.JnseclSci. Applic. Vol. 14. No. 3, pp. 273-284, 1993 Printed in Kenya. All r~ghtsreserved

WORLD REVIEW OF THE NATURAL ENEMIES AND DISEASES OF LITURA (F.) (: NOC'TUIDAE)'

G. V. RANGA RAO, J. A. WIGHTMAN and D. V. RANGA RAO Legumes Entomology, International Crops Rcscuch lnst~tutefor the Semi-And Tropics. (ICRISAT), Patancheru P.O. 502 324. Andhra Pradesh, India

(Received 5 February 109 I; clcceprr~lI I'chr uclry 1992)

Abstract-Published information was used to compile a summary of natural enemies(parasitoids, predatorsand diseases) reported attacking Spodoptera litirra (F)under field conditions. (71) of parasitoids in seven families of and two families of Diptera were listed as parasitoids of difTerent stages of S. litura. Predatory (36) belonging to 14 families and 12 species of spiders from six families have been reported to feed on this species. Four prolozoan, four fungal, seven bacterial, four each of viral and nematode species were also reported to be the pathogens of this species. Published information suggested that periodic releases of large number of egg parasites could help in suppressing populalions of this pest. There is a considerable scope for increased attention to the role of natural enemies as component of integrated pest management programmes of S. litura.

Key Words: Spodoprera lirura, natural enemies, diacascs

ResumC - Compte rendu mondial des ennemis naturels el des nialadies de Spodoplera litirra (F.) (Lbpidoptbres: Nwtuides): Des informations publiees ont servi de source pour la compilation de ce compte rendu des ennemis naturels (parasites et predateurs) et des maladies qui s'attaquent P Spodoptero lifura (P.)en milieu reel. ~oixante-neufespkes d'insectes parasitoid parmi sept familles des HymknoptCres ainsi que de familles des Dipthres sont classees comme parasites des difTkrenls stad~de~.l~ura.~rente-~ixinsectes~r~dateurs~a~~artenant~14familleset12esp&ccsd'araingnks provenant de six familles sc nourriraient de cette espcce. Ouatre protozoares, uuatre moisissures, iept bactbries, quatre virus et cinq nematodes seraient ~~illeme~tdes agents iathogi.nes de cette espkce. Les information publikes laissant P croire que des introductions pkriodiques des parasites des oeufsen grands nombres permettraient de supprimer la population de Spodoplera. Les ennemis naturels ofTrent des possiblites importantes pour jouer un r61e rlt! dans des programmes de lutte integrke contre S. lilura.

