cyst nematodes in equatorial and hot tropical regionsfrom: cyst nemat'qdes edited by f....

From: CYST NEMAT'QDES Edited by F. Lamberti and C, E. Tayfor (Plenum Publishing Corporation, 1986)

CYST NEMATODES I N EQUATORIAL AND HOT TROPICAL REGIONS

Michel Luc I Musiehm n a t i o n a l d ' H i s t o i r e n a t u r e l l e Labora to i re des Vers, 61 rue de Buffon, 75005 P a r i s France

INTRODUCTION

In the p re sen t communication, t h e term "cys t nematodes i n e q u a t o r i a l and t r o p i c a l reg ionsf t i s used in s t ead of t h e usua l " t r o p i c a l c y s t nematodes" f o r two reasons :

- i n add i t ion to those spec ie s of c y s t nematodes t h a t are c l o s e l y assoc ia ted wi th h o t t r o p i c a l crops and a r e a s , t h e r e are o t h e r s t h a t a r e common i n temperate a r e a s which may a l s o be occas iona l ly found i n t he t r o p i c s ;

- t h e topographical meaning of " i n t e r t r o p i c a l " encompasses some mountainous a r e a s , namely i n Cent ra l and South America, where c l ima te , vege ta t ion and crops are q u i t e similar to those of temperate r eg ions : t hese a r e the "cold t ropics" .

This paper dea l s on ly ,wi th t h e "hot t rop ic s" , and the t e r m " t rop ic s" and f%ropica l f l r e f e r here only t o these c l i m a t i c regions.

Regarding the earliest r eco rds of %"teroderart i n e q u a t o r i a l and t r o p i c a l a r e a s , i t must be kep t i n mind t h a t p r i o r t o Chitwood's (1949) r e s u r r e c t i o n of t h e anc ien t gener ic name MeZoidogyne Goeldi, 1877, t h e genus Hetmodera contained both c y s t forming spec ies of Heteroderidae ( o r Heterodera sensu l a t o ) and var ious r o o t knot spec ies under the name Heterodera marioni (Cornu). i n t he t rop ic s were given wi thout morphological d e t a i l o r d e s c r i p t i o n of symptoms on the h o s t p l a n t , i t i s n o t poss ib l e to determine whether t he spec ies i n ques t ion were MeZoidogyne o r cyst-forming spec ies .

A s most of t h e e a r l y records of "HeterOdt?ra"

Also, t he re a r e numerous records of IlHeterodera-like juven i l e s " i n t h e l i t e r a t u r e concerning t r o p i c a l crops bu t such juveni les may belong t o spec ies of non-cyst forming Heteroderinae genera which a l s o occur i n the t rop ic s . For example, i n the Ivory Coast Heterodera-like juven i l e s were f r equen t ly found i n some a reas of primary o r anc ien t f o r e s t s bu t ex tens ive inves t iga t ion f a i l e d t o f ind c y s t s . Eventually it w a s discovered t h a t these juven i l e s belonged t o a non-cyst forming spec ies of Heteroderinae p r e s e n t on some t r e e s and t h a t they represented a new genus and spec ie s , subsequently described under the name of HyZonema ivorense by Luc e t a l . (1978). Therefore records only of TTHe$eroderatl j ____- - -- - _ _ __ --

Q O 10025052

356 M. LUC

The f i r s t r e p o r t of a c y s t nematode a t t ack ing a t r o p i c a l crop i n a t r o p i c a l a r e a concerns sugar cane i n the Hawaiian I s lands (Muir and Henderson, 1926). because the au thors s t a t e d "It is only r e c e n t l y t h a t the two spec ie s * [of ItHeterodera"] have both been recognised i n Hawaiian Islandsll. A d e s c r i p t i o n i s given of t he damage symptoms caused t o t h e h o s t p l a n t by the "two spec ies" and the d i s t i n c t i o n between Heterodera and MeZoidogyne i s c l e a r l y s t a t e d . Concerning Heterodera they say " In sugar cane t h i s nemaode does n d t show a preference f o r t he t i p of r o o t s bu t a t t a c k s them a t any po in t ; ... i t does no t produce a g a l l on the root". c i t e d .

It i s c l e a r t h a t t he re was no confusion with MeZoidogyne

The "brown c y s t stagell is a l s o

Muir and Swezey (1926) a l s o recorded IlHeterodera schachtiill on pine- app le , i n t he Hawaiian I s l ands , bu t here t h e t e x t i s no t a s c l e a r and doubts remain. These two records a r e c i t e d aga in i n a l a t e r paper by Henderson (1926).

The r e p o r t 6f a c y s t nematode on sugar cane was no t considered s e r i o u s l y by subsequent au thors because of the p reva i l i ng thought t h a t c y s t nematodes were t o be found only i n temperate a reas . For example, i n t he compilation by Goodey (19561, t h i s record i s c i t e d as 1IHet. schacht i i = ? Mel. sp . l t .

This ItHeterodera" has n o t been r epor t ed again in Hawaiian sugar cane p l a n t a t i o n s and thus the spec ie s , and perhaps the genus, remain unknown. However, I consider t h a t it probably was n o t H . sehacht ic , as sugar cane has been shown t o be a non-host f o r t h i s spec ie s (Skarbilovich and Kurramov, 1473).

