBUSTUS IN SENEGAL nematode, no multiplication was recorded with this H. pararobustus is a ubiquitous species in the soils of plant in 1985 and 1987 on the plots under the peanut­ Senegal where it occurred during the dry and the rainy millet rotation (Fig. 1). 3 seasons in low numbers (0-2 000 nematodes per dm ) associated with pluvial crops [Arachis hypogea L., Penni­ VerticaL distribution in the saiL setum typhoides Rich., Sorghum vuLgare L., Vzgna un­ The nematode ...vas erratically distributed in the soil, guiculata (L.) Walp., Hibiscus sabdariffa L.] and several From the surface down to a depth of80 cm; it was mosùy annuai and wild plants [Cenchrus biflorus Roxb., Ses- predominant in the upper layers of the soil (Fig. 3).

438 FU/ulam. appl. Nematal. Ecology ofHoplolaimidae. 6

5 5

10 10

15 15

20 20 Ê Ê ~ 25 ~ 30 J:...... J: Cl. 30 Cl. 40 W W 0 0 35 50

40 60

45 70

50 80 0 D

5 5

10 10

15 15

20 20 Ê Ê ~ 25 ~ 30 ...J: ...J: Cl. 30 Cl. 40 W w 0 0 35 40 :: ] 45 70

50 80 G G

5 5

10 10

15 15

20 20 Ê Ê ~ 25 ~ ...J: ...J: Cl. 30 Cl. 40 W W 0 0 35 50

40 60

45 70

50 80 E F 0 10 20 30 40 50 60 70 80 90 100 0 10 20 30 40 50 60 70 80 90 100 PERCENTAGE OF TOTAL POPULATION PERCENTAGE OF TOTAL POPULATION

Fig. 3. Vertical dismlnmon ofHoplolairnus pararobusrus according la lhe cullural praclices and lhe season ofobservation. (See Fig. 1 for the legend; left column : dry season; right column : rainy season).

Vol. 18, n° 5 - 1995 439 P. Baujard & B. Marliny

30 A LABORATORY STUDIES Observations on laboratory stock cultures 25 H. pararobustus : during me flfSt period of stock cul­ >- '"~ cr: 20 ture, multiplication rates varied from 0.12 to 2.77 z o (x =2.22 ± 1.76, n =29) and during me second period, ~ 15 ü from 1.01 to 41.6 ex = 9.61 ± 10.09, n = 36). Sex ratio ~ >= varied from 36 to 150 % (x =87 ± 28, n = 12). ~ 10 H. seinhorsti: me nematode reproduced on , ~illet, sorghum and kenaf wim multiplication rates of 3-15,20-31.9,9.84-74.22, and 46.65 respectively. No

30 32 34 36 males occurred in me stock cultures. SaIL TEMPERATURE (·C)

30 Factors affecting the muliiplication rate B Soil temperature affected significantly me multiplica­ 25 tion rate of bom me species in me same way, higher >- multiplication occurring at 32-34 oC (Figs 4, 7). Soil '"~ 20 cr: z moisture affected significantly me multiplication rate o ~ 15 only of H. pararobustus on millet and sorghum (Figs 4, ü ::; 7). The use of millet or sorghum as host plant for H. ~ ~ la pararobustus in mese experiments did not modify mese trends. The lower multiplication rates observed for H. pararobustus under sorghum in comparison wim millet can be related ta me effect of host plant ancIJor of me

30 32 34 effect of me duration of me experiment (75 days wim .,SOll TEMPERATURE (-C) millet vs 60 days wim sorghum). Host plants had a sig­ 30 nificant effect on me multiplication rate of bom species. c Ali me plants allowed me reproduction of H. seinhorsti, 25 whereas only millet and sorghum allowed it for H. para­ ta >- robustus. Millet appeared be me best host for H. para­ '"~ 20 cr: robustus and cowpea for H. seinhorsti (Figs 5, 6). Mul- z o ;:: ~ 16 ü ~ ;:: g 10 30

