Root-Parasitic Nematodes of Rice

Articlebibliographique

ROOT-PARASITIC NEMATODES OF RICE

Renaud FORTUNERand Georges MERNY

ORSTOM, Laboratoire de Nématologie, B.P. TT 51, Abidjan, Côte d’Ivoire and ORSTOM, Laboratoire de Biologie des Sols, 70-74 route d’Aulnay, 93140 Bondy, France

Geographicaldistribution of nematodes Hirschmanniella of whichseveral species have associated with rice been observed associated with rice. H. oryzae is the most frequently encountered inal1 countries whererice is grown, except Europe. Another More than one hundred species of nematodes species, H. spinicaudata, iscommon inWest have been reported from upland and paddyrice Africa andhas been observed once inSouth in many countries (Tab. 1).Their frequency and America. In WestAfrica a geographical gradient importance are very variable and, in mostcases is observed in the distribution of both species : the existence of a parasitic relationship withrice H. spinicaudata is highly prevalent in the humid is probable but has not been demonstrated. countries ,like Ivory Coast whereas H. oryzae is Manyspecies of rootnematodes have been foundmostly in the Sahelian regions (North observed both in dry and irrigatedfields but very Senegal) ; a balanced mixture of both species is few species are found in both situations. Several observed in intermediategeographical areas surveys made by the authors in WestAfrica have (Gambia). shown that a relatively low number of species Ten recognizedspecies of Pratylenchus have are adapted to permanently flooded conditions. been identified parasitizing rice. The most fre- When the field is only temporarily flooded, the quent is P. brachyurus, rather common in Afri- number of species present is higherand the can upland rice fields, which has been observed nematode fauna tends to ressemble that observedonce in South America ; P. zeae isa common in uplandrice fields, which is composedof species parasite of paddy in Africa and South America ; also common in other crops. Unfortunately, in P. indicus is common in India and single a species many papers, the authors fail to indicate with of Pratylenchus (P. vulnus) was observed in the whatkind of rice (paddyor upland rice) the Far East. parasite was associated. Nematodes attacking roots can be dividedinto three categories. SEDENTARYENDOPARASITES Polyphagouspopulations of Meloidogyne MIGRATORYENDOPARASITES belonging tothe ‘7avanica-incognita-arenaria group” have often been observed in the vicinity The most important migratory endoparasites of rice roots in many countries. When they are associatedwith paddy belong tothe genus found in irrigated paddy fields, their relation to

Revue Nématol. 2 (1) : 79-102 (1979) 79 R. Fortuner & G. Merny rice is doubtfull for they are known to be very species ,of wh.ich have been observed in rice. The susceptible to flooding and they are most proba- most wide-spread species is T. martini known to bly associated withplants growingwhen the parasitise rice in the USA, the Middle and Far field is free of water. One species,M. graminicola East and Africa. T. mashhoodi, which is perhaps isespecially correlated withgramineae and, a synonym of the former, is a frequent parasite recently, M. oryzae hasbeen discoveredasso- of rice in West Africa. ciated with irrigated rice. Since 1961, four species of Heterodera have Five species of Criconemoides havebeen been found to be associated with rice : H. oryzae reported from rice. One of them, C. onoensis, a and H. sacchari arepresent in West African polyphagousspecies, discovered in Africa but pgddy, H. graminophila is associated with rice not associated with rice in this area, has been inthe USA and H. elachista has beenfound foundin the U.S.A. where it seems to cause parasitising upland rice in Japan. losses in paddy.

ECTOPARASITES Ten species of Helicotylenchus were reiorted invarious countries in the vicinity of rice, The most frequently encountered ectoparasite mostlyupland. Their actual relationship with belongs to the genus Tylenchorhynchus, several rice is not known.

Table 1 List, and distribution of nematodes associated with rice roots 1 Species Records

Aglenchus agricola (De Man, 1884) Meyl, 1961 USSR (MASLENNIKOVA,1965 b) A. costatus (De Man, 1921) Meyl, 1961 Côte d’Ivoire (MERNY, 1970) A. bryophilus (Steiner, 1914) Meyl, 1961 USSR (MASLENNIKOVA,1965 b)

Aphelenchoides asterocaudatus Das, 1960 USSR (MASLENNIKOVA,1965 b), India (KKERA& CHATURVEDI,1970) A. bicaudatus (Imamura, 1931) Filipjev & Schuur- Japan (IMAMURA,1931), Bengla Desh (TIMM & AMEEN, mans Stekhoven, 1941 1960),Côte d’Ivoire (MERNY, 1970),Brunei (ANON., 1972 a) A. helophilus (De Man, 1880) Goodey, 1933 USSR (MASLENNIKOVA,1963) A. kuehnii Fischer, 1894 USSR (MASLENNIKOVA,1963) A. lagenoferrus Baranovskaya, 1963 USSR (MASLENNIKOVA,1965 b) A. limberi Steiner, 1936 Côte d’Ivoire (MERNY, 1970) A. parietinus (Bastian, 1865) Steiner, 1932 USSR (MASLENNIKOVA,1963) A. subparietinus Sanwal, 1961 USSR (MASLENNIKOVA,1965 b) A. subfenuis (Cobb, 1926) Steiner & Buhrer, 1932 USSR (MASLENNIKOVA,1963) Aphelenchoides sp. Venezuela (LOOF,1964) Aphelenchus avenue Bastian, 1865 BenglaDesh (TIMM, 1955) Côt:e d’Ivoire, SBnkgal and Guin6e (Luc & DE GUIRAN,1960), Venezuela (LOOF,1964) USSR (MASLENNIKOVA, 1965 b), Korea (CHOI, 1972), Cameroun (SAMSOEN& GERAERT,1975) Aphelenchus sp. Guyana (ANON., 1971) Basiria aberrans (Thorne, 1949) Siddiqi, 1963 USA (ATICINSet al., 1955 a) B. gracilis (Thorne, 1949) Siddiqi 1963 Côte d’Ivoire (MERNY, 1970)’ B. graminophila Siddiqi, 1959 Iran ( KHEIRI,1972)

80 Revue Nématol. 2 (1) : 79-102 (1979) Nematodes of rice

Species Records

Boleodoroides oryzae Mathur, Khan & Prasad, 1966 India (MATHUR et al., 1966)

Caloosia exilis Mathur, Khan, Nand & Prasad, 1969 India (MATHUR et al., 1969) C. paxi Mathur et al., 1969 India (MATHUR et al., 1969)

Criconemoides curvatus Raslii, 1952 Guinée (Luc, 1959 b), Thailand(TAYLOR, 1968), Côte d’Ivoire (MERNY, 1970),Sénégal (FORTUNER& MERNY, 1973), Cameroun (SAMSOEN& GERAERT, 1975) C. komabaensis (Imamura, 1931) Taylor, 1936 Japan (IMAMURA,1931) (species inquirenda) C. onoensis Luc, 1959 USA (EMBABI,1967), Côte d’Ivoire (Luc, 1970) C. ornatus (Raski, 1952) Raski 1958 Sénégal (FORTUNER,1975) C. palustris Luc, 1970 Côte d’Ivoire (MERNY, 1970),Sénégal (FORTUNER& MERNY, 1973) C. rusficus (Micoletzki, 1915) Taylor 1936 Bengla Desh (TIMM, 1955) C. sphaerocephalus Taylor, 1936 Emp.Centre Africain (Luc, 1970) Japan (MOMOTA & OSHIMA, 1973) Criconemoides sp. Taiwan(LIN, 1970), El Salvador (INTERIANO Mufioz, 1970), Ghana (ADDOH, 1971), Guyana (ANON.,1971)

Ditylenchus intermedius (De Man, 1880) Filipjev, 1936 Japan (IMAMURA,1931) USSR (MASLENNIROVA,1965 b)

Ditylenchus sp. USA (ATKINSet al., 1955), Bengla Desh (TIMM& AMEEN, 1960), Venezuela (LOOF, 1964), Panama (YEPEZ& MERE- DITH, 1970), Cameroun (SAMSOEN& GERAERT, 1975)

Filenchus pliformis (Biltschlii, 1873) Meyl, 1961 Japan ( IMAMURA, 1931),USSR (MASLENNIKOVA,1965 b), India (KHERA& CHATURVEDI,1970)

Helicotylenchus cavenessi Sher, 1966 Nigeria (CAVENESS, 1967), Iran (KHEIRI,1972) H. crenacauda Sher, 1966 Indonesia (SHER, 1966), Taiwan (LIN, 1970) H. dihystera (Cobb, 1893) Sher, 1961 = H. crenatus Madagascar (Luc, 1959 a), Venezuela (LOOF,1964), Côte Das, 1960 d’Ivoire (MERNY, 1970),India (NANDAKUMAR & RAO, 1974) H. erythrinae (Zimmerman, 1904) Golden, 1956 Thailand (TAYLOR, 1968),USA (HOLLIS,1969 a), Venezuela (YEPEZ& MEREDITH, 1970) H. exallus Sher, 1966 Cameroun (SAMSOEN& GERAERT, 1975) H. flatus Roman, 1965 Côte d’Ivoire (MERNY, 1970) H. microcephalus Sher, 1966 Côte d’Ivoire (MERNY, 1970) H. multicinctus (Cobb, 1893) Golden 1956 Bengla Desh (TIMM, 1955), Ghana (ADDOH, 1971) H. pseudorobustus (Steiner, 1914) Golden, 1956 Nigeria (CAVENESS,1967) Helicotylenchus sp. USA (ATKINSet al., 1957), Sénégal and Guinée (Luc & DE GUIRAN,1960) Emp. Centrafricain (Luc, MERNY & NETSCHER, 1964), USSR (IVANOVA, 1968), Guyana (ANON. 1971),Brunei (ANON., 1972 a),’ Gambia, (FORTUNER& MERNY, 1973), Rhodesia (.ANON., 1973a)

Hemicriconemoides cocophilus (Loos, 1949) Chitwood Côte d’Ivoire (MERNY, 1970) Sénégal and Gambia (FOR- & Birchfield, 1957 TUNER & MERNY, 1973) H. intermedius Dasgupta Raski &Van Gundy, 1969 Korea (CIIOI, 1972)

Revue Nématol. 2 (1) : 79-102 (1979) 81. R. Fortuner h G. Merny

~~ Species Records

Hemicycliophora belemnis Germani & Luc, 1973 Sénégal ( FORTUNER,1975) H. diolaensis Germani & Luc, 1973 Sénégal (GERMANI& Luc, 1973) H. nigeriensis Germani & Luc, 1973 Nigeria (GERMANI& Luc, 1973) H. oostenbrinki Luc, 1958 Côte d’Ivoire (MERNY, 1970), Ghana (ADDOH, 1971) H. paradoxa Luc, 1958 Côte d’Ivoire (Luc & nE GUIRAN,1960) H. similis Thome, 1955 Madagascar (Luc, 1959 a) Hemicycliophora sp. Guyana (ANON., 1971),Cameroun (SAMSOEN& GERAERT, 1975)

Heterodera elachista Ohshima, 1974 Japan (OKADA,1955) H. graminophila Golden & Birchfield, 1972 USA (BIRCIIFIELD,1973) H. oryzae Luc & Rerdon, 1961 Côte d’Ivoire (Luc & BERDON, 1961) Sénégal (FORTUNER & MERNY, 1973) H. sacchari Luc & Merny, 1963 Côte d’Ivoire (MERNY, 1970), Sénégal and Gambia (FOR- TUNER & MERNY, 1973)