Th i r ty f i v e years l a t e r , Luc (1961) repor ted two t r o p i c a l Heterodera s p e c i e s , one p a r a s i t i s i n g swamp r i c e i n t h e Ivory Coast and the o t h e r sugar cane i n the Congo. They were described as H . oryzae Luc & Berdon - Br izue la , 1961 and H . sacehari Luc & Merny, 1963 and t h i s c l e a r l y e s t ab l i shed t h a t c y s t nematodes could occur i n the t rop ic s . Subsequently s e v e r a l o the r spec ies have been described from t r o p i c a l crops and weeds. Never the less , it appears t h a t t h e cyst-forming nematodes a r e not as common nor as numerous, and probably n o t a s economically important, i n t h e t r o p i c s as they are i n temperate a reas .

GENUS HETERODERA Schmidt, 1871

Heterodera oryzae Luc & Berdon-Brizuela, 1961

H . oryzae w a s repor ted from swamp rice f i e l d s a t Bokakouamekro i n the c e n t r a l p a r t of the Ivory Coast (Luc and Berdon-Brizuela, 1981). This s p e c i e s , of t he schacht i i group, i s cha rac t e r i s ed mainly by the presence of a voluminous egg-mass, a t t ached t o t h e c y s t , which o f t e n conta ins more eggs than the c y s t i t s e l f . The o r i g i n a l d e s c r i p t i o n has been supplemented by Luc and Taylor (1977).

*Prior to t he beginning of 1928, only two spec ies of IIHeterodera~l were recognised: H . sdkacht i i Schmidt ( the cyst-forming spec ie s ) and H . marioni (Cornu) ( t h e r o o t knot spec ie s ) . A t h i r d spec ie s , H . javanieu Treub, 1885, a l s o e x i s t e d bu t a s it was described from an exo t i c h o s t p l a n t ( sugar cane) and from a t r o p i c a l country (Indonesia) l i t t l e a t t e n t i o n w a s given to it a t t h a t t i m e .

,. . . r

CYST NEMATODES IN EQUATORIAL AND HOT TROPICAL REGIONS 357

The r o l e of t he egg-mass i n r e l a t i o n t o population dynamics has been s tudied (Merny, 1966; Merny, 1972a, Reversa t , 1975d). Juveni les i n the egg- mass ha tch immediately without need of a s t imu la t ion by r o o t exudates and t h i s enables two o r t h ree genera t ions t o develop on the same r i c e p l a n t (Merny, 1972a). This ready ha tch ing of egg-masses f a c i l i t a t e s t h e c u l t u r e of the nematode, even under monoxenic condi t ions (Reversat , 1975a). Thus H. oryzae has been used t o examine var ious aspec ts of the biology of Heterodera such as ovigenesis and reproduct ion , which does no t d i f f e r from other amphimictic spec ies of t h e genus (Netscher, 1969); h i s topa thology of a t tacked roo t s (Berdon-Brizuela and Merny, 1964); f a c t o r s in f luenc ing the sex r a t i o and sex determination (Cadet e t a l . , 1975; Cadet and Merny, 1978a); gene t ic f a c t o r s r e l a t e d t o the length of j uven i l e s (Netscher and Pernes , 1971) ; production of a r t i f i c i a l mutants (Cadet, 1976) ; f a c t o r s in f luenc ing pene t r a t ion by the juven i l e s (Reversat and Merny, 1973; Cadet and Merny, 1976; Merny and Cadet, 1978); chemical composition of j uven i l e s (Reversa t , 1977a); oxygen uptake by j u v e n i l e s (Reversat , 1975c; 1977b) and t h e i r su rv iva l (Reversat , 19801, p a r t i c u l a r l y i n anaerobic condi t ions (Reversat , 197513).

The spec ies i s known only from some swamp a reas i n the Ivory Coast (Merny, 19701, and i n Senegal where it p a r a s i t i s e s banana t r e e s (Taylor , 1978). Reports of H . oryzae from Japan (Hashioka, 1964; Shimizu, 1971, 1972, 1973; Nishizawa e t a l . , 1972) r e f e r t o a d i f f e r e n t spec ie s , descr ibed by Ohshima (1974) as H . e lachis ta . However, t he two spec ies appear t o be c lose ly r e l a t e d and t h e i r r e l a t i o n s h i p s r equ i r e f u r t h e r study.

I n add i t ion t o r i c e and banana, few o the r p l a n t s have been shown t o be hos ts . Maize and Mariscus wnbellatus a r e suscep t ib l e (Merny and Cadet, 1978) whereas sugar cane, Sorghum uuZgare, Pennisetwn typhoides and Panicum maximum a r e n o t , and l a rge d i f f e rences between var ious r i c e c u l t i v a r s have been observed (Merny and Cadet, 1978). could expla in the l imi ted d i s t r i b u t i o n of t h e spec ies . I t is a l s o l imi t ed i n flooded r i c e f i e l d s by competit ion with H. sacchari; experiments wi th juven i l e s marked with 35 have shown t h a t H. oryzae i s much l e s s aggress ive than H. sacchari (Reversat and Bois , 1982).