25

7 9 11 SOll MOISTUAE (-I.) W 1- 30 « 20 o a: z 26 o i= 15 >- « '"~ 20 ü cr: z :J o Cl. ~ 16 i= ü :5 10 ~ ~ ;:: :5 la '" 5

c­ 7 9 " o SOll MOISTUAE (-/.) MILLET SORGHUM COWPEA Fig. 4. Effew o/soillemperalUre (A. B), and soil moislUre (G, D) HOST PLANT on tfle multiplication raie o/ on millet Fig. 5. Effects o/hast plants on tfle multiplication rate o/Hoplolai­ (A-G) and sorghum (B, D). (In each experiment, data followed mus pararobustus. (In each experiment, data followed by the by the same lerrer are not significantly different at P ~ 0.05). same lerrer are not signiflcantly different at P ~ 0.05).

440 Fundarn. appl. NernalOl. Ecology ofHoplolaimidae. 6

tiplication rates recorded for H. seinhorsti under sor­ HEMATOOE COI'OTlOH ~1 ANotYOROOlOnc • ttvOAOlltOnc ghum in this experiment were four to five times higher 1200 A than in the experiments on soil temperature and soil ./ moisture (Fig. 6). This difference could really be related 1000 only to the origin of the inoculum : soil and root popu­ 800 lations in the experimenrs on soil temperature and mois­ ture 'Vs only root populations in the experimenr on host 600 plant. The multiplication rate of H. pararobustus increased 400 with an increase in the duration of the culture; the dura­ tion of the previous culture did not affect the multiplica­ 200 tion rate of the nematode (Fig. 6).

Root population ievels with millet and sorghum were PEANUT MILLET SORGHU~OWPEA constant in these experimenrs; they were not affected by HOST PLANT environmenral factors and differed according to the spe­ 100 B cies: 20-68.5 % for H. pararobustus 'Vs 66.1-85.8 % for ------H. seinhorsti; with poor host plants, these levels were 80 below 10 % (Table 1). ~ w ~ 80 < Ability to enter anhydrobiosis cr: ;i > Cessation of watering induced a decrease of soil mois­ '> 40 cr: mre down to 0.2 % in 5 days for H. pararobustus and ::> down to 0.5 % in 15 days for H. seinhorsti. Nematodes '" 20 survived soil desiccation over a 60-day period according to the size of the population before watering was stop­ ped for the four host plants for H. pararobustus (Fig. 6);

Table 1. Percenlages of lhe populaLion of Hoplo1aimus pararo­ 30 c busnls and Hoplolaimus seinhorsti in lhe rOOls al lhe end of lhe 25 dijferenl expen'menls on muliiplualion l'ale. (ND = nor deter­ w ~ mined). ~ 20 /"

Z Q / < 1/5 <.> Experiment Treaonent Root population as a%of ii' ~ 10 tota! tube population ::> ~

H. pararobustus H. seznilrmri

600AYS 75DAYS 100DAYS MilIee Sorghum CUL TURE LENGTH

30 Sail temperature 30'C 68.5 31.8 822 D 32 'C 63.3 74.4 52.2 25 34°C 68.4 31.3 78.3 w ~ 36'C 54.9 45.4 66.1 ~ 20 z Sail moisrure 5% 280 20.0 781 Q 15 7% 41.4 34.5 73.2 ~ Host plants peanut 1.5 9.1 millet 38.5 69.1 sorghum 44.2 80.1 cowpea 60/60 75/60 100/60 75175 100/75 2.8 85.8 PREVIOUSJACTUAL CULTURE LENGIH (numb61 01 da ys) Culture length 60/60 42.1 ND 75n5 41.4 ND Fig. 6. EjfeCls ofsail desiccalion on salt popu/alion (.,1) and surviv­ 100/100 35.6 l-m al l'ale (8) and ejfecl ofculiure lenglh on muliiplicaLion rate (C, D) ofHoplo1aimus pararobustus.