Hirschmanniella belli Sher, 1968 USA (SHER, i968) H. caudacrena Sher, 1968 USA (%ER, 1968) H. gracilis (De Man, 1880) Luc & Goodey 1963 Taiwan (LIN, 1970), India (MATHUR& PRASAI),1971) H. imamuri Sher 1968 Japan (IMAMURA,1931), Iiorea (CHOI, 1972) H. mangaloriensis Mathur & Prasad, 1971 India (MATIIUR & PRASAD,1971) H. mucronata (Das, 1960) Luc & Goodey, 1963 India ( DAS, 1960), Philippines, Thailandand Bengla Desh (SIIER,1968) H. orgzae (Van Breda de Haan, 1902) Indonesia (VANBREDADE HAAN,1902), Japan (IMAMURA, 1931),Bengla Desh (TIMM,1955), USA (ATKINSet al., 1955 a), SriLanka & Malaysia (JOHNSTON,A., 1958), Madagascar (Luc, 1959 a), Thailandand Philippines (THORNE,1961), India (BIRAT, 1965), Sierra Leone & Nigeria (SHER, 1968),Venezuela (SHER, 1968),El Salvador (SHER, 1968), Taiwan (SHER,1968), Korea (PARKet al., 1970), Ghana (ADDOH, 1971),Sénégal and Gambia (FOR- TUNER & MERNY, 1973), Mauritanie (FORTUNER, 1975). H. spinicaudata (SchuurmansStekhoven, 1944) Cameroun (Luc,1957), Venezuela (LOOF,1964), Nigeria Luc & Goodey, 1963 (SHER, 1968), Côted’Ivoire (MERNY, 1970),Sénégal and Gambia (FORTUNER& MERNY, 1973) H. thornei Sher, 1968 Indonesia ( SHER, 1968) Hirschmanniella spp. Hong Kong & New Zeland (SIIER,’1968)

Hoplolaimus clarissimus Fortuner 1973 Senegal (FORTUNER& MERNY, 1973) H. tylenchiformis von Daday, 1905 -- H. coronatus Bengla Desh (TIMM, 1955) Cobb, 1923 H. indicus Sher, 1963 India (Birat, 1965)

Hoplolaimus sp. USA (ATKINSet al., 1957)

Hypsoperine sp. Costa Rica (FIGEROA& JIMENEZ, 1974)

Lelenchus discrepans (Andrassy, 1954) Meyl, 19.61 Côte d’Ivoire (MERNY, 1970) i L. inprmus (Andrassy,1954), Meyl, 1961 , USSR (MASLENNIKOVA,1965 b) L. leptosoma (De Man, 1880) Meyl, 1961 Japan (IMAMURA,1931) Nematodes of rice

Species Records

Longidorellaparva Thorne, 1939 (SURYAWANSHI,India 1971)

Longidorus sp. USA (ATKINS, FIELDING& HOLLIS,1955 a), Emp. Cen- trafricain (Luc et al., 1964), Senegal (FORTUNER,1975)

Malenchus andrassyi Merny, 1970 Côte d'Ivoire (MERNY, 1970)

Meloidogyne arenaria fhamesi Chitwood, 1952 South Africa (VAN DER LINDE,1956) M. exigua Goeldi, 1887 Thailand (KANJANASOON,1964) M. graminicola Golden & Birchfield, 1965 Laos (GOLDEN& BIRCHFIELD,1968), India(PATNAIIC, 1969), Thailand (BUANGSUWONet al., 1971) M. incognita (Kofoid & White, 1919) Chitwood, Egypt(IBRAHIM, IBRAHIM & REZK,1972) 1949 M. incognifaacrita Chitwood & Oteifa 1952 Japan(ICIIINOE, 1955), South Africa (VANDER LINDE, 1956), Côte d'Ivoire and Guinée (Luc& DE GUIRAN,1960) M.javanica (Treub, 1885) Chitwood 1949 South Africa (VANDER LINDE,1956), Thailand(LEAUM- SANG & KANJANASOON,1961), COmOI'eS(VUONG, 1972), Egypt (IBRAHIM, IBRAHIM& REZK, 1972) M.oryzae Maas, Sanders & Dede, 1978 Indonesia (MAAS, SANDERS& DEDE, 1978) Meloidogyne sp. USA (TULLIS,1934, STEINER,1934), Brasil (MONTEIRO, 1968), Taiwan (LIN, 1970), Ghana (ADDOH,1971) Bengla Desh (Ou, 1972) Sénégal (FORTUNER& MERNY; 1973)

Paralongidorusberyllus Siddiqi & Husain, 1965 India(SIDDIQI & HUSAIN,1965) P. cifri (Siddiqi, 1959) Siddiqi,Hooper & Khan,India (BIRAT, 1965) 1963 P. oryzae Verma, 1973 India (VERMA,1973)

Paraphelenchzzs'pseudoparietinus (Micoletzky, 1922) BenglaDesh (TIMM, 1955), URSS (MASLENNIROVA, Micoletzky, 1925 1965 b)

Paratylenchus aquaticus Merny, 1966 Côte d'Ivoire (MERNY,. 1966)

Parafylenchus sp. USA (ATKINSel al., 1957), Sénégal and Gambia (FORTU- NER & MERNY, 1973), Cameroun (SAMSOEN& GERAERT, 1975)

Pelfamigratus nigeriensis Sher, 1963 Nigeria (CAVENESS,1967), Sénégal ( FORTUNER&'MERNY, 1973)

Pratylenchus brachyurus Godfrey, 1929 Madagascar (Luc, 1959 a), CGte d'Ivoire(Luc & DE GUIRAN,1960) Egypt(OTEIFA, 1962), Emp.Centrafri- cain & Congo (Luc, MERNY & NETSCIIER, 1964), Brasil (MONTEIRO, 1968), Nigeria (BRIDGE,1972), Senegal ( FORTURNER,1975) P. delaffrei Luc, 1958 Emp. Centrafricain & Congo (Luc, MERNY & NETSCHER, 1964) P. goodeyi Sher & Allen, 1953 Egypt ( OTEIFA,1962) P. minyus Sher & Allen, 1953 Egypt (OTEIFA,1962), Korea (CHOI, 1972) P. penetrans (Cobb, 1917) Chitwood & Oteifa, 1952 Egypt ( OTEIFA,1962) P. prafensis (De Man, 1880) Filipjev, 1936 Egypt ( OTEIFA,1962) P. sefaensis Fortuner, 1973 Senegal (FORTUNER,1973) P. fhornei Sher & Allen, 1953 Egypt (OTEIFA,1962)

Revue Nématol. 2 (1) : 79-102 (1979) 83 R. Fortuner & G. Merny

Species Records

P. vulnus Allen & Jensen, 1951 Korea (CHOI, 1972) P. zeae Graham, 1951 USA (ATKINSet al., 1957), Egypt (OTEIFA,1962), Venezuela (LOOF,1974), Brasil (MONTEIRO, 1968), CBte d’Ivoire (MERNY, 1970), Cuba (GATEVA& PENTON,1971), Rhodesia (ANON., 1972), Cameroun (SAMSOEN& GERAERT, 1975), Sénégal (FORTUNER,1975) Pratylenchus sp. Sénégal and Guinée (Luc & DE GUIRAN,1960), Bengla Desh(TIMM & AMEEN,1961), Taiwan (LIN, 1970) ‘

Psilenchus hilarulus De Man, 1921 USA (ATIIINSet al., 1955 a), Egypt (ELMILIGY& GERAER?, 1971)

Psilenchus sp. Venezuela (LOOF, 1964), Guyana (ANON.,1971)

Rotylenchulus borealis Loo€ & Oostenbrink, 1962 Côte d’Ivoire (MERNY, 1970) Roiylenchulus sp. Emp.Centrafricain (Luc, MERNY & NETSCHER,1964), Sénégal (FORTUNER& MERNY, 1973)

Rotylenchus sp. USA (ATKINSet al., 1957), Rengla Desh (TIMM& ARIEEN, 1960), Guyana (ANON., 1971)

Seinura propora Siddiqi,’Husain & Khan, 1967 India (SIDDIQI, HUSAIN& KHAN,1967) S. diversa (Paesler, 1957) Goodey, 1960 USSR (MASLENNIKOVA,1966) S. oxura (Paesler, 1957) Goodey, 1960 USSR (MASLENNIKOVA,1966) S. speciosa (Andrassy, 1958) Goodey, 1960 USSR (MASLENNIKOVA,1966)

Scutellonema cauenessi Sher, 1963 Gambia (FORTUNER& MERNY, 1973) S. clathricaudatum Whitehead, 1959 Emp.Centrafricain (Luc, MERNY & NETSCHER,1964), Nigeria (CAVENESS,1967), Côte d’Ivoire (MERNY, 1970) Scutellonerna sp. Cameroun (SAMSOEN& GERAERT, 1975)

Telotylenchus sp. Sénégal (FORTUNER,1975)

Tritlersus annulatus (Merny, 1964) Sher, 1973 Côte d’Ivoire (MERNY, 1970)

Trichodorus christiei Allen, 1957 Brasil ( MONTEIRO, 1968)

Trichodorus sp. Taiwan (LIN, 1970), Ghana (ADDOII, 1971),Sénégal (FOR- TUNER & MERNY, 1973)

Trichotylenchus falciformis Whitehead, 1959 Sénégal (FORTUNER& MERNY, 1973)

Tylenchorhynchus brassicae Siddiqi, 1961. India (CHIIABRA et al!, 1974) , T. clarus Allen, 1955 USSR ( IVANOVA,1968), Egypt (ELMILIGY & GERAERT, 1971) T. gladiolatus Fortuner & Amougou, 1973 Sénégal (FORTUNER& AMOUGOU, 1973) T. martini Fielding, 1956 USA (FIELDING,1956), Bengla Desh (TIMM & AMEEN, 1960),Sierra Leone (HOOPER& MERNY, 1966),Taiwan (LIN, 1970), Iran (KIIEIRI, 1972) T. mashhoodi Siddiqi & Basir, 1959 India(SIDDIQI, 1961), Thailand (TAYLOR, 1968), CAte = T. crassicaudatus Williams, 1960 d’Ivoire (MERNY, 1970), Sénégal and Gambia (FORTUNER = T. elegans Siddiqi, 1961 & MERNY, 1973),Mauritanie (FORTUNER,1975), Came- roun (SAMSOEN& GERAERT, 1975))

84 Revue Ndmatol. 2 (1) : 79-102 (1979) II Nemafodes of rice

Species Records

T. phaseoli Sethi & Swarup, 1968 Cameroun (SAMSOEN& GERAERT,1975) T. sulcatus de Guiran, 1967 Nigeria (BRIDGE,1972) Tylenchorhynchus sp. Guinée (LIJC & DE GUIRAN, 1960),Venezuela (YEPEZ6 MEREDITH, 1970),Guyana (ANON., 1971)

Tylenchus baloghi Andrassy, 1958 Côte d’Ivoire (MERNY, 1970)

T. kirjanovae Andrassy, 1954 , USSR (MASLENNIKOVA,1965 b) T. parvus Siddiqi, 1963 Côte d’Ivoire (MERNY, 1970) T, thornei Andrassy, 195’4 USSR (GAGARIN, 1972) Tylenchus sp. . USA (ATKINSet al., 1955 a) Venezuela (LOOF,1964), Ghana (ADDOH, 1971), Guyana (ANON., 1971), Cameroun (SAMSOEN& GERAERT, 1975)

Uliginotylenchus palustris (Merny & Germani, 1968) Côte d’Ivoire (MERNY & GERMANI,1968) Sénégal and Siddiqi, 1971 Gambia (FORTUNER& MERNY, 1973) U. rhopalocercus (Seinhorst, 1963) Siddiqi, 1971 Cameroun(SEINHORST, 1963) Côte d’Ivoire (MERNY, 1970), Nigeria (BRIDGE,1972) Sénégal and Gambia (FOR- TUNER & MERNY, 1973)

Xiphinema attorodorurn Luc, 1961 Senegal ( FORTUNER,1975) X. bergeri Luc, 1973 Côte d’Ivoire (MERNY, 1970) Sénégal and Gambia (FOR- TUNER & MERNY, 1973)