This r e l a t i v e l y narrow h o s t range

s

Heterodera sacchari Luc & Merny, 1963

H. sacchari, of the schacht i i group, w a s o r ig ina sugar cane ' in the Niari Val ley , Congo (Luc and Merny, subseqúently been repor ted from t h e Ivory Coast where been found i n flooded r i c e f i e l d s i n the nor thern and

l y described from 1963). I t has it has f r equen t ly c e n t r a l reg ions

and wi ld g ra s ses (Merny, 19701, i n Niger ia on sugar cane ( J e r a t h , 1968 (Odih i r in , 19751, i n flooded r i c e f i e l d s of Casamance Province, Senegal and i n Gambia (Fortuner and Merny, 1973) and on sugar cane i n Burkina Faso (Cadet and Merny, 1978b). The spec ie s is a l s o repor ted t o have been found on Saccharum spontaneum i n Ind ia (Swarup e t a l . , 1964).

I n addi t ion t o sugar cane and r i c e , var ious wild Graminaceae a r e considered a s hos t s by Odih i r in (1975): PaspaZum eonjugatum, Axonopus compressus, Mariscus umbellatus, Cynodon dactylon, Eleusine indica and Brachiaria brizantha. Odihi r in (1975) considers t h a t H. sacchmi i s indigenous i n Niger ia and w h i l s t I agree I a l s o th ink t h a t t h e a rea of d i s t r i b u t i o n is probably wider than ind ica ted by present r epor t s .

The morphology of H. sacchari has been described and t h e o r i g i n a l desc r ip t ion was supplemented by Luc (1974). The major d i agnos t i c charac te r of the spec ies i s a very dark , s t rong ly developed underbridge

358 M. LUC

of the female c y s t which has f inge r - l i ke p ro jec t ions . The apparent ly non- func t iona l male has been descr ibed by Netscher e t al . (1969). (1969) has shown t h a t H . sacchari i s a t r i p l o i d par thenogenet ic spec ies .

Netscher

The hatching of juveni les from c y s t s i s similar i n sugar cane r o o t d i f f u s a t e and tap water (GarabKdian and Hague, 1984) but Riversa t (1981) has found potassium permangate t o be u s e f u l as a hatching agent. Aldicarb, oxamyl and carbofuran prevent hatching from c y s t s but t h i s e f f e c t i s decreased by the addi t ion of sugar cane r o o t d i f f u s a t e (Garabedian and Hague, 1984). 8

J e r a t h (1968) observed t h a t i n fe s t ed sugar cane p l a n t s a r e s tun ted and t h i n , and secondary roo t s a r e less abundant than hea l thy p lan ts . In sugar cane p l a n t a t i o n s , c y s t s of H. sacchari may be t ranspor ted by the water i n i r r i g a t i o n cana ls f o r a t l e a s t 5 to 8 km (Odih i r in , 1977).

i

On r i c e , experiments show (Babatola , 1983a) t h a t i n fe s t ed p l a n t s a r e c h l o r o t i c and t h e i r growth r e t a rded , r o o t s are nec ro t i c and blackened, t i l l e r numbers a r e reduced and the g r a i n y i e ld i s lower. H. sacchari a t t a c k s both swamp and upland r i c e b u t the l a t te r appears more suscep t ib l e to damage (Babatola , 198313). Rice c u l t i v a r s r e a c t very d i f f e r e n t l y . t o in fe s t a - t i o n by the nematode (Babatola , 1983b).

I n western areas of Af r i ca , H. sacchari may c o n s t i t u t e a danger f o r sugar cane, and both upland and swamp r ice . F o l i a r app l i ca t ions of oxamyl have been shown experimental ly t o prevent invasion of young sugar cane p l a n t s (Garabedian and Hague, 1982) but f i e l d tr ials have no t been reported.

H. cyperi Golden, Rau & Cobb, 1963

H. cyperi was f i r s t found i n f e s t i n g Cyperms escuZentus a t Sanford, F l o r i d a , USA, and w a s descr ibed by Golden e t a l . (1963). known and i t s d i s t r i b u t i o n w a s thought t o be l imi ted to two coun t r i e s i n F lo r ida and one i n Gèorgia (Mulvey and Golden, 1983). Recently, however, Kumar (1980) recorded it on Cynodon dactyZon and Bambusa sp. i n a cof fee e s t a t e , i n Karnataka, India . Eroshenko and Kazachenko (1972) considere'd it t o be a spec ies inquirenda, but t he re does not seem s u f f i c i e n t reason f o r t h i s .

No o the r hos t was

Heterodera mothi Khan & Husain, 1965

H. mothi; found on Cypems rotundus, w a s t he f i r s t c y s t nematode Cysts of t h i s spec ie s , descr ibed from India (Khan and Husain, 1965).

which is of the goettingiana group, a r e cha rac t e r i s ed by the th inness of t h e i r c u t i c l e , the prominence of the a n a l a r e a , and the . s l ende rness of the vu lva l br idge. b r idge i s r a r e l y present . An egg-sac has been observed (Khan and J a i r a j p u r i , 1975).