Mots Clks: Spodoprero liruro, ennemis naturels, maladies

INTRODUCTION known to cause severe damage to many crops including tobacco and groundnut in India (Moussa et The tobacco caterpillar, Spodoprera litura (F.) is one al.. 1960; Ayyanna et al., 1982). of the most important insect pests of agricultural The "green revolution" in Asia brought with it an crops in the Asian tropics. This species is widely increased awareness of the potential of inscclicides distributed throughout tropical and temperate Asia, for increasing the sustainability of rice production. Australasia and the Pacific Islands (Feakin, 1973; Unfortunately, the involvement of farmers in Kranz et al., 1977). It is a polyphagous pest and insecticide related technologies did not proceed as fast z the rate of subsidy spread and the overspill of *~ubmittedas~ournal ArticleNo.7076 by the International Crops Research Institute for the Semi-Arid Tropics, insecticide usage into the fields of legume growers Patancheru 502 324, Andhra Pradesh. India. and horticulturalists. Legume pests are increasing in 274 G. V. RANGA RAO et al. g economic importance all through Asia due to the furnished in this paper may suggest new sources of destruction of natural control systems, and the build- natural enemies for different geographic regions. It up of insecticide resistance following the may also indicate the potential importanceofdifferent "spraymania" of many farmers. If this is to be natural enemies which may be relevant to future counteracted, natural control needs to be given biocontrol and IPM projects. increased emphasis as acomponent of IPM approach. S. litura populations in groundnut fields (our study EGG PARASITOIDS crop) are increasing in number and intensity, especially in fields where insecticides have been Four species of trichogrammatids, one scelionid applied (Ranga Raoand Shanower, 1988; Stechmann and one braconid which had been reported as egg and Semisi, 1984). parasitoids of S. lirura, Chelonus sp, and Telenomus In the past, the control of depended spp., have also been reported as both egg and larval mostly on inexpensive and efficient insecticides. But parasitoids. A total of 10 egg parasitoids have becn in recent years populations of many pests including reported from different parts of the host distribution S. lirura have developed resistance to many (Table 2). Among the trichogrammatids, T. commercially available pcsiicides (Ramakrishnan et austrulicum and T. chilonis from India (Joshi et al., a]., 1984). Moreover, outbreaks of secondary pests, 1979; Patel et al., 1971), T. japonicum from and the effect of pesticides on non-target organisms Indonesia (Chu, 1979), T. dendrolimi from China is becoming increasingly common. Because of these (Chiu and Chou, 1976), are the most common. These reasons, the control of pests is becoming species are often reported from eggs of several other increasingly difficult and it is vital that all biological hosts. alternatives to insecticides need to be given greater Mass releases ofan indigenous egg-larval parasite priority, both in research and application. Chelonw helipae in 1971-1973 in Anand, Gujarat, Although S. lilura is known to cause substantial India, against S. litura in cauliflower crop proved damage to a number of crops, there has been no ineffective in controlling the pest. During 1974, attempt to review the literature on the role of natural weekly release of Telenomus remus Nixon, an exotic enemies in regulating the abundance of this pest. The egg-larval parasitoid, in a tobacco nursery did not main purpose of this paper is to review the status of result in any parasitism. However, five weekly natural enemies of S. litura on global basis. S. lit~tra releases of 50,000 parasitesl0.2 ha and two ieleases is known to be attacked by many of r!atural enemies of 15,000 parasitoids/0.2 ha in cauliflower, resulted at various life stages. Altogether, about 13 1 species in 60% parasitism (Pate1 et al., 1979). of natural enemies have been reported from different Rao et al. (1979) observed 8% parasitization by parts of the world (Table 1). The list of parasites, Chelorius blackburni Cameron an eggllarval predators and diseases was compiled from published parasitoid of S. litura in Karnataka, India. An insect literature, and arranged in a systematic order that survey conducted in JawaTimur, Indonesia revealed includes host stage attacked, scientific name, family the occurrence of the parasitoids Trichogramma and geographic distribution, followed by the most japonicum Ashm, and Telenomus dignus (Gah) pertinent references (Table 2). The information attacking S. lilura eggs (Chu, 1979). Chiu and Chou

Table 1. Spodopteru lilura natural enemies and diseases worldwide Number of natural enemy species and diseases reported from different countries Parasitoids predators* Nematodes Diseases India Australia Japan China Indonesia Western Samoa Papua New Guinea Philippines New Zealand Total 7 1 48 4 20 *spiders are included. Natural cncrnles of Spodoprera lirura