Vol. 18, n° 5 - 1995 441 P. Baujard & B. Marliny

30 Table 2. Percenlages of the populalion ofHoplolaimus pararo­ A busrus and Hoplolaimus seinhorsti in lhe raols al lhe end oflhe 25 dif.[erenl experirnenls on palhogenicùy. (ND = not determined). ;~ ~ 20 '0: l~ Experiment Treatment Root population as a % of ,~ 15 total rube poplÙation u ii';:: ~ 10 ~ H. pararobustus H. seinhorsli

Peanut * 500 2.9 ND

* 1000 2.3 ND 32 3. 38 ~OIL TEMPERA TURE CC) Millet * 220 36.0 ND ---_.- * 440 29.1 ND 30 * 500 61.3 ND * 1000 58.5 ND .. w >- Sorghwn * 500 32.3 ND ~ 20 or * 1000 30.8 ND z 0 * 325 ND 81.4 ;:: ,. ~ u * 650 ND 85.6 ii' ;:: Cowpea * 350 56.4 ND :5 10 :l' * 700 61.4 ND * 500 ND 91.3 * 1 000 ND 90.4

7 • " InocuJum per tube. SOll MOISTURE (el.) " 150 c for H. seinhorsti, the number of nematades surviving soil 125

w desiccation was constant for millet, sorghum and cow­ >- < 100 pea (Fig. 7). Survival rates varied according ta the host or plant and to the nematade species : 13-36 % for H. para­ oZ ~ 75 robuSlus vs 9-22 % for H. seinhomi (Figs 6, 8). u ii' Palhogenicity ~ 60 :l' Neither species had any significant effect on growth of plants at the inoculum levels tested on peanut, millet, 25 sorghum or cowpea (Tables 3,4). Gnly H. pararobuslUs induced a reduction of cowpea fresh root weight at the PEANUT MILLET SORGHUM COWPEA rate of 1000 nematades per plant. Root population HOST PLANT /EUATODE cotClTION levels varied as in the previous experiments (Table 2). . 12000 .Q A~06lOnc • HVDAOCIIOTlc ", Discussion 10000

Laboratory experiments conducted with H. pararo­ -g SOOO bUSlUS conflTmed the field observations on population . 1 eooo dynamics and supported the currenùy known distribu­ z tion ofthis species from wet to semi-arid tropics of West ~ ; 4000 Africa. Cereals are good hosts for this species which is a. o able ta support high soil temperature levels and ta enter a. < 2000 anhydrobiosis. Reproduction is dependenr directly and z ;;: indirecùy (duration of activity in the soil) on the soil moisture. PEANUT MILLET SORGHUM COWPEA HOST PLANT These experimenrs showed that H. seinhorsti exhib­ Fig. 7. Efjects of sail temperalure (A), sail moiswre (B) and hast ited the same behaviour as H. pararobuslUs in relation ta planls (C) on lhe mulliplicalion rate and efjeets of soil desiccalion temperature and soil moisture. It appeared better adapt­ on sail populalion (D) ofHoplolaimus seinhorsti. (ln each exper­ ed ta the ecological conditions of the sahelian zone of iment, data followed by the same letter are not significantly West Africa since it is more polyphagous and its multi- different at P ,,:: 0.05).

442 Fundam. appl. NemalOl. Ecowgy of Hoplolaimidae. 6

Table 4. Multiplication rate and effects ofHoplolaimus seinhor­ 100 ,/ sri on peanut and millet. (Numbers followed by the same letter are not significantly different at P < 0.05).