Mulvey (1972) emphasised the f a c t t h a t the under-

Sharma and Swarup (1984) s t a t e t h a t t h i s spec ies reproduces only on the type h o s t whereas o ther records a r e from Chrysanthemum carinatum and Hibiscus syria'cus i n India (Yaqoob and Khan, 1969); Cyperus escuZentus, GZyCine m u , GOSSypiUm hirsutum, i n Georgia, USA (Minton e t a l . , 1973); i n s o i l from sugar-beet f i e l d s i n I r an (Tala t sch ian and Achyani, 1976); sugar cane (Maqbool, 1981) and onion ,(Maqbool e t a l . , 1983) i n Pakis tan ; and c y s t s i n s o i l a t V i t i Levu, F i j i I s l ands (Kirby e t a l . , 1978).

',


Sharma and Swarup (1984) gave information on the morphometrics and l i f e cyc le of H . mothi but nothing i s known about i t s pathogenici ty .

H . indocyperi Husain & Khan fi Khan e t a l . , 1964 ( c i t e d by Hesl ing, 1978) i s considered by Hesling t o be a nomen nudum and a synonym of H . mothi.

Heterodera cajani Koshy, 1967 (= H . V igni Edward & Misra, 1968)

This spec ie s , of the schacht i i group, was f i r s t reported under the name of H . t r i f o z i i i n a pigeon-pea f i e l d of t he I A R I a t New Delhi , Ind ia (Swarup e t a l . , 1964). It w a s descr ibed l a t e r as H. cajani by Koshy (1967). H . v i g n i descr ibed from r o o t s of Vigna unguicuzata by Edward and Misra (1968) has been recognised (Kalha and Edward, 1979) as a jun io r synonym of H . cajani.

H. cajani is the most widespread cyst-forming nematode i n India and has been reported i n the s t a t e s of Andhra Pradesh, Bihar , Delhi , Haryana, Rajasthan. Tamil Nadu and Uttar Pradesh (Sharma and Swarup, 1984). The . . spec ie s has a l s o been repor ted on cowpea i n Egypt (Aboul-Eid and Ghorab, 1974).

i'

On the bas i s of f i e l d observa t ions and experiments by var ious au thors (Koshy, 1967; Koshy and Swarup, 1972; B h a t t i and Gupta, 1973; Janarthanan, 1972; Verma and Yadav, 1975; Gaur and Sing, 1977) the following h o s t p l a n t s have been e s t ab l i shed : Cajanus cajan, Cicer arietinwn, Cyamopsis te t ra- gonoloba, DoZichos bifZorus, D. ZabZab, GZycine max, PhaseoZus atro- purpureus, P. aureus, P. caZcaratus, P. Zathyroides, P. Zunatus, P. mungo, P. vulgar is , P i s w n sativwn, Sesamwn indicum, S. orientaze, V i v i a narabenesis cv. sa t iva , Vigna s inens is and Zea mays. t he l as t spec ie s , a l l these h o s t p l a n t s belong t o Leguminosae o r Pedal iaceae. Koshy and Swarup (1972) e s t ab l i shed t h a t 82 crop and weed spec ie s were non-hosts including Cyamopsis tetragonoloba, GZycine max, Phaseozus CaZcaratus, P i swn sativwn cv. Ear ly Wisconsin and seve ra l V i c i a spec ie s . Koshy e t a l . (1971) and Sharma and Swarup (1984). The biology of H . cajani i s s i m i l a r t o o the r Heterodera spec ies . Host r eac t ions have been descr ibed by Aboul-Eid and Ghorab (1974) and the l i f e cyc le by Koshy and Swarup (1971a.l. More than nine generat ions per year have been repor ted (Koshy and Swarup, 1971b). The hatching of j uven i l e s from cys t s and egg-masses i s opt imal a t 29°C; r o o t leacha tes of Cajanus cajan and DoZichos ZabZab have a p o s i t i v e e f f e c t ; hatching of j uven i l e s from egg-masses a t 20 days i s 100 pe r cen t i n c o n t r a s t t o only 16 per cen t from brown c y s t s (Koshy and Swarup, 1971d). The presence of egg-masses, and the rap id hatching of j u v e n i l e s , expla ins the high number of genera t ions observed. I t has a l s o been observed t h a t H. cajani may a t t a c k the a e r i a l p a r t s of young seedl ings of pigeon-pea (Koshy and Swarup, 1971e).

With the exception. of

The o r ig ina1 ,desc r ip t ion of the spec ies has been supplemented by

Although the re i s no p rec i se da t a on the damage t o crop p l an t s caused by H. cajani , it must be considered a dangerous spec ies . Its presence i n Egypt shows t h a t it could become es t ab l i shed i n t r o p i c a l , sub t rop ica l and even Mediterranean count r ies .

Heterodera graminis Stynes, 1971

H . graminis w a s descr ibed from r o o t s of Cynodon daetyzon on a bowling green a t New Castle, New South Wales, A u s t r a l i a (Stynes, 1971). spec ie s has r ecen t ly been recorded on undetermined spec ies of grasses on

The

360 M. LUC

a gol f course a t Delh i , Rajasthan, Ind ia (Sharma and Swarup, 1984) and has been found i n s o i l a t Vanua Levu, F i j i I s lands (Kirby e t a l . 1978) and on Cynodon dactylon i n a sugar cane p l an ta t ion i n Trinidad (Stone, i n l i t t . ) .