Tablc 2. Natural enemies and dlicascs of Spodqrera lirura (Fabricius) Stagc attackcd/parasi~c/ Geographic prcdatorlpathogcn Family rarlgc Sclcctcd rcfcrcnces PARASITES Ese Chelonus helipae Gupta lndia Patel et al. (197 1) Tricho~rummaausrralicum Girault Trichogrammatidac lndia Joshi et al. (1979) Trichogromnto rhilonrs lshii Trichogramrnatidac India Bhatnagar ( I98 1 ) Z~Land Kushwaha (1983) Trictro~rantmadendrolimi Mats. Trichogran~matidae China Chiu and Chou ( 1976) Telcnomus dignus (Gah) Scclionidae Indonciia Chu (1979) Trichogrumma japonicum Ashm. Trichogramma~idae Indonesia Chu ( 1979) Egg-larval Chclottrts hlac.khurni Camcron Braconidac lndoncsia Rao ct al. (1979) Chelonus carhonu~orMarshall Braconidae Indiii Rao and Satyanaraya (1984) Chelontcs fi~rmosanrcs(Sonan) Braconidac lrldia Patcl et al. (1971) Rai (1974) Scclionidac lndia Joshi ct al. (1979) lndia Patel ct al. (1979) lndia Rao and Pate1 ( 1976) India Zaz and Kushwaha (1983) Ncw Zealand Anon (1977) Western Samoa Braune ( 1982) Larval Aponre1e.c sp. Braconidae Wcstcrn Samoa Braune et al. (1981) Wcstcrn Samoa Stechmann and Semisi (1984) Apanteles ncar A. rltficrrts (Hal.) Braconidae Wcstcrn Samoa Braunc ct al. (1981) Aponreles sp. ncar Colenlani Vicr Braconidae lndia Patel et al. (1971) Apanreles sp. (ocronarius group) Braconidac lndia Joshi ct al. (1979) Apanteles chilottis Mats. Braconidae Indonesia Chu (1979) Apan~elesrolemoni Vicrcck Braconidac India Sathe (1987) Apanteles marginiventris Braconidac Australia Michacl ct al. (1984) Apanteles plutellae Kurd. Braconidac China Chiu and Chou (1976) ~pantelesprodcniae Viereck Braconidae lndia Sathe (1987) Apanteles rrtjicrus (Hal.) Braconidac China Chiu and Chou (1976) Western Samoa Braune ct al. (1981) lndia Zaz and Kushwaha (1 983) Apattteles kazuk Telenga Braconidae Australia Michael et al. (1984) Bracon hrevicornis Wesmacl. Braconidae lndia Thontadarya and Nangia (1983) Cotesia (Apanreles) morginiventris (Cresson) Braconidac lndia Jalali (1987) Diadegnru ar~mteopilosaCamcron lchncurnonidae India Sathe (1987) Echrhromorpha sp. Braconidae India Sathe (1987) Enicospilus sp. Braconidae India Sathc (1987) Microgasrer sp. Braconidae China Xu and Yang (1983) Micropliris sp. Braconidae lndia Zaz and Kushwaha (1983) Microplitis demolitor Wilk. Braconidae Australia Hafcz (1951) Micropliris paNidipes Szepl. Braconidae China Chiu and Chou (1976) Micropliris prodiniae R.S.K. Braconidac lndia Sathe (1987) Micropliris ruhcrculifera (Wesm) Braconidae China Chiu and Chou (1976) Rogas sp. Braconidae lndia Bhatnagar (1981) Snellenius manilae (Ashm) Braconidae China Chiu and Chou (1976) Zele chlorophrhalma Nees. Braconidae lndia Bhamagar ( I98 1) Rao and Satyanaraya ( 1984) Brachymeria lasus (Wlk.) Chalcididae India Narcndran and Joseph (1 977) Lusiochalcidia ? erythropoda Chalcididae lndia Bhatnagar (198 1) Cameron Liromasrh maculara Isheii Encyrtidae China Sheng and Shen ( 1983) Euplectrus sp. Eulophidae China Chiu and Chou (1976) Euplecrrus gopimohani Mani. Eulophidae India Patel (1944) India Pate1 ( 1980) G. V. RANGA RAO et al.