80 V Plant Inoculum Multiplication Fresh weight Dry weight g rate (g) (g) w 60 ./' 1-« Roots Shoots Shoots OC ...J « -. Peanur 0 2.71 a 5.07 a 0.80 a > ;; 40 V 325 15.12 2.37 a 5.20 a 0.77 a OC ::> 650 13.99 2.21 a 4.89 a 0.76 a en Cowpea 0 3.76 a 6.32 a 1.28 a 20 V 500 2.29 4.29 a 6.54 a 1.21 a 1000 1.88 4.00 a 6.82 a 1.20 a <

o ./ ~ "- .- PEANUT MILLET SORGHUM COWPEA HOST PLANT survival rate ofH. seinhorsli is 50 % that ofH. pararobus­ Fig. 8. Effects ofhost plants and sail desiccation on survival rate of lUS and independent of the size of the population before­ Hoplolaimus seinhorsti. soil drying. Survival abilicy to the long dry season (up to 9 months) occurring in semi-arid tropics of West Africa remains unknown since the experiment was conducted Table 3. Multiplication rate and effeclS ofHoploJaimus pararo- only for 60 days and without any evaluation of the re­ bustus on peanut and millet. (Numbers followed by the same production capacicy of the nematodes after anhydrobio­ letter are not significantly different at P ~ 0.05). SIS. H. pararobus/Us appeared pathogenic only on cowpea, Plant Inoculum Multiplication Fresh weight Dry weight inducing a reduction of fresh weight of roots. Little is rate (g) (g) known about pathogenicicy of these two species. Good­ ey (1957) and Whitehead (1959) respectively reponed Roots Shoots Shoots the presence of H. pararobuslus under the names H. proporicus and H. auguslulaws in the roots of oil palm and banana exhibiting rotting symptoms. No laboratory Peanut 0 2.34 a 7.95 a 1.37 a data are available on the pathogenicicy of this species. 500 067 2.03 a 6.96a 1.15 a 1000 0.48 2.03 a 7.71 a 1.32 a Bridge (1973) showed that H. seinhorsLi parasitized cow­ pea in Nigeria, whose roots exhibited necrosis in the Millet 0 4.19 a 4.10 a 0.64 a field. The nematode was also found associated with 220 0.70 2.99 a 4.41 a 0.72 a poor growth of pigeon pea in India (Sikora & Greco, 440 0.29 2.98 a 4.95 a 0.85 a 1990). 0 3.08 a 5.63 a 0.78 a 0.78 a 500 4.68 2.53 a 5.82 a References 1000 3.96 2.52 a 5.76 a 0.85 a Sorghum 0 4.91 a 5.75 a 1.00 a BAU}ARD, P. (1995). Laboratory methods used for the study 500 119 4.71 a 6.06 a 1.02 a of the ecology and pathogenicity of , Longidori­ 1000 1.27 4.87 a 6.12 a 1.10 a dae and Trichodoridae from rainy and semi-arid tropics of West Africa. Fundam. appl. NemalOl.) 18: 63-66. Cowpea 0 4.98a 5.07 a L02a 350 1.14 4.79 a 5.98 a 1.18 a BAU}ARD, P. & MARTlNY, B. (1994). Études nématologiques 700 1.33 4.01 b 6.28 a 1.28 a au Mali, Afrique de l'Ouest. 1. Prospections de deux zones arachidières sur l'arachide et le mil. J. afro Zool., 108 : 217­ 226. plication rate was not affected by soil moisture. There­ BAU}ARD, P. & lv1ARTINY, B. (1995 a). Études nématolo­ fore, the absence of this species from the semi-arid trop­ giques en Mauritanie et au Niger, Afrique de l'Ouest: né­ ics of West Africa remains unexplained. The experi­ matodes associés au mil sauvage (Pennisetum violaceum). J. ment on anhydrobiotic survival showed that the afro Zool.) 109: 51-69.