’ X. cavenessi Luc, 1973 Côte d’Ivoire (Luc, 1973) X. seredouense Luc, 1975 Guinée (Luc & DE GUIRAN,1960) X. insigne Loos, 1949 India (BIRAT, 1965), Rengla Desh (TIMM& AMEEN, 1960) X. orbum Siddiqi, 1964 India (SIDDIQI,1964) X. rotundatum Schuurmans Stekhoven & Teunis- Côte d’Ivoire (MERNY, 1970) sen, 1938 X. setariae Luc, 1958 Emp. Centrafricain (Luc, MERNY & NETSCHER, 1964) Xiphinema sp. Taiwan (LIN, 1970), Ghana (ADDOH, 1971) Xiphinemella caudata Andrassy, 1970 Côte d’Ivoire (ANDRASSY, 1970)

Nematodes and rice plant NEMATODEPOPULATIONS

Penetration of endoparasites Therelationship between parasites .and rice plant have been investigated for endoparasites. In Hirschmanniella oryzae, Van der Vecht and Their penetration in the roots and subsequent Bergman (1952) noted that males and females development as well-as their reproduction have enter the young roots through the epidermis at been studied for species belonging to the genera some distance from the tip and move .in both Hirschmanniella,Meloidogyne and Heterodera. distal and proximal directions through the air For some of the species Our knowledgeis res- channels. The thin lateral roots, having no air tricted to fragmentary observations in thefields channels, are not infected. Mathur and Prasad whereasforothers laboratory experiments (1972 a) observednematodes near the central havebeen done and details are lmown about cylinder, producing hollows in the cortex. their relations with the plant. Hirschmanniella mucronata enters the roots at

Revue Nématol. 2 (1) : 79-102 (1979) 55 R. Fortuner & G. Merny

a distance of about 0,5 cm from the tip (Anon., the roots is observed at heading time whatever 1972). the date of planting. Hirschmanniella imarnuri can enter the roots Populationsdensity in soil decreasesafter alongtheir whole lengthbut most frequently plantingand increases afterharvest when old between 11 to 60% of the total length from the roots begin to rot (Yokoo & Su, 1966). In Sene- tip. After penetration, nematodes move into the gal, Fortuner (1976) observedthat the maximum cortex at the base of root. Adults can migrate percentage of the total population present in the as far as 6.3to 10.3 mm (Goto, 1973). roots is reached between tillering and heading : Merny (1972 a, b) noted that al1 stages of he also noted in pot experiments that the maxi- Hirschmanniella spinicaudata can enter or leave mum final populations was obtainedwith an rice roots and proposed a mode1 derivedfrom inoculum of 300 individuals and more. Nicholson's competitioncurve to express the Kuwahara and Iyatomi (1970) observed that number of individuals which penetrate in terms juveniles of Hirschmanniellaimamuri hatched of thenumber of inoculatednematodes. The in the roots can escape after a certain time and equation : become adult in the soil : only one generation occurs every year in Japan. Inpot experiments Merny(1970 b) studied the development of populations of Hirschman- niella spinicaudata in roots and soil in relation in which II is the inoculum, N the maximum to inoculum.He observed that, whensmall number of nematodesable toenter the root inocula are involved, the increase of population system of a plant and a the probability for a did notfit perfectly the Verhulst-Pearl equation, single nematode to enter the root system, fits formulated by Oostenbrink (1966) : the experimental results. P't Ibrahim,Ibrahim and Rezk (1973) have Pf = observed that juveniles of Meloidogyne incognita b+cPf enter the roots at the distal end and seldom in the zone, of elongation. They are found in any Instead of the inoculumhe considered as the part of the apical meristem or in vascular bun- initial population (Po the number of individuals dles without any particular orientation. havingpenetrated in the roots and the final l?Ieloidogyne graminicola penetrationtakes population (P,) should be expressed bythe a minimum of 41 hours, when hypertrophy and equation : hyperplasybegin in cortical cells, gallsbeing d formed in 72 hours (Patnaik, 1969). pz f; Al1 stages of Hoplolaimrzs indicus are able to = Pf a b penetrate but 4thst,age juveniles and adults are -Pt+- most often observed in the roots. They produce C C inter-and intracellular galleries in thecortex d which loses its rigidity : they also cause defor-. in which - - represents the smallest inoculum ma'tions in vascular bundles (Das & Rao, 1970). C for which the subsequent developmentof a popu- &Iultiplication lation canbe expected because at smaller inocula there is a low probability of gettinga gravid Van der Vecht and Bergman (1952) observed female or a female and a male having a chanceto that the multiplication rate of Hirschmanniella mate. Merny(1972 a) furtherobserved that oryzae could reach thirteen per generation and nematodes start to be observed in soil 90 days Kuwaharaand Iyatomi (1970) noted that, in aft,erinoculation and the soil-population in- Japan,two generations occured in one year, creases during maturation of rice as if, lacking whereas Fortuner (1976) stated that three gene- room and food, the nematodes escaped from the rations occur in Senegal during a rice crop. In decaying roots. Japathe maximumpApulatio-n ofarasitesin In-India, Dasand Rao (1971) have noted that

. 86 Nématol. Reoue (1979)2 (1) : 79-102 Nematodes of rice

Hirschmanniellamucronata waspresent in generationappear at 35 days.The nematode greater number in soil at heading time. induces an hypertrophy and hyperplasy of cells In Meloidogynegraminicola, Raoand Israel inthe meristem, cortex endodermis and peri- (1972 c).observed that low inocula led to higher cycle of the roots and giant cells form in the multiplication rates, dueto the fact that, highat meristem, cortex and xylem after 10 days. As initialdensities, multiplication is hindered by many as six giant cells per juvenile have been competition for nutrients within the roots. observed (Ibrahim, Ibrahim & Rezk, 1973). Merny (1972 a) noted that the reproduction Roy (1975)observed that rootexudates of rate in pots of Heteroderaoryzae expressed as resistantor susceptible rice cultivarshave no the ratio of the number of second stage juveniles effect on the hatching of eggs of Meloidogyne hatching in the second generation and the num- graminicola. ber of second stagejuveniles inoculated, was Netscher (1969) studied ovogenesis and repro- aboutsix for initial densities between 50 and duction of the two Heterodera species known in

200. The same low multiplication was observed West Africa and 1 observed that H. oryzae is as in Hirschmanniella spinicaudata at low initial diploïd (2n = 18) and amphimictic while H. sac- densities. Chari is triploïd (3n = 27) and parthenogenetic. Populationdevelopment of Heterodera ela- Cadet, Merny and Reversat (1975) observedthat, clzista (often mistaken for H. oryzae) on upland in H. oryzae sexwas already determined in rice in Japan gave a series of peaks, due to the second stagejuveniles, relative abundance of hatching of eggs in successive generations : three males in cases of overcrowdingbeing due to peaks were observed on late maturing cultivars preferential death of female juveniles, whereas andtwo peaks on early or normal cultivars Cadetand Merny (1978) have shown thatin (Shimizu, 1973). H. sacchari sex was determinedlater, each The population of Pratylenchus zeae, in Cuba, second-stage juvenile being able to develop into is maximum at headingand not at maturity a male or a femalè adult according to external (Gateva & Penton, 1,971). factors. Populations of Criconemoidesonoensis obser- The life cycle of H. oryzae wasstudied by ved early in the crop inUSA disappear at flood- Berdon-Brizuelaand Merny(1964) Who estab- ing and reappear after the drainage preceding lished that thesecond moult occured after three harvest indicating that this species is not well to four days and the third moult nine to ten adapted to flooded conditions(Hollis, 1969 b). days after penetration, the first males appearing after thirteen days and the first females after sixteendays. Hatching of eggs in egg-masses Reproduction and development , and cysts has been studied byMerny (1972 a) in Most of the studies on reproduction concern water and in soil. In water, hatching in cysts is sedentary endoparasites of the genusHeterodera. stimulated by rice-root exudates. In soil, hatch- In Hirschmanniella oryzae, Van der Vecht and ing incpts lasts longer and reaches a higherlevel Bergman(1952) noted that ovipositionbegins than in water while hatching in egg-masses is some days'after penetration of adults and eggs delayed so that egg-masses kept in soil for nine ~ laid in the roots hatch after four to five days. months are still able to liberate infective juve- Nothing is known on the ovogenesis and sexual- niles. ity of the different Hirschmanniella species. Eggs of Meloidogyne incognita hatch between two and eight days in distilled waterbut second INFLUENCEOP NEMATODES ON THE RICE PLANT stage juveniles are obtained in one day in rice pots. Third stage juveniles were observed after eight days and fourth stage juveniles ten days Nematodes living in roots of rice or, at least, afterinoculat.ion. The larval head is nearthe feeding on them inevitably induce troubles in endodermis and its body in the cortex and youngthe physiology of the plant. Symptoms induced females appearedfrom the 15th day and egg to rice plant by root nematodes are neverspeci- massesafter 20 days ;' juveniles of thenext fic. The most frequentis a reduction in tillering.

Revue Némaiol. 2 (1) : 79-102 (1979) 87 R. Fortuner & ‘G. Merny

Hirschmanniella oryzae by Luc(1957) in Cameroon, associated with yellowingrice plants and the same symptoms Van der Vecht and Bergman (1952) considered have beenobserved in Upper-Volta(Germani, this species to beresponsible for the disease pers. comm.). knownin Indonesia as “Omo Mentek”. Later, Hirschmanniella spp. stunted rice in Thailand Van der Vecht (1953) was less affirmative and (Buangsuwon et al., 1971). onlystated that the disease appeared when InIndia Meloidogynegraminicola causes a H. oryzae was present and rice growth conditions yellowing of the, upper partof the leaves. Plants were veryunfavourable. More recently, Ou arestunted, roots proliferate, are thin and (1965) stated that Omo Mentek was not caused forked and bear characteristic galls (Roy, 1973). by H. oryzae but by a virus. However, Van der Similarsymptoms werefound in inoculated Vecht and Bergman (1952) had established that seedlings, tillers were smaller, heading occured H. oryzae couldinduce troubles in rice plant, earlier thannormal and the grains werede- namelygrowth retardation and reduction of formed, but rice will grow satisfactorily with up tillering which could be beneficial for the culti- to 1 785 galls (Patnaik, 1969). In Laos the main vars grown in Java which suffer from an excess attack is on seedlings in nurseries ; they become of tillering due to poor soil fertility. yellow, brown and dry out : galls are fornled on Thorne (1961) estimatedthat, under favorable upland rice but have notbeen observed onpaddy agronomic conditions there is little evidence of although they will appear as soon as the irriga- injury to rice by H. oryzae becauserice roots tion water is removed. develop atthree stages of growth(seedling, In pot experiments in Thailand, Meloidogyne tilleringand end of tillering)and nematodes sp. caused a poor growth, reduction of tillering, developing in the roots during one period remain delayed maturation and a yield decrease (Chan- thereduring the followingone so thatthe tanao,1962). Similar symptoms caused by amount of healthy roots is always sufficient to Meloidogyneincognita and M. javanica were secure nourishment of the plant. observed in Egypt (Ibrahim, Ibrahim & Rezk, InJapan, decreasein tillering induced by 1972). H. oryzae is more important insoils with low rH, Joshi, Ibrahim and Hollis (1975) have noted resulting in a disease called “Akiochi” and yield that oxygen released by the roots was reduced losses are more important in thesesoils. (Kawas- in three-week-oldinfected rice seedlings : the hima, 1964 b). Infested seedlings undergo growth number of galls increased with the size of inocu- retardation, height and weight of the plant are lum although the growth was not affected. decreased and, at t.he same time, the browning Ina pot experiment, in Costa-Rica, Hypso- ’ of theroots ishigher at highinitial densities perine (n. sp. ?) interfered with the growth of (Kawashima & Fujinuma, 1961). According to rice seedlings, roots showing apical thickenings the same authors, nematodes interfere with the withgrowth retardation : the aboveground physiology of the roots, decreasing their oxidiz- symptoms,similar to nutritional deficiencies, ingcapacity and inducing their coloration by consisted of leaf chlorosis and necrosis,poor iron oxide. tillering, stunting and lack of vigour, nutrient H. oryzae decreased tillering and root weight uptake being deficient in infested plants. inIndia (Mathur & Prasad,1972 b). Inthe Heteroderaelachista at initial densities of O U.S.A. the parasite caused decay of the tip of to 10,000 cysts, decreasedtillering from 7to primary roots in the layerof the soil with a high 2.7 tillers per plant (Odaet al., 1963) : the height rH (Hollis, 1967). of plantsand root weight were markedly In India Hirschmanniellamucronafa reduces reduced and grain yield decreased from 7.82 g yield but not ear number and does not retard to 3.15 g atthe highestinfestation level. In tillering but length and weight of roots increase seedlings, there was leaf chlorosis and a reduc- with the size of inoculum, indicating that the tion of tillering and rooting and it was noted plant compensates for the attack by the nema- that the capacity of roots to absorb iron and tode (Panda & Rao, 1971). nitrogen was reduced and they turnedbrownish. -Hi~*chma~niella_s~inicaIldaba,‘wasrecorded Theappearance of symptomsina field was very