H . graminis is of t he goe t t inghna group, and appears t o be c lose ly r e l a t e d t o H . cyperi.

In po t experiments H . graminis reproduced on Setaria i t a l i c a , Secale cereale and maize (Sharma and Swarup, 1984). Nothing i s known about i t s pa thogenic i ty but h o s t p l an t s w e r e descr ibed as yellowish and s tun ted (Kirby e t a l . , 1978; Sharma and Swarup, 1984). Data on t h e l i f e c y c l e a r e given by Sharma and Swarup (1984).

i Heterodera zeae Koshy, Swarup & S e t h i , 1971

This spec ies of t h e schacht i i group w a s described from specimens found i n a s soc ia t ion wi th maize a t Chapl i , Rajasthan, Ind ia (Koshy e t a l . , 1971). Surveys showed t h a t H . zeae i s widely d i s t r i b u t e d throughout India and i n t h e following s t a t e s : Andhra Pradesh, Bihar , Delh i , Gujara t , Madhya Pradesh, Punjab, Rajasthan, Uttar Pradesh (Koshy and Swarup, 1 9 7 1 ~ ) and Karnataka (Narayanaswamy, e t a l . , 1982). Subsequently i t was recorded i n Pakis tan (Maqbool, 1981), i n Egypt (Ote i f a , unpublished r e p o r t , 19781, and i n the USA (Sa rdane l l i e t a l . , 1981). H . zeae i s considered t o be one of t he most economically important nematodes a t t ack ing c e r e a l s i n Ind ia (Sharma and Swarup, 1984).

In add i t ion t o the o r i g i n a l desc r ip t ion , d a t a on morphology and morphometries have been repor ted from Indian populations by Sharma and Swarup (1984) and f o r American popula t ions by Golden and Mulvey (1983). Hutzel1 (1984) has r e c e n t l y descr ibed t h e m a l e . provide some information on the biology of t he spec ies b u t t he re has been no published information on pa thogenic i ty . Hosts recorded f o r H. zeae inc lude seve ra l c u l t i v a r s of Zea mays, (Misra e t a l . , 19771, Echinochloa cozona, Panicum spp., Setaria italics and Tri t icum aestivum. (Sr ivas tava and Swarup, 1975; Sharma and Swarup, 1984) t h e following p l a n t s have been found suscep t ib l e t o H . zeae: Oryza sa t iva; Secale cereale , Sorghum vulgare, Zea mexicana, Avena sa t ivaand Hordeum vulgare. vulgare w a s , however, considered r e s i s t a n t by Sr ivas tava and Swarup (1975) a s were Paspaluni spp. and Pennisetuni typhoidem. Maqbool (1981) r epor t ed non-graminaceous c rops (gram, c i t r u s , pea r , g a r l i c ) a s h o s t s , bu t it i s n o t c l e a r i f c y s t s were found on t h e r o o t s of these p l an t s o r i n the surrounding s o i l .

Sharma and Swarup (1984)

Experimentally

Sorghum

Heterodera gaembiensis Merny & Netscher, 1976

Heterodera gambiensis, belonging t o the schacht i i group, has been repor ted on Sorghum vulgare and Pennisetum typhoideum i n var ious loca t ions i n Gambia, West Af r i ca (Merny and Netscher, 1976). Su rp r i s ing ly , i n the b

Senegalese reg ions surrounding Gambia, ex tens ive surveys f a i l e d t o l o c a t e t h i s spec ies . It has , however, r e c e n t l y been recorded on Eleusine caracana i n Karnataka, Ind ia (Narayaswamy e t a l . , 1982). There i s no information on the biology, hos t range o r pa thogenic i ty of the spec ies .

Heterodera oryzicoza Rao &,Jayaprakash , 1978

This spec ies of t he schacht i i group is known only from upland r i c e i n Kerala S t a t e , Ind ia (Ra0 and Jayaprakash, 1978). Although it has been observed t o pene t r a t e t he r o o t s of s eve ra l graminaceous p l a n t s , according

CYST NEMATODES IN EQUATORIAL AND HOT TROPICAL REGIONS 36 1

t o Sharma and Swarup (1984) cys t s are produced only on r i c e , mainly on the c u l t i v a r s J aya , Ratna and T (N)-l.

H . oryzieoZa causes s tun t ing and ch lo ros i s of leaves (Rao and Jayaprakash, 1977) and f i e l d experiments i nd ica t e y i e ld lo s ses of 21-42 per cent (Usha and Kurien, 1981).

Morphologically H . oryzicola i s very c lose t o H . oryzae, from which i t d i f f e r s mainly i n var ious measurements of t he s t y l e t and sp icu le s , and length of t he hya l ine terminal p a r t of t a i l of t he second s t age juven i l e (Bao and Jayaprakash, 1978). Sharma and Swarup (1984) provide add i t iona l morphometric d a t a and information on the l i f e cyc le .

Heterodera deZvii J a i r a j p u r i , Khan, Se t ty & Govindu, 1979

This s p e c i e s , belonging t o the goettingiana group, w a s descr ibed from I1ragiv1 (Eleusine coracana) i n Bangalore, Karnataka, Ind ia ( J a i r a j p u r i , 1979). var ious EchinochZoa spec ie s , Sorghum vulgare, maize, p e a r l m i l l e t , f i nge r m i l l e t , Setar ia i t a z i c a , wheat and Pan?hm miliaceum.