Vol. 18, n° 5 - 1995 443 P. Baujard & B. Martiny

BAU]ARD, P. & MARTlNY, B. (1995 b). Charaeteristics of the Luc, M., SIKORA, R. A & BRlDGE,] (1990). Plant parasitic soil nematode populations from the peanut cropping area of nematodes in subtropical and tropical agriculture. Wallingford, Senegal, West Africa. J. afro Zool., 109 : in press. UK, CAB International, Institute of Parasitology, XVli + 629 p. BAU]ARD, P. & MARTINY, B. (1995 c). Ecology and pathogen­ iciry of the Hoplolaimidae from the sahelian zone of West SHARA-IA, S. B. (1990). Further investigations on the role of Nrica. 1. Field studies on cavenessi Sher, plant parasitic nematodes in crop growth variabiliry of 1964. Fundam. appi. NemalOl., 18: 261-269. groundnut in Niger. Legumes Pathol. Progr. Rep., ICRJSAT, lndia, 8, 61 p. BAU]ARD, P. & 1'v1ARTlNY, B (1995 d). Ecology and patho­ SHARA1A, S. B., SUBRAHMANYAM, P. & SARR, E (1990). geniciry of the Hoplo1airnidae from the sahelian zone of Plant parasitic nematodes associated with groundnut in Nig­ West Africa. 2. Laboratory studies on Scutellonema cave­ er. Trop. Pest Manag., 36 : 71-72 nessi Sher, 1964. Fundam. appl. NemalOl., 18: 335-347. Sl-LARMA, S. 8, SUBRA..H.MAl'-TYAM, p., SARR, E. & Vc\;-'; RIEL, BAU]ARD, P. & MARTINY, B. (1995 e). Ecology and pathogen­ H. (1988). Plant parasitic nematodes associated with iciry of the Hoplolairnidae (Nemata) from the sahelian zone groundnut at Sadoré in Niger. lnt. Arachis Newsl., 4 . 10-11. of West Africa. 3. SCUlelionema clathricaudatum \'(1h.itehead, SI-iARJ\1A, S. B., WALIYAR, F., SUBRAHMANYAM, P. & NDUN­ 1959. Fundam. appl. NemalOi., 18: 347-353. GURU, B] (1992). Role of Scutel/onema clathricaudawm in BAU]ARD, P. & MARTINY, B. (1995 f). Ecology and pathogen­ etiology ofgroundnut grovvth variabiliry in Niger. Pi. & Soil, iciry of the Hoplolaimidae (Nemata) from the sahelian zone 143: 133-139. of West Africa. 4. The genus Aphasmatylenchus Sher, 1965. SIDDIQI, M. R. (1974). Hoplolaimus pamrobuslUs. CfH. De­ Fundam. appi. NemalOi., 18: 355-360. script. Pi. parasit. Nematodes. Set 3, No. 33 : 3 p. BAU]ARD, P. & MARTI1'-TY, B. (1995 g). Ecology and pathogen­ SIKORA, R. A. & GRECO, N. (1990). Nematode parasites of iciry of the Hoplolaimidae (Nemata) from the sahelian zone food legumes. ln: Luc, M., Sikora, R. A. & Bridge,] (Eds). of West Africa. 5. Aorolaimus macbethi (Sher, 1964) Forrun­ Plant parasitic nematodes in subtropical and tropical agricul­ er, 1987. Fundam. appi. Nematoi., 18: 427-433. ture, Wallingford, UK, CAB International, Institute of Para­ sitology: 181-235. BRlDGE, (1973). Hoplolaimus seinhorsti, an endoparasitic J. VAN DEN BERG, E. (1976). Hoplolaimus seinhoTSti. CfH De­ nematode ofcowpea in Nigeria. PI. Dis. Reptr, 57 : 798-799. script. PI. parasit. Nematodes. Set 6, No. 76 : 3 p. GOODEY, J. B. (1957). Hoplolaimus proporicus, n. sp. (Hoplo­ WI-DTEHEAD, A. G. (1959). Hoplolai/nus angustulatus n. sp. laimidae : Tylenchida). NemalOlogzca, 2 : 108-113. (Hopiolaimidae: Tylenchida). NemalOlogica, 4: 99-105.

444 Fundam. appl. NemalOl.