88 Nématol. Revue (1979)2 (1) : 79-102 Nematodes of rice

variable, being different in every year (Wata- plants grown in pots on heat-sterilized soil and nabe, 1963). compare with non inoculated controls. Heterodera sp. (djfferent from H. elachista and H. oryzae) has been found on the roots of chloro- Field treatment tic rice plantsin India (Ra0 & Jayaprakash, 1977). Treatmentswith D-D and DBCP ina field Criconemoidesonoensis. Rootsattacked by infested with Hirschmanniella sp., in Thailand parasites belonging to thisspecies have a charac- gave increases in yield of 28 to 30%. Treatments teristic knotty appearance. They are shorter and of nurseries,infested with +Meloidogyne sp., less forked and the size and weight of roots and resultedin better seedling growthbut to no aerial parts of inoculated seedlings are reduced increase in yield (Taylor et al., 1966). (Embabi, 1967). Deformationof the roots caused InJapan, D-D treatment of fields infested by this nematode mainly occurs in the layer of with Heterodera elachista increased yield by 30% soil whith a high rH beneath the biochemically (Nishizawa,Shimizu & Nagashima,1972). reduced zone (Hollis, 1967). Against the same parasite, D-D and EDB gave Interiano Mufioz (1970)could notascertain an increase of 20 to 70% (Yanaka etal., 1962) the pathogenicity of Criconemoides sp. but the authors thought theincrease is too high Tylenchorhynchusmartini. Seedlingsinocu- to be explained by the destruction of nematodes lated in pots with 3,000 individuals per pint of and suggested that the chemicals had some side soil (about 5,300per liter) showed, after two effect,favorable to plant growth. H. elachista weeks, significant reductions in length and dry caused heavy losses in rice over a period of four weight of roots compared with non-inoculated years ; yields were 2,870 kg/ha in the first year controls.The effects were moresevere when and 950 kglha in the fourth year (Watanabe et inoculatedwith “swarming” than with L‘non- al., 1963) andthus, these authors considered swarming” nematodes : plants were stunded and H. elachista responsible forthe failure of continu- chlorotic,nematode aggregates were observed ous upland rice cropping in Japan. in vitro around the roots and it was suggested The use of Dasanit(fensulfothion) against thatthe nematodesinhibit the absorption of Criconemoides onoensis in the USA, resulted in nutrients or excrete toxins (Joshi& Hollis, 1976). an increaseof 2.0% in paddyyield (Hollis, 1969a) Pratylenchusindicus. Rao and Prasad (1977) and, against Tylenchorhynchus martini and other stated that this parasite can cause the death of nematodes,methyl bromide gave increases of rice 40 to 50 days after germination. 45% at harvest (Atkins, Fielding& Hollis, 1957).

Inoculation- experi~nents YIELDLOSSES In Senegal, the vigour of paddy in fertilized microplots infested with Hirschmanniella oryzae, Determination of yield losses caused by nema- was much reduced as compared with plants in todesis very difflcult. Treatment of infested non-infested plotswith yieldincreases of the soils withnematicides often givegood yield order of 42%.However, thecharacteristics of increases. But soil treatments are drastic, other grains (weight of 1,000 grains, germination, etc.) pathogens can be killed and examples are known was the samein both conditions (Fortuner, of yieldincreases in nematicide-treated soils 1974). Fortuner (1977) experimented on micro- which had not been caused by the destruction plots, insoil free of nematodes. Half of the microplots were inoculatedwith H. oryzae. In each # of nematodesbut of otherpathogens such a fungimicro-arthropods.or Although yield category,inoculated or control, half of the increaseswith nematicides are, in most cases, microplotsreceived fertilizers. He ascertained caused by nematode destruction, results of such that : trials must be interpreted with caution. (a) in non-fertilized plots, the yield losses due A better way to determine the part played by to nematodes were higher (31%) than in ferti- nematodesin yielddecreases is toinoculate lized plots (19%),

Revue Nématol. 2 (1) I 39-102 (1979) 89 R. Fortuner & G. Merny

(b) the yieldincrease due to fertilization is between two rice crops : soil moisture and pre- higher in inoculated (36%) than in non-inocu- sence or absence of host roots. As these nema- lated plots (17%). todesare endoparasites, after rice has been Inoculation with 5,000 and 10,000Hirschman- harvested high populations are still inside the niellamucronata to one-day-old rice seedlings roots and a variable number are found in the decreased the yield by 50% and 70% respec- soil. tively. If parasites were inoculated to40-day-old In the laboratory Van der Vecht. and Berg- plants, the decrease was still important (45 and man (1952) observed that, in every conditionsa 65%). On 70-day-old plants there was no effect highproportion of theinitial population of (Prasad & Rao, 1971). Hirschrnanniella oryzae was still alive after ten ' Shimizu (1971) reported yield losses fronl 7.2 weeks. They noted that the death rate ina dry- to18.7% after inoculation with Heterodera ing soil was about the same as in a wet soil or elachista: the losseswere higher when plants Sand and that it 'is generally higher in soil or were inoculatedbetween tillering and heading Sand withoutroots than with roots. Thorne (Shimizu, 1972).'In pottedrice plants inoculated (1961) found numerous quiescent H. oryzae indi- with O, 400 and 4,000juveniles of Heterodera vidualsin the dried up Clayof rice fields in elachista grain yield was'71.2 g, 67.2 g and 59.5 g Thailand and Philippines during the dry season. respectively.Maximum losses 'occuredwhen In Japan, Kawashima (1962) States that nema- nematode penetration took placebefore heading. todes survive in winter as juveniles or adults in Under fields conditions losses were more severe dead rice roots ; in very wet fields they often when rice was sownearly, theeffect of the nema- hibernate in the egg stage but they seldom sur- ' todes being more severe in the last stages of rice vive in well drained fields due to drying of soil growth(Shimizu, 1976). in spring. In Korea,H. oryzae can survive for up Inoculation of Meloidogyne graminicola 1,000 to sevenmonths in the water of flooded rice to 8,000 per plant decreased yield by 2.6 to 8% fields (Park, Han & Lee, 1970). In India, Mathur a) per1,000 nematodes added (Rao , & Biswas, and Prasad (1973 have shown that, in aerated 1973). water, 50 to 60% of thenematodes were still alive after eight weeks but only O to 20% sur- Associations ulith other pathogens yivedafter sixteen weeks : in a soil sample allowed to dry for ten months and containing "Rootbrowning" of rice,mainly caused by rice roots, 240 nematodes per liter of soil were soil micro-organismswas slightly increased by still alive while a soil without roots contained the presence of 'Hirschmanniella oryzae (Lee & no more living nematodes after the same period. Park, 1975). Fortuner (1977) statedthat in the Senegal River Delta H. oryzae c,an withstand very severe dry seasonsbetween Januaryand August. Nema- Nematodes and environment todes leave the roots in August when the water cornes. Gotô '( 1969) has reported similar effects and noted that most adults of Hirschmanniella For a parasitic nematode, the main factor in imamuri after having hibernated inold rice roots the environment is the presence of a host plant. are released in soil at, planting time, in June. In However,other factors such as soil conditions a microplot trial with H. oryzae if soil remains canhave an influence on their development, wet after the harvest, roots decay and the nema- multiplication and also on their survival in the todes in the roots arereleased into the soil ;popu- absence of a host. lations decrease slowly and become eradicated after one year. In dry soil, both root and soil SURVIVALBETWEEN RICECROPS populations decrease slowly until, after flooding, populations are released from the roots into the Hirschmanniella spp. soil (Fortuner, 1977). Merny(1972 a) observed Two factors are especially important for the that, in IvorvCoast, the'survival of Hirschman- survival of Hirschmanniella spp. in the interval niellaspinicaudata is better in roots than in soil

90 Nématol., Revue 2 (1) : 79-102 (1979) Nemafodes of rice

and is much longer if the soil containing roots EFFECTOF IRRIGATION ON NEMATODE POPULA- becomes dry thanif it renlains flooded. TIONS The results obtained by Merny (1972 a) and Fortuner (1977) are not in agreement with Van Rice nematodes are more or less adapted to derVecht and Bergman (1952) Who observed flooded conditions. Fortuner (1977 b) observed that soil moisture had no effect on survival of an hydro-topographical gradient in the distribu- nematodes.However, thelatter authors have tion of the nematode fauna of Senegaleserice studied the survivalof nematodes insoil, without fields. Comparing paddyfields and uplandfields, roots and the esperiments were made in Petri as the water table goes down, there isa progres- dishes, in thin layersof soil, witha good aeration sivedecrease, ending in a disappearance, in while Merny and Fortuner worked in microplots upland ricefields, of species characteristic of and pots and hence the results are difficult to flooded rice andother species, usually found compare. associated with other crops, appear. Fortuner’s experiments (1977) have shown that in Sénégal It can be concluded that : species belonging to the genus Hirsclzmunniella are the only ones which are perfectly adapted 1) Nematodes belonging to the genus Hirsclz- to constant flooding. Tylenchorhynchusmash- tna~zniellasurvive longer in roots than in soil. hoodi is adapted tofields where the water table 2) Survival of root populations is shorter in is close to the ground level ; during a flooded ricecrop its populations decrease and increase flooded soil due to quick decaying of the roots. againafter the harvest, when the soil is well 3) In soil, the survical of mobile individuals drained but wet enough to allow rice shoots to is better in wet conditions. grow. Yokoo and Su (1966) observed that popula- 4) Due to anhydrobiosis,the persistance of tions of’Hirsch~nanniella oryzaedecrease rapidly populationsout of the, roots is betterin dry afterwater has beenremoved and thatthis conditions. nematode was found in the 10 to 20 cm layer before drainage and 0-20 cm after. Fukazawa, Little is known of the effect of temperature Mobayashi and Nakata (1963) noted that more onnematode survival. Ip India,Mathur and nematodes were found in fields badly or poorly Prasad(1973 a) reported that, in soil samples drained. Another example of the specialization kept in sealed plastic bags for eight weeks, at of Hirschrnanielle oryzae is given by Kawashima different temperatures, 57.6% of H. 0ryzu.e indi- (1962) Who notes thatthis parasite doesnot viduals were still alive in bags kept at 150, 90% attack uplandrice except when grown inflooded at 350 and none at - 20. In fallow fields, they conditions. H. oryzae commonly occurs in badly have observed that the parasitecould withstand drainedpeaty and humiferous fields where high temperatures (35 to 450) in May and June. “Akiochi” diseaseiscommon (Kegasawa & Kawashima, 1962). Heterodera.oryzae Meloidogyne graminicola is poorly adapted to flooded conditions. Itspenetration in roots is Juveniles of this species survive less than best when soil moisture is 32% and its develop- 30days in soil inIvory Coast and they die ment is favoured bya mois ture of 20 to 30 % and quicker in flooded than in wet soil ; eggs can by soil dryness at rice tillering and earing (Rao survive in egg-masses for nine months and for & Israel, 1971 a, 1972 b)< more than two years in cysts (Merny, 1972 a). Like Tylenchorlzynclzusmashhoodi, T. mu.rtini Reversat (1975 a) has shown that, although this develops best at low soil moistures : 40 to 60% species can live in flooded fields in heavy soils of field capacity (Johnston, T.M., 1958). whereoxygen is scarce, the juveniles have an aerobic metabolism. Under anaerobic conditions, EFFECTOF SOIL CONDITIONS they become inactive but, hence, they survivea longer time. Hirsclz~namiellaoryzae appearsto develop