Other hos t s repor ted by Krishna Prasad e t a l . (1980) include

H . dezv i i i s c lose t o H . eyperi , H . mothi and H . graminis, but d i f f e r s mainly i n the s t r u c t u r e of the underbridge which shows a subci rcu lar mass a t tached t o i ts c e n t r e ; t h i s is absent i n the o the r spec ies . No informa- t i o n is a v a i l a b l e on the biology and the pa thogenic i ty of t h i s spec ies . It i s not known ou t s ide the type loca t ion , and at tempts t o f ind it again a t t h i s s i t e have been unsuccessful (Sharma and Swarup, 1984).

Heterodera sorghi J a i n , S e t h i , Swarup & Sr ivas t ava , 1982

This spec ie s , of the schacht i i group, has been found on Sorghum vulgar@ a t Raispur , Utah Pradesh, Ind ia ( J a i n e t a l . , 1982); it has a l s o been recorded, without i nd ica t ion of h o s t , a t Hayatpur, Haryana S ta te (Dhawan e t a l . , 1983). I n pot experiments H . sorghi reproduced on EchinocZoa colona, Eleusine caracana, Se tar ia i ta ' l i ea , Paspazwn scrobiculatum. and Swarup, 1984).

Zea mays, Cypems rotundus and Ponnisetwn typhoides (Sharma

H . sorghi is cha rac t e r i s ed by a very t h i c k , prominent and dark brown underbridge. Sharma and Swarup (1984) bu t nothing i s known about i t s pathogenici ty .

Data on morphometrics and l i f e cyc le have been provided by

Heterodera r a s k i i Basnet & Jayaprakash, 1984

H. r a s k i i w a s r e c e n t l y descr ibed from Hyderabad, Ind ia , on CypeWS bulbosus Vab., which i s p re sen t ly the only known h o s t (Basnet and Jayaprakash, 1984). It is i n the goettingiana group and considered t o be c lose t o H . cyper i Golden e t a l . , 1962. Cysts are very elongated, somewhat resembling those of Doliehodera Mulvey & Ebsary, 1980. Nothing i s known about pa thogenic i ty o r hos t -pa ras i t e r e l a t ionsh ips .

Other Heterodera spec ies

Heterodera schacht i i Schmidt, 1871: This t y p i c a l l y temperate spec ies has been recorded itl Western Tropical Afr ica i n two loca t ions c lose t o each o ther . In Senegal , H . schacht i i -was found i n a s m a l l vegetable garden i n the cen t r e of Dakar, where it caused damage t o bee t s and cabbage (Luc and Netscher, 1974); ex tens ive surveys i n the surrounding area were negat ive.

362 M. LUC

It w a s recorded l a t e r i n Banjul , Gambia, a l so on bee ts (Bridge and Manser, 1980). with rooted p l a n t s and i s n o t considered to pres-ent a danger t o crops i n Senegal and Gambia.

In both cases , H . schachtii w a s probably introduced from Europe

Heterodera graminophiZa Golden & Birch f i e ld , 1972: Although found by Golden and B i rch f i e ld (1972) ou t s ide of a s t r i c t l y t r o p i c a l area (Louis iana. USA), H . graminophiza i s c i t e d here as B i rch f i e ld - ( l973) s t a t e d t h a t the spec ies "may not be n a t i v e t o the USA s ince the hos t p l a n t , EchinochZoa cozona and o ther c l o s e l y r e l a t e d forms, a r e cu l t i va t ed i n t r o p i c a l A s i a and Afr ica f o r food". H . graminophiza w a s s a id by B i rch f i e ld (1973) t o be a typ ica l of the genus because of t h e absence of g i a n t c e l l s a t the feeding po in t wi th in the hos t roo t : the females feed i n tlie pe r i cyc le and phloem t i s s u e s of the h o s t ; no g a l l i n g of r o o t t i s s u e s and no f o l i a g e symptoms occurred. B i rch f i e ld (1973) considered it an example of a well-balanced hos t -pa ras i t e system, poss ib ly ind ica t ive of a very o ld p a r a s i t e r e l a t ionsh ip .

'

Other hos t s a r e r i c e and SOPghwn hatepense.

H. graminophiza belongs t o the goettingiana group, and is c lose t o H . cyperi and H . graminis.

Heterodera avenae Wollenweber, 1924: This i s a cosmopolitan spec ies present i n some w a r m p a r t s of t he world. It has been a s soc ia t ed with severe d iseases i n Ind ia known as llmolyatl. Many Indian pub l i ca t ions r e f e r t o H . avenue but it occurs only on temperate c e r e a l s such as bar ley and wheat. On the con t r a ry , t r o p i c a l c e r e a l s such as EchinochZoa coZona, Sorghum vuzgare, EZeusine coracana, Oryza sa t iva , Setaria italica, Zea mays a r e non-hosts ( G i l l and Swarup, 1971; Sharma and Swarup, 1984). Thus s t r i c t l y H . avenue does n o t qua l i fy as a t r o p i c a l c y s t nematode.