Revue Nématol. 2 (1) : 79-108 (1979) 91 R. Portuner h G. Merny

~~~~ ~

better in Clay soils of North Senegal than in the mediate or dry nurseries or from directly sown sandy soils of the South (Fortuner, 1977) and fields (Miura & Shoji, 1964). Higher populations the highest populations in India, are found in are found inplanted up rice thanin directly heavy Clay soils (Mathur & Prasad,1971) On sown rice roots(Nakazato, Kawashima & theother hand, Fortuner (1977) noted that Kurosawa, 1964). Hirschmanniella spinicaudata developed equally Heteroderaelachista normally attacks upland well, in Senegal, on both types of soil. rice. However,Kawashima (1964 a) observed Rao and Israel (197.1 a) reported that soil pH that flooded rice could be attackedif it was sown had no effect on the developmentof Meloidogyne directly into dry land and flooded one month graminicola but both movement and penetration later : rice planted up traditionally in an adjoin- into roots were better in coarse textured soils ing field remained free of nematodes. (particles over 53 pm). Clay soils are less favour- ableand a linear relation has been observed Crop rotations ' betweenpercentage of sand in soil andthe activity of nematodes,the increase of their In Japan,a crop of soybean reduced the popu- effect onroot growth and the appearance of lations of Heterodera elachista; three successive galls (Ra0 & Israel, 1972 d). crops did not lead to complete eradication but In in uitro experiments, penetration of Hete- the following upland rice crop yielded 2.8 to 3.7 roderaoryzae is best in artificial soil composed times more than compared with rice which had of particles between 100 and 160 pm and very been cultivated for more than four years(Nishi- poor if particles are bigger than 250 pm. On the zawa, Shimizu & Nagashima, 1972). After non- other hand, with Hirchmanniella spinicaudata, hostplants had been cultivated forone year, penetration is best with particles between 160 H. elachista populations were reduced to 74% of the initial population and to 2-3% after two and 250 pm and still occurs up to 630 pm : the optimum particle size for penetration is appar- years, then agood upland rice yield wasobtained ently related to thesize of the nematode (Rever- butnematode populations built upagain sat & Merny, 1973). (Hoshino etal., 1961) ' . Inpot experiments, soybean, sweet potato and Cotton haveproved to bebad hosts of Control Crisonemoidesonoensis in USA (Alhassan & Hollis, 1969). One crop of soybean reduced the Although Van der Vecht and Bergman (1952) population of nematodes and a good rice yield stated that cleaning the fields, keeping a good was obtained : the beneficial effect was still fertility and proper agronomic practices should obvious on the next ricecrop. Hollis (1969 b) allow rice to withstand attacks by Hirschman- reported that maize is attacked by this nema- niellaoryzae withoutnoticeable losses, control tode. of rice nematodes is diacult and several kinds of control measures should be considered includ- Effect of fertilization ingagricultural pratices, varietal resistance, InJapan, Tomonaga and Kurokawa (1964) biological control and chemical control. havenoted that calciumsilicate and compost reduce Hirschmanniella sp. populationsand CULTURALCONTROL increase yield. Ishikawa (1965)stated that nitrogenousfertilizers favour the development of Planting conditions nematodepopulations (without stating which Earlyplanting decreases populations of specieswere involved)and increase the decay Hirschmanniella oryzae and the number of juve- and browning of roots. The height and tillering niles of H. imamuri but did not seemto influence of plantsare improved. Inversely, potassic thenumber of adults of H. imamuri (Sato, fertilizers andcompost keep nematodepopu- Koyama & Koshihara,1970). Higher popula- lations at a low level. tions of Hirschmanniella oryzae are reported in Fertilizerscontaining respectively 64.3 and sesdLngsfrom flooded nurseries than frim inter- 30_0/, of silicic acid and varibus micro-elements

92 Ne'matol. Revue (1979)2 (1) : 79-102 Nematodes of rice

reduced .' Hirschmanniella sp.by 15% and xylem and a high content of aspartic acid and increasedyield by 5% (Yokoo & Morimitzu, alanine (Jena & Rao. 1974). Penetration is more 1969). rapid and life cycle shorter in susceptible than In India, Mathur and Prasad (1972 b) noted in resistant cultivars (Jena & Rao, 1976). that fertilizers improve plant growth as well as Thecultivar International is resistant to nematodepopulations. However, the same Meloidogyne incognita acrita (Ibrahim, Ibrahim authors (1971) stated that populationsof Hirsch- & Rezk, 1972) but Kumazawa (1965) found no manniellaoryzae were lower in fields fertilized cultivar resistant toHeterodera elachista. Twelve with urea. cultivars tested by Figueroa and Jimenez (1971) Nitrogen at the rate of 40 kg/ha and/or phos- are al1 susceptible to Hypsoperine sp.(syn. of phorus increase the reproduction of Meloidogyne Meloidogyne). graminicola (Ra0 & Israel, 1971 a).

BIOLOGICALCONTROL VARIETALRESISTANCE In Japan, afertilizer containing 3% of spores Kawashima (1963) tested seventeen rice cul- of Arfhrobotrys sp., a fungus predator of nema- tivars for their susceptibility to Hirschmanniella todes, applied at the rateof 60 kg-ha reducedthe oryzae and found no differencebetween them. populations of Hirschmanniellaoryzae and

Withthe sameparasite, Park, Han and Lee Hirschmanniellaimamuri, and increasedtillering , (1970) tested 270 cultivars and found that five by 7%, number of grains and straw weight by of them were relatively resistant (less than one 10% and yield by 6% (Yokoo, 1971).

nematode per gram of roots). I Gemma, Shibuya and Kikuchi (1964) observed TheLe is a considerable literature on testing that tadpoles could eat nematodes.

rice cultivars resistant to Meloidogyne gramini- When rice fields are flooded the soil becomes , cola : anaerobic and populations of sulphate reducing Golden and Birchfield (1968) tested 30 cultivars bacteria increase : under these conditions popu- and Manser (1968, 1971) tested 80 cultivars and lations of Tylenchorhynchusmpriini decrease. no resistance was observed in both cases. Laboratory tests have confirmed that H,S con- Sampath, Rao and Roy (1970) and Roy (1973) centrations produced by bacteria were toxic to foundtwo resistant cultivars and Rao et al., nematodes (Rodriguez-Kabana, Jordan & Hollis, (1969)reported that cultivarsTKM and6 1965).The existence of the samephenomenon Patna 6 were resistant while four others were in Senegal has been shown in field observations moderately susceptible. In addition, the cultivar and laboratory trials (Fortuner & Jacq, 1976). TKM 6 was also resistant to stem borer. Sam- Microplot experiments have given good results path, Rao and Roy (1970) stated that resistance and it might be possible to artificially increase to Meloidogynegraminicola haschemicala bacteriaactivity after the harvest to further origin. reduce the number of viable nematodes present Penetration of juveniles of thisparasite is in the soil. However, some or al1 eggs laid inside the same in susceptible (IR 8, Pusa 2-21) as in the roots are notkilled (Jacq & Fortuner, 1979). resistantcultivars (IR 20, BasantBahar) but subsequent development is reduced in resistant cultivars, a smaller number of galls being pro- CHEMICALCONTROL duced (Roy, 1975). . According to Jenaand Rao (1974) the resis- Fumigants tanceto M. graminicola of cultivarsHamsa, IR 5, IR 47, IR 2,Manaharsali and Baharsia In Japan, numerous soil-treatment trials have has no chemical origin but is due to the fact that been donswith D-D. (dichloropropane-dichlo- they have a smaller number of roots, a denser ropropene),EDB (ethylene dibromide), DBCP rootletsystem, a sclerifed exodermis, a thin (dibromochloropropane),methyl bromide and cortex and central cylinder, little phloem, much metham-sodium. Results have been summarized

Revue Némafol. 2 (1) : 79-102 (1979) 93 R. Portuner Ce- G. iierny

by Ichinoe (1968) Who noted that yield increases Hayward 327-1 and carbofuran are less efficient caused by D-D cannot always be explained by than fensulfothion. Aldicarb treatment appears the reduction innematode population. Yield to increase nematode populations and weight of increases with DBCP are lower and EDB is phy- plants. (Hollis, 1969 a.) totoxic,causing a dark green discoloration of Applications of phenamiphos, Dupont D-110 leaves and excessivetillering. Al1 chemicals and fensulfothion incorporated by cover-cropp- tested reduce nematode populations during rice ing or flooding has caused the development of crop and the treatments have a beneficial effect adventicious plants and unfortunately weeding on nitrifying bacteria and increase the quantity with propanil was prohibited because it reacts of nitrogen available in soil. Hence, the amount with the nematicidesand becomes phytotoxic of nitrogenousfertilizers to be appliedcan be (Hollis, 1972). reduced. In addition various positiveor negative InIndia, Samantaray and Das (1971)have effectshave been observed on theseverity of treated Hirschmanniella sp. and Hoplolaimus sp. attacksby other pathogens, e.g.insects or with thionazin (12,5 l/ha) dichlofenthion(24 l/ha) fungi. andtwo fumigants : DBCP (11,5 l/ha)and In Thailand, D-D, DBCP and methly bromide metam-sodium (393 l/ha). After 45 days,’ both have beenused tocontrol Meloidogyne sp.in systemicshad eliminated al1 nematodes, while nurseriesand Hirschmanniella sp.under field DBCP had eliminated Hoplolaimus sp. and mar- conditions.In nurseries vigour and weight of kedly reduced Hirschmanniella sp. populations ; plants were increased and planting up could be metam-sodium was not effective after 60 days made ten to forteen days earlier and in thefield, and no information was given about yield. yield was increased by 24 to 36% but the treat- Fensulfothion, phorate and cytrolane are the ments wereuneconomic (Taylor etal., 1966 ; most efficient chemicals against Hirschmanniella Taylor,1968). mucronata (Anon. 1973 b). In India Hirschmanniella oryzaewas cont,rolled by D-D applications at the rate of 400kg/ha The following treatments are efficient against injected at a depthof about 22 cm and by adding Meloidogyne graminicola : DBCP to irrigation water at the rateof 120 cm3 - Soaking rice seeds for twelve hours in 500 per 100 m2. Treatments were done three weeks p.p.m. of oxamyl or phorate orfor 24 hours before sowing in nurseries and five weeks before in 1,000 p.p.m. of fensulfothion or carbofu- planting out (Mathur & Prasad, 1973 b). ran (Prasad & Rao, 1976 a). In USA treatments against Tylenchorhynchus - Soakingseedling roots in100 p.p.m. of martini, Hirschmanniella oryzaeand other nema- oxamyl. (Prasad & Rao, 1976 b). ‘todes with methyl-bromide applied undercovers - In pots, after inoculation,. application of (100 g/mz)is better than with EDB (113 l/ha at 50 p.p.m. oxamyl or fensulfothion or 100 83%),D-D (378 l/ha) or DBCP (47 l/ha).MethyI- p.p.m. of carbofuran and DBCP (Prasad & bromide was equally active against Helminthos- Rao, 1976 c). poriumoryzae and Echinochloa but did not prevent a slight attack of “straighthead”. Yield Prasadand Rao (1973) planted rice inpots was increased by 45% (Atkins& Fielding, 1956 ; containing Tylenchorhynchus sp. treated imme- Atkins, Fielding & Hollis, 1957). diately with oxamyl, DBCP, dursban and phorate at rates from 100 to 2,000 p.p.m., either by Non fumigants foliar application or directly in the soil. Foliar appliedoxamyl reduces thenematodes from In USA, treatmentsagainst Criconemoides 59.&74.3% even at low concentrations while the onoensis withfensulfothion allows the highest other products are only efficient at 2,000 p.p.m. yield increase whenthe chemical is appliedearly, On the other hand, in thesoil application, other on 25 cm high plants, before flooding of the field. products give control of 94 to 100% at al1 con- Yield increase is then 20%, about 1,000 pounds centrations while oxamyl reduces the nematode populationby 68-95% at concentrationsfrom

94 Nématol. Revue (1979)2 (1) : 79-102 Nematodes of rice

Chhabra, Sajjan and Singh (1974) found that ALHASSAN,S. A. & HOLLIS,J. P. (1969). Ring nema- carbofuran (1.5 kg/ha)fensulfothion (5 kg/ha) toderatings of ricearea rotation crops. (Abstr.). andaldicarb (3 lcg/ha) appliedone week after Phytopathology, 59 : III. sowing were al1 efficient against Tylenchorhyn- ANDRASSY,1. (1970). Einigeneue Nematodenarten ausWestafrikanischen Reisfelder. AnnlsUniv. chusbrassicae in nurseries but phorate had no Scient. bpest. Rolando Eotvos, Sect. biol., 12 : 243-254. effect. ANON.(1971). Annual report for the year 1970. Guyana In dapan, Nishizawa, Shimizu and Nagashima Ministry of Agriculture : III 8 ; III 18-19. (1972) applied methomyl granules (5%) at the ANON.(1972 a). Annual report of the plant pathologist. rate of 40 kg/lO ares at sowing. Brunei, Department of Agriculture : 14-16. ANON.(1972 b). Report of thesecretary for agriculture Miscellaneous for the period lsfOctober 1970 to 30th September 1971. Salisbury, Rhodesia : 26. Yokoo, Ookuchi and Teramachi (1967), noted ANON. (1973 a). Abridgedreport for theyear endetb that applyingan herbicide : 4-dichloro- 2-me- 30th June 1973, h‘utsaga Research Station. Tobacco ResearchBoard, Salisbury, Rhodesia : 15-17. thylphenoxyacetic acid(MCPA) increased the ANON.(1973 b). Annual technical report, Central Rice populations of Aphelenclzoides sp. while penta- Research Institute, Cuttack. New Delhi, India, 301 p. chlorophenol (PCP) anda diphenylether (MO) ATICINS,J. G. & FIELDING,M. J. (1956). A preliminary increased those of Hirschmanniellaoryzae. report on the response of rice to soi1 fumigation for Mustard and “neem” oil-cakes reduce Hirsch- thecontrol of styletnematodes, Tylenchorhynchus Inanniella oryzae populations and improve plant martini. Pl..Dis. Reptr, 40 : 488-489. growth (Mathur & Prasad, 1973 b). ATKINS,J. G., FIELDING,M. J. & HOLLIS,J. P. In pots, maize-cakesincrease n-butyric acid (1955 a). Parasitic or suspected plantparasitic concentrationsup to nematicidalproportions nematodesfound in rice soils fromTexas and Louisiana. Pl. Dis. Reptr, 39 : 231-222. andpropionic acid, which isalso formed,‘is ATKINS,J. G., FIELDING,M. J. & HOLLIS,J. P. active against nematodes (Hollis & Rodriguez- (1955 b). A newnematode on ricein Texas and Iiabana, 1966).The effects of fattyacids on Louisiana. Pl. Dis. Reptr, 39 : 69. Tylenclzorhynchus lnartini depends on theirmole- ATKINS,J. G., FIELDING,M. J. & HOLLIS,J. P. (1957). cular weight ; the most efficient is butyric acid Preliminary studies on root parasitic nematodes of (Johnston,1959). rice in Texas and Louisiana. Pl. Prot. Bull. F.A.O., ’ Controlling Criconernoidesonoensis in Loui- 5 : 53-56. sianastimulates weed growthmore than rice BERDON BRIZUELA,R. & MERNY, G. (1964). Biologie growth. The control results in lower rice yields d’Heteroderaoryzae Luc & Berdon, 1961. 1. Cycle duparasite et réactions histologiques de l’hôte. in fields infestedwith Cyperusesculentus. On Revue Path. vég. Ent. agric. Fr., 43 : 43-53. ,the other hand, large and significantincreases RIRAT,R. B. S. (1965). New records of parasitic in yield of rice were obtained when weeds were nematodes on rice (Oryza satiaa L.) in Bihar. (Cor- removedand nematodes controlled (Hollis, respondance). Sci. Cult., 31 ; 494. 1977). BIRAT, R. B. S. (1968). Occurrence of Hirschmanniella oryzae (Van Breda de Haan, 1902) Luc & Goodey, 1963, on rice roots in Bihar. (Correspondance). Sci. Cult., 34 : 484-485. ACKNOWLEDGEMENTS BIRCI-IFIELD,W. (1973). Pathogenesisand host- parasite relations of the cyst nematode, Heterodera The authors are grateful to Dr. N. G. M. Hague who graminophila, on grasses. Phytopathology, 63 : 38-40. revised the English text. BISWAS,H. & RAO,Y. S. (1970). Studies on nematodes of rice and rice soils. II. Influence of Afeloidogyne graminicola, incidenceon yields of rice. Oryza, 7 : 59-60. BIBLIOGRAPHY BISWAS,H. & RAO,Y. S. (1971). Toxicological assays of nematicideswith rice nematodes. lndianPhy- ADDOI-1,P. G. (1971). Thedistribution and economic topath., 24 : 159-165. importance of plant parasitic nematodes in Ghana. BRIDGE, J. (1972). Plantparasitic nematodes of Ghana J. agric.Sci., 4 : 21-32. irrigatedcrops in the northern States of Nigeria.

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SamaruMisc. Papers, Ahmadu Bello University, (Nematoda : Tylenchida). Cah. ORSTOM, Sér. Biol., Zaria, Nigeria, No. 42, 17 p. NO. 21 : 25-34. BUANGSUWON,D., TONBOON-EK, P., RUJIRACHOON,G., FORTUNER,R. (1974). Evaluationdes dégâts causés BRAUN,A. J. & TAYLOR,A. L. (1971). Nematodes. par Hirschmanniellaoryzae (VanBreda de Haan, In : Rice diseases and pesfs of Thailand. Ministry of 1902) Luc & Goodey, 1963, nématode endoparasite Agriculture, Thailand : 61-67. des racines du riz irrigué. Agron. trop., Nogent, 29 : CADET, P. & MERNY, G. (1978). Influence of some 708-714. factorson sex-ratio in Heteroderaoryzae and H. FORTUNER,R. (1975). Lesnématodes parasites des sacchari (Nematoda : Heteroderidae). Revue Néma- racines associés au riz au Sénégal (Haute-Casamance fol., 1 : 143-149. et régionsCentre et Nord) et enMauritanie. Cah. CADET, P., MERNY, G. & REVERSAT,G. (1975). Fac- ORSTOM, Sér. Biol., 10 : 147-159. teurs affectant le déterminisme du sexe chez Hete- FORTUNER,R. (1976). Lesnématodes parasites des roderaoryzae (Nematoda : Tylenchoidea). Cah. racinesassociés au riz au Sénégal. These Docteur- ORSTOM, Sér. Biol., 10 : 207-214. Ingénieur, Univ. Cl. Bernard, Lyon, France, no 249, CAVENESS, F. E. (1965). Nematologystudies 1960- 51 p. 1965. End of tour progress report on the nematology FORTUNER,R. (1977). Etude écologique des nématodes project. Lagos,Ministry of Agriculture & natural des rizières du Sénégal. Cah. ORSTOM, Sér. Biol., Ressources, Nigeria, vi + 135 p. 11 (1976) : 179-191. CAVENESS,F. E. (1967). Shadehousehost range of FORTUNER,R. (1977). Fertilisationdu riz etdégâts some nigerian nematodes.Pl. Dis. Reptr, 51 : 115-119. causéspar lenématode Hirschmanniellaoryzae CAVENESS, F. E. (1973). Two nematodeobservations (VanBreda de Haan) Luc & Goodey, C. r. hebd. from Nigeria. (Abstr.). Nematropica, 3 : 1. Séanc. Acad. Agric. Fr., 58 : 624-630. CHANTANAO,A. (1962). [Nematodes of rice and some FORTUNER,R. & AMOUGOU,J. (1973). Tylenchorhyn- other plants in Thailand]. Entomol. Pl. Path. Bull, chusgladiolatus n.sp. (Nematoda : Tylenchida), Thailand, 1 : 1-15. nématode associé auxcultures du Sénégal et de Gambie. Cah. ORSTOM, Sb. Biol., no 21, 21-2.4. CHHABHA,H. K.,SAJJAN, S.S. & SINGH,H. (1974). Occurrence andcontrol of thestunt nematode FORTUNER,R. & JACQ,V. (1976). In uitro study of Tylenchorhynchusbrassicae Siddiqi, 1961 infesting toxicity of solublesulphides to t.hree nematodes rice nurseries in Punjab. Curr. Sci., 43 : 632. parasiticon rice in Senegal. Nematologica, 22 : 343-35 1. CHOI, Y. E. (1972). Astudy on theplant parasitic nematodes(Nematoda : Tylenchida)inKorea. FORTUNER,R. & MERNY, G. (1973). Les nématodes Korean J. Pl. Prot., 11 : 69-84. parasites des racines associés au riz en Basse Casa- mance (Sénégal) et en Gambie. Cah. ORSTOM, Sér. DAS, P. K. & RAO, Y. S. (1970). Life hist,oryand Biol., na 21 : 3-20. pathogenesis of Hoplolaimusindicus andits incidence in rice. Indian Phytopath, 23 : 459-464. FUKAZAWA,N., KOBAYASHI, Y. & NAKATA,M. (1963). [On therice root nematode Radopholus].Proc. DAS, P. K. & RAO, Y. S. (1971). On theoptimal Kansai Pl. Prot. Soc., 5 : 70-72. sampling time for assessment of nematode populations in rice soils. (Correspondance). Curr. Sci., 40 : GAGARIN,V. G. (1972). [On thenematode fauna of 17-18. perennialFar East rice in the Uchinskian water- DAS, V. M. (1960). St.udies on the nematode parasites reservoir]. (Abstr.).In Nematodnyeboleyni sel' of plantsin Hyderabad (Andhra Pradesh, India). skokhozyaistvennykhkul'tur i merybor'by s nimi. Terisy soveshchaniya Moskva, dekabr' 1972 : 48-49. 