Heterodera chaubattia Gupta & Edward, 1973: This spec ie s was found i n the s o i l of an orchard. a t Chaubat t ia . Uttah Pradesh. Ind ia (Guota and . L

Edward, 1973). This is the only r e p o r t and Sharma and Swarup (1984) f a i l e d t o red iscover the nematode. a t the type l o c a l i t y . Moreover types are apparent ly n o t a v a i l a b l e f o r study and thus taking i n t o account the . discrepancies and lack of information i n the o r i g i n a l d e s c r i p t i o n , Sharma and Swarup (1984) considered H . chaubattia to be a spec ies inquirenda.

GENUS CACTODERA K r a l l 6 K r a l l , 1979

Cactodera cacti ( F i l i p j . & Sch. Stekh. , 1941) K r a l l & K r a l l , 1979

This spec ies w a s repor ted on Echinopsis spp. i n Mysore, Ind ia (Kumar, 1964); the nematode could have been introduced with the h o s t p l a n t , which i s an ornamental.

Cactodera amaranthi (Stoyanov, 1972) Krall & Kral l , 1969

This spec ies w a s descr ibed by Stoyanov (1972) as Heterodera amaranthi from roo t s of Amaranthus v ir idis growing i n experimental f i e l d s a t Sant iago de las Vegas, near Hayana, Cuba. Heavy i n f e s t a t i o n of the hos t w a s repor ted . It w a s found e a r l i e r i n F lor ida by G. J. Rau (unpublished r e p o r t , 1957). Golden and Raski (1978) who a l s o s tudied the d i f f e r e n t s t a g e s of the nematode.

Details of t h i s and subsequent f ind ing a r e given by

Hosts a r e l imi ted t o spec ies of Amaranthus (A. v i r id i s , A. spinosus, A. r e t r o f z e x u s ) . The d i s t r i b u t i o n includes Cuba, Mexico, and the USA


(F lo r ida , Colorado, I l l i n o i s , Kansas, Missouri , Tennessee) according t o Mulvey and Golden (1983). However, Reis (1982) r epor t ed a c y s t forming spec ies "c lose ly resembling" G. maranth i and heavi ly p a r a s i t i s i n g Amaran thus putulus i n a c i t r u s orchard near Maputo, Mozambique. Thus, t h e d i s t r i b u t i o n of t h i s spec ies may be r a t h e r wide.

GENUS GLOBODERA Skarb i lovich , 1959

4 (Stone, 1973) Behrens, 1975

The pota to c y s t nematodes have been repor ted i n t r o p i c a l coun t r i e s bu t only on r a r e occasions. The f i r s t r e p o r t i s t h a t of Tiago (1956) i n Mozambique; bu t a t t h a t t i m e t h e r e p o r t w a s considered ques t ionable . Ind ia , G. rostochiensis and/or G. pazzida a r e considered t o be se r ious p e s t s of po ta toes i n t h e N i l g i r i s reg ion , which i s a h i l l y p a r t of Madras S t a t e and does n o t have a t r o p i c a l c l imate (Seshadri and Sivakumar, 1962; Prasad and Chawla, 1965; Logisvaran and Menon, 1969). (1983) r e p o r t t he presence of both spec ies i n T a m i l Nadu S t a t e , t h e southernmost s ta te of Ind ia and t r o p i c a l . G. rostochiensis had a l s o been recorded i n South A f r i c a , where i t s d i s t r i b u t i o n appears t o be r e s t r i c t e d t o t h e Transvaal (Kleynhans, personal communication).

In

Sharma and Swarup

Apparently, G. rostochiensis and G. pazzida are n o t adapted t o the t r o p i c s ; they may su rv ive i n some loca t ions ( a s does H. schachtii) bu t they do n o t pose as severe a t h r e a t t o pota to crops i n h o t t r o p i c a l a r eas as do var ious root-knot nematode spec ies .

GENUS AFENESTRATA Baldwin & B e l l , 1985

Afmestrata africana (Luc, Germani & Netscher, 1973) Baldwin & B e l l , 1985)

A. africana w a s descr ibed by Luc e t a l . (1973) as t h e second spec ie s of Sarisodera Wouts & Sher, 1971, as the two spec ies apparent ly shared seve ra l important cha rac t e r s inc luding t h e presence of c y s t s devoid of f e n e s t r a and a p e c u l i a r vu lva l cone formed by t h e hypertrophied and very pro tuberant vu lva l l i p s . hydrophiza Wouts & Sher , 1971, by Baldwin and B e l l (1985) l ed these au thors t o cons ider t h a t t h e females do n o t transform i n t o c y s t s ; consequently, they e rec t ed the genus A f e n e s t r a t a (without c y s t s ) .

Restudy of t he type spec ie s , Sarisodera

(with c y s t s ) , d i s t i n c t from Sarisodera

Morphology, caryology (n = 9 , as i n many Heteroderinae) and h i s to logy of t h e syncytium produced by A. africana are wel l documented (Luc e t a l . , 1973; Taylor and Luc, 1979; Baldwin and B e l l , 1985). The only repor ted h o s t i s Guinea g r a s s Panicum m a x i m w n , and the only l o c a t i o n i s the type l o c a l i t y - Experimental F i e l d s , ORSTOM Centre, Adiopodoume near Abidjan, Ivory Coast. However, A. africana came from Kenya, East Af r i ca and the loca t ion i n the Experimental F i e l d s where the nematode w a s found had been p lan ted wi th rooted seedl ings of Guinea g ras s obtained by p l a n t breeders during an expedi t ion i n Kenya.