2. Parasitkde, 19 : 553-605. ELMILIGY,1. A. & GERAERT,E. (1971). Occurrence GATEVA,S. & PENTON,G. (1971). Faunade frtone- of someplant parasitic nematodes belonging to matodos en diferentes etapas de dos variedades de Tylenchida(Nematoda) Egyptin and Congo- arrozy fauna de malahierbas encontradas en el Kinshasa. Biol. Jaarb., 39 : 150-156. arrozal. CienciasAgropecuarias, serie 1, Ingeniera Agronomica, Cuba, 10 : 19 p. ENIBABI,M. S. (1967). Quantitativenematode-root relations in rice and Cotton. Diss. Abstr., 27 : 2569. GEMMA,T., SHIBUYA,T. & KIKUCHI,S. 1. (1964). FIELDING,M. J. (1956). Tylenchorhynchusmartini, a [Investigationon nematode parasitism in roots of new nematode species found in the sugar cane and riceplants and its injury]. YamagataNôrin Gak- rice fields of Louisiana and Texas. Proc.helminth. kaihô, 22 : 15-20. SOC.Wash., 23 2 47-48. GERMANI,G. & Luc, M. (1970). Contribution à l'étude FIGUEROA,A. & JIMENEZ,M. F. (1974). Infection and du genre Hemicriconemoides Chitwood & Birchfield, symptom induction in rice plants by the nematode 1957 (Nematoda : Criconematidae). Cah. ORSTOM, Hypsoperine n. sp. Phytopathology, 64 : 767-768. Sér. Biol., no 11 : 133-150. FORTUNER,R. (1973). Descriptionde Pratylenchus GERMANI.G. & Luc. M. (1973). Contribution à l'étude sefhensis_n^_sp,etdplolaim~~~~a~i~s~~~s~n~s~.d_u-ge&e Hemicuc1;ophora De Man, 1921 (Nematoda : -

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, Tylenchida)comportant la description de cinq nou- IMAMURA,S. (1931).Nematodes in the paddy field vellesespèces. Cah.ORSTOM, Sir. Biol., no 21 : withnotes on their population before and after 67-84. irrigation. J. Coll. Agric.imp. Univ. Tokyo, 11 : GOLDEN,A. M. & BIRCIIFIELD,W. (1968). Rice root 193-240. hot nematode (Meloidogyne graminicola) as a new INTERIANOMUROZ, J. D. (1970).Identification of Pest of rice. Pl. Dis. Reptr, 52 : 423. nematodes found in lands of the “Escuela Nacional GoTÔ, M. (1971). [Population fluctuations of therice deAgricultura” in El Salvador, and pathogenicity root nematodes in the Shônai District]. Comm. Ann. of ring nematode (Criconemoides spp) on rice (Oryza Meeting Japanese Soc. appl. E12to1nol. Zool., Fuchû, sativa L.) undergreenhouse conditions. Nematro- Japan, 7-9 April 1971 : 29. pica, 1 : 2-3. GoTÔ, M. (1973).[Invasion of riceroots in paddy ISHIICAWA,M. (1965).[Relation between fertilization field bythe riceroot nematode (Hirschmanniella and the rice root nematode]. Proc. Kantô Tôsan Pl. ima~nuri)] (Abstr.). Cornm. 17th Am. Meeting Prot.Soc., 12 : 115. Japanese Soc. appl. Entomol. Zool., Nagano, Japan, ISRAEL,P. & RAO, Y. S. (1971).Isolation of sources 3-5 April 1973 : 128. for nematoderesistance in ricc. SABRA0 News- HASHIOIIA,Y. (1964).Nematode diseases of ricein lefters, Mishima, 3 : 7-10. the world. Riso, 13 : 139-147. IVANOVA,T. S. (1968). [Nematodesof cereals from the HOOPER,D. J. & MERNY, G. (1966). Tworice nema- Seravshan Valley of Tadzhikistan] Dushambe, Izda- todesnew for Africa. Pl. Prot. Bull.F.A.O., 14 : telstvo LLDonish”,84 p. 25-26. IYATOMI,IL & NISHIZAWA, T. (1970). Growth response HOLLIS,J. P. (1967). Nature of the nematode problem of rice to soil fumigation .In: Toussoun, T.A., Bega, in Louisiana rice fields. Pl. Dis. Reptr, 51 : 167-169. R. V. & Nelson, P. E. (Eds) 1700t diseasesand soil bornepathogens. Berkeley,USA, University of HOLIJS,J. P. (1969 a). Chemical control of soil nema- California Press : 226-228. todesin rice fields. (Abstr.) Phytopathology, 59 : 1031. JACQ,V. & FORTUNER,R.(1979). Biological control of rice nematodes using sulphate reducing bacteria. HOLLIS,J. P. (1969 b). Gpzesis of a soil nematode Revue Nernatol., 2 : 41-50. problem in Louisiana rice. Il2t. Rice Comm. Newsl., 18 : 19-28. JENA,R. N. & RAO,Y. S. (1974). Root-knot nematode resistance in rice. Indian J. Genet. Pl. Breed., 34 A : HOLLIS,J. P. (1972). Nematicide-weeds interaction in 443-449. rice fields. Pl. Dis. Reptr, 56 : 420-424. JENA,R. N. & RAO,Y. S. (1976). Nature of root-knot HOLLIS,J. P. (1977). Loss in yield of rice causcd by (flfeloidogyne graminicola) resistance in rice (Oryza the ring nematode Criconelnoidesonoensis revealed sativa). 1; Isolation of resistantvarieties. Proc. by elimination of yellow nutsedge, Cyperus esculen- Indian Acad. Sci., 83 B : 177-184. tus. Nemafologica, 23 : 71-78. JOIINSTON,A. (1958). Diseases of paddy. Malay. IIOLLIS, J. P. & RODRIGUEZ-KABANA,R. (1966). agric. J., 41 : 10-17. Rapidkill of nematodesin flooded soil. Phytopa- JOIINSTON,T. M. (1958). The effect of soil moislure on thology, 56 : 1015-1019. Tylenchorhynchusmartini andother nematodes. I~OSHINO,M., YANAIIA,S., TAICITA,Y. & ODA,K. Proc. La Acad. Sci., 20 : 55-60. (1961). [On thecontrol of the ricecyst nematode JOI-INSTON,T. M. (1959). Effect of fatty acid mixlures of upland rice]. Proc.Kantd-Tôsan Pl. Pro€. Soc., on therice stylet nematode (Tylenchorhynchus ’ 8 : 74. rnarfini Fielding,1956). Nature,Lond., 183 : ‘1392. IBRAHIM,1. K. A., IBRAHIM,1. A. & REZIC,M. A. JOSIII,M.M. & HOLLIS,J. P. (1974).Pathogenicity (1972).Pathogenicity of certainparasitic nema- of Tylenchorhynchzzsmartini ontwo rice varieties. todes on rice. Alex. J. agric. Res., 20 : 175-181. (Abstr.). Proc. Amer. phytopath. Soc., 1 : 162. IBRAHIM,1. A., IBRAIIIM,1. K. A. & REZIC,M. A. JOSHI,M. M. & HOLLIS,J. P. (1976).Pathogcnicity (1973).Host parasite rolationship of Meloidogyne of Tylenchorhynchusmartini swarmerstorice. incognita’(Kofoid & White) Chitw. on rice. Nemafol. Nematologica, 2.2 : 123-124. medit. 1 : 8-14. JOSHI,M. M., IBRAHIM, 1. K. A. & 1-IOLLIS,J. P. (1975). ICIIINOIIE,M. (1955). [Twospecies of theroot knot Oxygen release from rice seedlings : effect of Meloi- nematodes in Japan]. Japanese J. appl. Zool., 20 : dogyneincognita and Helminfhosporiurnoryzae 75-82. . infections. ActaPhytopath. Acad. Sci. Hungar., ICIIINOHE,M. (1968). Prcsent status of research on the 10 : 51-53. rice-infestingnematodes in Japan. Rev. Pl. Prot. ICARIMOVA, S. M. (1972). [Investigation on nematodes Res.,Tokyo. 1 : 26-38. of rice and maize in the Uzbek SSR]. (Abstr.). In : ICIIINOHE,M. (1972). Nematode diseases of rice. In : Nematodnyebolezni sel’ skokhozyaistvennyldlkul’tur Webster, J. M. (Ed.) Economic nematology, London, i mery bor’ by s nimi. Tezisy soveshchaniya Moskva, U.K., Academic Press : 127-143. dekabr’ 1972. Tashkent State IJniv.

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KAWASHIMA, K. (1962). Ecologyof Radopholus oryzae LORDELLO,L. G. E.(1976). A semente do arroz dis- attacking rice roots. J. Plant Prot., Tokyo, 16 : 57- seminanematoides. RevtaAgric., Piracicaba, 51 : 59. 102. KAWASHIMA, K. (1963). [Investigationson Hirschman- Luc, M. (1957). Radopholus lavabri n. sp. (Nematoda : niaoryzae. 1. Varietalsusceptibility to the nema- Tylenchidae) parasite du riz au Cameroun FranGais. t.ode]. AnnualRept. Soc. Pl. Prot.North Japan, Nematologica, 2 : 144-148. 14 : III. Luc, M. (1958). Xiphinema. del'Ouest Africain : KAWASIIIMA,K. (1964 a). [Injuryto lowland rice descriptionde cinq nouvelles espkces (Nematoda : caused by Heterodera oryzae]. Annual Rept. Soc. Pl. Dorylaimidae). Nematologica, 3 : 57-72. . Prot.North Japan, 15 : 133-134. Luc, M. (1959 a). Nematodes parasites ou soupsonnés KAWASIII&rA, K.(1964 b). [Studies on Hirschmannia deparasitisme envers les plantesde Madagascar. oryzae. IV. On the soi1 reduction and the nematode Bull. Inst. Rech. agron. Madagascar, 3 : 89-102. injury]. AnnualRept. Soc. Pl. Prot.North Japan, Luc, M. (1959 b). NouveauxCriconematidae de la 15 : 131-132. zone intertropicale (Nematoda : Tylenchida). Nema- KAWASHIMA,K. & FUJINUMA,T. (1965). [On the tologica, 4 : 16-22. injuryto the rice plant caused by the rice root LUC, M. (1970). Contribution à l'étude du genre Cri- nematode (Hirschmanniaoryzae) : injuryto the conemoides Taylor, 1936 (Nematoda : Criconema- rice seedlings]. Res.Bull. Fukushima Pref. agric. tidae). Cah. ORSTOM, Sér. Biol., no 11 : 69-131. Exp. Stn., 1 : 57-84. Luc, M. (1973).Redescription de Xiphinemahallei KEGASAWA,K. & KAWASHIMA,K. (1962). [On the Luc, 1958 et descriptionde six nouvelles espèces geographical distribution of the rice root nematode de Xiphinema Cobb,1893 (Nematoda : Dorylai- Hirschmanniaorgzae (V.Breda de Haan)]. Proc. moidea). Cah. ORSTOM, Sr. Biol., no 21 : 45-65. Kantô Tôsan Pl. Prot. Soc., 9 : 72. Luc, M. & BERDONBRIZUELA, R. (1961). Heterodera KEOBOONRUENG,S. (1972). Effects of rice-root nema- oryzae n. sp. (Nematoda : Tylenchoidea)parasite tode, Hirschmanniellaoryzae (VanBreda de Haan du riz en Côte d'Ivoire. Nematologica, 6 : 372-279. 1902)Luc and Goodey,1963 on rice seedlings. Dissert. Abstr. infern., 32 B : 3729-3730. Luc, M. & GUIRAN,G. de (1960). Les nématodes asso- ciés aux plantes de l'Ouest Africain. Liste prélimi- KIIAN, A. 'ïVI. (1972).Studies on plantparasitic naire. 15 : 434-449. nematodes associated with vegetable crops in Uttar Agron. trop., Nogent, Pradesh. Final techn. Rep. ' Aligarh Muslim Univ., Luc, M. & GOODEY,B. (1962). Hirschmannia n. g. Botany Dept, 238 p. differenciated fronl Radopholus Thorne, 1949 (Nemn- KI-IEIRI, A. (1972). Plant. parasitic .nematodes (Tylen- toda : Tylenchoidea). Nematologica, 7 : 197-202. chida) from Iran.Biol. Jaarb. Dodonaea,40 : 222-239. LUC,M., MERNY, G. & NETSCHER,C. (1964). Enquête KIIERA,S. & CHATURVEDI,Y. (1970). On some tylen- sur les nématodes parasites des cultures de la Répu- c,hid nematodesfrom Orissa. Records zool. Survey bliqueCentrafricaine duet Congo-Brazzaville. India, 68 : 287-295. Agron.trop., Nogent, 19 : 723-746. KUMAZAWA,T. (1965).(Ecology and control of t,he MAAS, P. W. T., SANDERS,H. & DEDE, J. (1978). ricecyst nematode). Proc. KantôTôsan Pl. Prot. Meloidogyneoryzae n. sp. (Nematoda : Meloido- SOC.,12 : 6-8. gynidae) infesting irrigated rice in Surinam (South I

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MASLENNIKOVA, V. F. (1965 b). [Dynamics of the rasites en fonctionde l’inoculum. Cah. ORSTOM, nematode fauna of rice in the Tashkent and Fergan Sér. Biol., no 16 : 129-140. regions inthe Uzbelr SSP,]. Truclygel’mint. Lab., MERNY, G. & DEJARDIN,J. (1970).Les nématodes 1G : 68-74. ’ phytoparasites des rizières inondées deCôte d’Ivoire. MASLENNIKOVA,V. F. (1966).[Nematode fauna of II. Essai d’estimation de l’importance des popula- rice in the Uzbek SSR]. 2001.Zh., 45 : 641-645. tions. Cah. ORSTOM, Sr. Biol., no Il : 45-67. MATHUR,V. K., KHAN, E., NAND,S. & PRASAD,S. K. MERNY, G: & GERMANI,G. (1968). Tylenchorhynchus (1969). Two newspecies of Caloosia Siddiqi & palustris n. sp. (Nematoda : Tylenchinae), hôte des Goodey (Nematoda : Hemicycliophoridae)from rizières de Côte d’Ivoire. Annls Epiphyt., 19 : 601- India. Bull. Ent. Loyola Coll., 10 : 27-31. 603. MATHUR,V. K., KHAN,E. & PRASAD,S. K. (1966). MIURA, II. & SHÔJI, H. (1964). [On thedistribution Boleodoroidesoryzae n. g., 11. sp.(Nematoda : of the rice root nematode in Yamagata prefecture]. Boleodorinae) from Bihar, India. Nematologica, 12 : Annual Rept Soc. Pl. Prot. North Japan, 15 : 132- 448-452. 133. MATIIUR, V. K. & PRASAD,S. K. (1971).Occurrence MOMOTA, Y. & 01-ISHIMA,Y. (1973). [Ring nematode anddistribution of Hirschmanniellaoryzae inthe detectedfrom continuouslya cropped field of Indian Union with description of H. mangaloriensis upland rice]. (Abstr.). 17th Annual Meeting Japanese n. sp. Indian J. Nematol., 1 ; 220-226. Soc. appliedEntomol. Zool., Nagano,Japan, 3-5 MATIIUR, V. K. & PRASAD,S. K. (1972 a). Embryonic April 1973, 129. developmentand morphology of larvalstages of MONTEIRO, A.R. (1968).Ocorrência no Brasil de therice root nematode, Hirschlnanniellaoryzae. importante nematoide fitoparasito. O Solo, 60 : 81. Indian J. Nematol., 2 : 146-157. NAKAZATO,H., KAWASIIIMA, K. & KUROSAWA,T. ’ MATHUR, V. K. JZ PRASAD,S. K. (1972 b). Role of (1964). [Seasonal occurrence of the rice root nema- th,erice root nematode Hirschmanniellaoryzae in tode]. Proc. Kantô-Tôsan Pl. Prot. Soc., 11 : 106. rice culture. Indian J. Nematol., 2 : 158-168. NANDAIWMAR,C. & RAO,Y. S. (1974). On the migra- MATHUR, V. K. & PRASAD,S. K. (1973 a). Survival tory behavior of some subterranean parasitic nema- and host range of the rice root nematode, Hirsch- todesto aerial part of riceplants. Nematologica, Inanniella oryzae. Indian J. Nematol., 3 : 88-93. 20 : 106. MATIIUR, V. K. & PRASAD,S. K. (1973 b). Control of Hirschmanniellaoryme associatedwith paddy. NETSCHER,C. (1969). L’ovogénhse et la reproduction H. : Indian J. Nematol., 3 : 54-60. chez Heteroderaoryzae et sacchari (Nematoda Heteroderidae). Nematologica, 15 : 10-14. MERNY, G. (1966 a). Biologie d’Heterodera oryzae Luc & Berdon, 1961. II. Rôle des masses d’œufs dans la NISHIZAWA, T., SHIMIZU,K. & NAGASIIIMA, T. (1972). dynamiquc des populations et la survie de l’espèce. [Chemical andcultural control of therice cyst ~nnls.Epiprzyt., 17 : 445-449. nematode Heterodera oryzae Luc & Berdon-Brizuela and hatching responses of the larvae to some root MERNY, G. (1966 b). Nématodes d’Afrique Tropicale : extracts]. Japanese J. Nematol., 2 : 27-32. unnouveau Paratylenchus (Criconematidae),deux nouveaux Longidorus et observations surLongidorus OD’A,K., HOSIIINO,M., YANAKA,S. & KUMAZAWA, laevicapitatus Williams, 1959 (Dorylaimidae). Nerna- T. (1963).[Studies on therice cyst nematode of tologica, 12 : 385-395. upland ricg. III. Analysis of the injury]. Proc. Kantô Tôsan Pl. Prot. Soc., 10 : 71. MERNY, G. (1970 a). Lesnématodes phytoparasites . desrizières inondées de Côte d’Ivoire. 1. Lesespèces OIISHIMA, Y. (1974). Heteroderaelachista n. sp. an observées. Cah. ORSTOM, Sér. Biol., no 11 : 3-43. uplandrice cyst nematode from Japan. Japanese MERNY, G. (1970 b). Loide croissance, sur plant de J. Nematol., 4 : 51-56. . riz, d’une population endophyte d’Hirschmanniella OKADA,T. (1955). [Onmorphological characters of spinicaudata (Nematoda : Tylenchoidea) enfonc- the upland rice nematode Heterodera sp.]. (Abstr.). tiond’un inoculum variable. Nematologica, 16 : Annual Meeting applied2001. entomol. Soc. Japarz,11. 227-234. OOSTENBRINR,M. (1966). Major characteristics in the MERNY, G. (1972 a). Les nématodesphytoparasites relationbetween nematodes and plants. Med. desrizières inondées de Côte d’Ivoire. III. Etudes LandbHogesch.Wageningen, 66 : 1-45. surla dynamique des populations de deux endoparasites : Hirschmanniellaspinicaudata et Hete- OTEIFA,B. A. (1962). Species of root-lesion nematodes roderaoryzae. Cah. ORSTOM, Sér. Biol., no 16 : commonlyassociated with economiccrops inthe 31-87. Delta of the U.A.R. Pl. Dis. Reptr, 46 : 572-575. MERNY, G. (1972 6). Lesnématodes phytoparasites Ou, S. H. (1965).Rice diseases of obscure nature in desrizières inondées de Côte d’Ivoire. IV. Essai Tropical Asia withspecial reference to “Mentek” d’interprétationmathématique de l’intensité de diseaseinIndonesia. Int. Rice Commn Newsl., l’infestation des racines par les nématodes endopa- 14 : 4-10.

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~ ~ ~~~~ ______- ~

Ou, S. H. (1972). Rice Diseases. Kew Commonwealth RAO,Y. S. & ISRAEL,P. (1971 b). Studies on nema- Mycological Institute, XIS + 368 p. todes of riceand rice soils. VI.Influence of soil PANDA,M. & RAO,Y. S. (1971). Evaluation of losses temperature on theprevalence and incidence of caused bythe rootnematode (Hirschmanniella Meloidogynegraminicola, the rice root-knot nema- mucronata Das) in rice (Oryza sativa L.). Indian J. tode. Oryza, 8 : 47-51. agric. Sci., 41 : 611-614. RAO,Y. S. & ISRAEL,P. (1972 a). Effect of temperature PARK,J. S., HAN,S. C. & LEE, Y. B. (1970). [Studies on hatching of e.ggs of the rice root-knot nematode on the ecology and feeding preferenceof Hirschman- Meloidogyne graminicola. Oryza, 9 : 73-75. niella oryzae]. Research Reports of the Office of Rural RAO,Y. S. & ISRAEL,P. (1972 b). Spatial distribution Development, Korea, Plant Environment, 13 : 93-98. of larvae of Meloidogyne graminicola, the rice roob PATNAIK,N. C. (1969). Pathogenicity of Meloidogyne knot nematode. Oryza, 9 : 87-91. graminicola (Golden & Bjrchfield, 1965) in rice. RAO,Y. S. & ISRAEL,P. (1972 c).'Influenceof inoculum (Abstr.). Al1 IndiaNelnatology Symposium, Neur density on the final populationof root knot nematode Delhi, 21-22 Aug. 1969: 12. (Meloidogyne graminicola) in rice. Indian J. Nema- PRASAD,K. S. & RAO,Y. S. (1973). Chemotherapy of fol., 2 : 72-76. the rice plant against the infestation by the stunt RAO,Y. S. & ISRAEL,P. (1972 d). Influence of soil nematode (Tylenchorhynchus sp.). Indian J. Nema- type on the activity of the rice root knot. nematode tol., 3 : 80. Meloidogyne graminicola Golden & Rirchfield. Indian PRASAD, K.S. K. & RAO,Y. S. (1976 a). Chemothe- J. agric. Sci., 42 : 744-747. rapy of theroot-knot nematode Meloidogynegra- RAO,Y. S. & JAYAPRAKASII,A. (1977). Leaf chlorosis minicola in rice. 1. Effect.s of soakingseeds in due to infestationby a new cyst. nematode. Int. nematicidesolutions. 2. PflEirankh.PflPath. Pfl- Rice Commn Newsl., 2 : 5. Schut~,83 : 665-668. RAO, Y. S. & PRASAD,J. S. (1977).Root lesion PRASAD,K. S. K. & RAO,Y. S. (1976 b). Chemothe- nematode damage in upland rice. In€. Rice Commn rapy of theroot-knot nematode (Meloidogyne Newsl., 2 : 6-7. graminicola) in rice. II. Effects of root-dip treatment REVERSAT,G. (1975 a). Etude préliminaire dela survie withnematicides oninvasion and development of en anaérobiose des juvéniles du nématodeHeterodera thenematode. 2. PflKrankh.PflPath. PflSchutz, oryzae (Tylenchida : Ileteroderidae). C.r. hebd. 83 : 730-735. Seanc. Acad. Sci., Paris, 280, série D : 2865-2868. PRASAD,K. J. K. & RAO,Y. S. (1976 c). Chemothe- REVERSAT,G. (1975 b). Méthodes pour lamesure de rapy of theroot-knot nematode (Meloidogyne la consommationd'oxygène des némat.odes par la graminicola) inrice. III. Evaluation of pesticides t.echnique duludion. Cah.ORSTOM, Se'r. Biol., as soildrench. 2. PflKrankh.PflPath. PflSchutz, 169-187.10 83 : 736-741. REVERSAT,G. & MERNY, G. (1973). Influence de quel- RAMANA,K. V., MATIIUR, S. C. & RAO,Y. S. (1974). quesfacteurs sur lapénétration du nématode Role of Hoplo1aim.u.s indicus on theseverity of Heteroderaoryzae dansles racines du riz. Cah. seedling blight of rice. (Correspondance). Curr. Sci., ORSTOM, Se'r. Biol., no 21 : 111-115. 43 : 687-688. REYES,R. D. (1971).Determinacion de la eficiencia RAO, V. N. (1972).0cc.urrence of Hirschuzanniella de diferentes variedades de arroz y de maiz como mucronata on riceleaves. Indian J. Nematol., 2. : hospederasde Pratylenchuszeae. In : Progreso de 214-215. LaboresdeInvestigaciones Agropecuarias, 1970. RAO,Y. S. (1970). Study of plant parasitic nematodes Panama,Facultad de Agronomia, Panama Uni- affecting ricc production in the vicinity of Cuttack versidad : 159-166. (Orissa),India. Finaltechnical report, Indian RODRIGUEZ-KABANA,JORDAN,R., J. W. & HOLLIS, Council of Agricultural Research, 115 p. J. P. (1965). Nematodes : biological control in rice RAO,Y. S., & BISWAS, H. (1973). Evaluation of yield fields : role of hydrogen sulfide. Science, N.Y., losses in rice due to the root-knot nematode Meloi- 148 : 524-526. dogyne incognita. Indian J. Nemafol., 3 : 74. ROY, A. K. (1973).Reaction of somerice cultrivars RAO,Y. S., BISWAS,H., PANDA,M., RAO,P. R. V. J. to the attack of Meloidogyne graminicola. Indian J. & RAO,V. N. (1969).Screening rice varieties for Nematol., 3 : 72-73. theirreaction to root and root knot nematodes. ROY, A. K. (1975).Studies on resistance of rice to (Abstr.) Al1 IndiaNematology Symposium, New theatt

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Accepte' pour publication le 25 octobre 1978.

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102 Revue Nématol: 2 (1) : 79-102 (1979)