Because of t h i s no f u r t h e r i nves t iga t ions of A. africana have been .undertaken and the o r i g i n a l f i e l d w a s t r e a t e d wi th a nemat ic ide , then p lan ted wi th a non-graminaceous crop. t r a n s f e r r e d t o a greenhouse a t the Nematology Laboratory, ORSTOM Cent re , t o be used only f o r taxonomic o r anatomical s tud ie s .

A population of A. ajcricana was

364 M. LUC

CONCLUSION

This paper somewhat resembles a catalogue and it i s d i f f i c u l t t o s epa ra t e i n t o sec t ions . Nevertheless , t h e following t e n t a t i v e c o l l e c t i v e conclusions can be made:

- with a few except ions , among the genera of cyst-forming Heteroderinae found i n temperate c l ima tes , only spec ies pe r t a in ing t o the genus Heterodera have been repor ted i n the ho t t r o p i c a l c l imates ;

- the twelve Heterodera spec ies considered he re in as t rop ' ical belong t o only two groups of the genus: the schacht i i group foryzae, sacehari, cajani , zeae, gamgiensis, oryzicola and SOPghi), and the goettingiana group (cyper i , mothi, graminis, deZvii and r a s k i i ) ;

- a l l these spec ies with the except ion of H . mothi and H . cajani show a c l e a r h o s t p reference f o r graminaceous p l a n t s ; s e v e r a l of them have been descr ibed from, o r experimental ly r a i sed on, r i c e b u t it must be kept i n mind t h a t r i c e i s the most c u l t i v a t e d c e r e a l i n the t r o p i c s , and con- sequent ly the most f requent ly sampled;

- these spec ie s , with a few except ions , have been repor ted only i n West Afr ica and i n Ind ia but t h i s could n o t be considered conclusive i n r e l a t i o n t o geographical d i s t r i b u t i o n because the re a r e no negat ive r epor t s from o the r t r o p i c a l p a r t s of t he world;

- cur ren t ly , on ly the genus Afenestrata can be considered as s t r i c t l y t r o p i c a l ;

- with the l imi t ed information on t h e biology of these spec ie s , i t can be s t a t e d t h a t t h e optimal temperature f o r hatching i s r a t h e r high, and there a r e seve ra l generat ions per year , t h i s being co r re l a t ed i n seve ra l spec ies with the presence of a voluminous egg-mass a t tached t o the c y s t ;

- as f a r as p resen t research observat ions permi t , i t may be assumed t h a t only H . ca jani , H . oryzicola, H . zeae and t o some e x t e n t H. sacchari a r e dangerous spec ie s ;

- c y s t forming Heteroderinae appear less important i n t r o p i c a l c l imates than i n temperate a reas but research on the group should no t be neglected because of t h e i r presence on var ious important c rops , p a r t i c u l a r l y ce rea l s .

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12

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t

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,

CYST NEMATODES IN EQUATORIAL AND HOT TROPICAL REGIONS 37 1

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i I

DISCUSSION I Noel s t a t e d t h a t , according t o d a t a of D r W. Hadisoeganda, H. gZycines

i i s p i e sen t i n Indonesia as a r e o the r spec ies of the genus and he asked i n what wqy.HH. glycines i s r e l a t e d t o H . cajani. Golden responded t h a t a l though the two spec ies a r e c lose ly r e l a t e d they are d i s t i n c t , l a rge ly on the b a s i s of s t r u c t u r e of the cys t w a l l . Stone s t a t e d t h a t populat ions of H. oryzicolQ h e i s s tudying with D r A . K. Koshy a r e very c l o s e , i f no t i d e n t i c a l , t o H. oryzae; they a t t a c k both r i c e and banana-trees , as does H . oryzae. Stone a l s o s t a t e d t h a t H. chaubattia i s probably a junior synonym of GZobodera mali .

B IChir reported t h a t c y s t nematodes a r e not economically important i n Tunis ia , as compared with MeZoidogyne or Pratylenchus, r e f l e c t i n g the genera l s i t u a t i o n i n the t r o p i c s , b u t GZobodera spec ies can be a problem when potatoes a r e grown continuously. This l ed to a d iscuss ion about t h e e f f e c t of temperature on the behaviour of po ta to c y s t nematodes from which it w a s concluded t h a t , based on observa t ions i n var ious coun t r i e s e.g. N e w Zealand, Tunis ia , Pakis tan , they become inac t ive when s o i l temperatures exceed 30°C but they may never the less c o n s t i t u t e a problem i n cooler seasons.

Edongali s t a t e d t h a t the H. trifolii reported i n Libya does no t a t t a c k subterranean clover and i n response t o a quest ion by Luc he s a i d t h a t H. Zatipons has no t y e t been recorded i n Libya, whereas i t i s present i n Tunis i a.

cyst nematodes in equatorial and hot tropical regionsfrom: cyst nemat'qdes edited by f....

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