Table 2 Contd. Stage attacked/parasite/ Geographic predator/pathogen Family range Selected references 1 Euplecrrus near E, xanrhocephalus Eulophidae Western Samoa Braune et al. (1981) Terrasrichus ayyari Rohwer Eulophidae lndia Sathe (1 987) Trichospilus pupivora Ferri Eulophidae India Sathe ( 1987) Compoleres sp. Ichneumonidae lndia Battu (1977) Compoleres chlorideae Uchida lchncumonidac India Battu (1977) lndia Sathe (1 987) China Chiu and Chou (1976) Charops bicolor (Szepi) Ichneumonidae China Chiu and Chou (1976) Charops obsrusus Morl. Ichneumonidae lndia Patel (1980) Hyposorer didymaror Thunb. Ichneumonidae Australia Michael er al. (1984) Eriohoru.~sp. Ichneumonidae lndia Bhatnagar ( I98 1 ) Ichneumon SD. Ichneumonidae lndia Anon. (1988) Nerelia ferruginea Cameron Ichneumonidae lndia Sathe (1987) Paniscus oroducrus Brulle Ichneumonidae Australia Hafez (1951) Temelucha bigurrula (Mats.) Ichneumonidae Indonesia Chu (1979) Fannia leucosrica Smith Muscidae lndia Zaz and Kushwaha (1983) Acria nigrirula Mall. Tachinidae Australia Hafez (1951) Parasarcophago misera (Walk.) Tachinidae lndia Bhattu ( 1977) Peribaea orhara (Wideman) Tachinidae lndia Jayanth and Nagarkatti (1984) Philippines Rao and Pate1 (1976) Tachinidae lndia Bhatnagar (1981) India Joshi et al. (1979) lndia Patel et al. (1971) Sturmia aequulis Mall. Tachinidae American Samoa Hoyt (1955) Tritaxys sp. Tachinidae Australia Hafez (I95 1) Winrhemio near dispar (Macq.) Tachinidae American Samoa Hoyt (1955) Pre-pupal Chelonus sp. Braconidae Western Samoa Braune and Kan (1981) Pupal Bracliymcria sp. Chalcididae India Thontadarya and Nangia ( 1983) Hyhorhoracini sp. Chalcididae India Rao et al. (I98 I) Sarcophaga albiceps Meigen Sarcophagidae lndia Bhatnagar (1981) Sarcophaga duThoms Sarcophagidae lndia Joshi et al. (1979) Sarcophaga peregrine Sarcophagidae lndia Zaz and Kushwaha (1983) (Robineauedesvoidy) Blepherella serigera Corti Tachinidae India loshi et al. (1979) Winthemia sp. Tachinidae Western Samoa Braune and Kan (198 1) PREDATORS Egg Chalcid wasp (unidentified) Chalcididae Western Somoa Braune and Kan (1981) Cardiocondyla nuda (Mayr) Formicidae Western Somoa Braune and Kan (1981) Tapinoma melanocephalum (F.) Formicidae Western Somoa Braune and Kan (1981) Liposcelis sp. Liposcelidae India Zaz and Kushwaha (1983) Egg-lar +a1 Chrysopa crassinervis lndia Rao and Satyanaraya (1984) Esben Peterson Small red ants (unidentified) Formicidae Wcstcrn Somoa Braune (1980) Nesidiocoris renuis Reuter Miridae India Pate1 ( 1980) Conocephalus sp. Tettigonidae China Deng and Jim ( 1985) Lflrval Carabid beetles (unidentified) Carabid Japan Ito et al. (1972) Crickets (unidentified) Japan Ito er al. (1972) Earwigs (unidentified) Japan Ito et al. (1972) Casnoidea indica (Thnb.) Carabidae Indonesia Chu (1979) Chrysopa sp. Chrysopidae lndia Sitaramaiah and Ramaprasad (1982) Coccinella sp. Coccinellidae India Sitaramaiah and Ramaprasad (1982) Epilachna sp. Coccinellidae India Cherian and Brahmachari (1942) Dieuches sp. Lygaeidae India Rao et al. (1981) Oechalia consocialis 4Bois.) Pentatomidae Australia Richard (1964) E!PUl uede r

PJ"!l qsnu p1mq qsng

(2x61) pese~dowr!~pue qc!etueJel!S (PX6 1 ) UJX (~861)pese~deuc~ put ~I!!!"UICJRI!S (CL61) !CqlUCMCH put! 0610~ (ZL61) U"xeH

fLZ UJIII!/~~rldopods jo ~s!uauj1eJn)eN 278 G. V. RANGA RAO et al.

Table 2 Contd. Stage attackedlparasitel Geographic predatorlpathogen Family range Selectcd refercnces Bacteila ha1 Bacillus cereus lndia Kore and Bhidc (1978) Frankland and Frankland Oblisami et al. (1969) Bacillus rhuringinsis Berliner lndia Zaz and Kushwaha (1983) Metarhizium anisopliae lndia Siddaramaiah et al. (1986) (Metchnikoff) Sorokin Micrococcus sp. Indiu Zaz and Kushwaha (1983) Serraria marccscctts Bizio lndia Ansari ct al. (1987) Pandey and Rangarajan (1967) Zaz and Kushwaha (1983) Streptococcus sp. lndia Zaz and Kushwaha (1983) Streptococcusfaecalis A.& H. India Battu ct al. (1972) Zaz and Kushwaha (1983) Virus Larval Baculovirus group Ncw Zealand Longworth (1976) Cytoplasmic polyhedrosis virus Japan Asayama and Osaki (1 970) Granulosis virus China Tsai et al. (1978) lndia Battu ct al. (1978) lndia Narayanan ( 1985) Nuclcar polyhedrosis virus lndia Battu et al. (1972) lndia Chari et al. (1985) lndia Krishnaiah et al. (1985) lndia Ramakrishnan and Tiwari (1969) China Tai (1973) Japan Okada ( 1974) Nematode Larval Hcsomer~niisspp. lndia Bhatnagar et al. (1985) Srinernemofelriae (DD 136) Japan Kondo and Ishibashi (1984) (Neoplecrana carpocapsae Wciscr) lndia Janardan Singh and Bardhan (1974) Ovomermis albicans (Sicbold) lndia Bhatnagar ct al. (1985) Penruronlimermis spp. lndia Bhamagar et al. (1985)

(1976) reported that T, dendrolimi is an egg of this species, Of thcse, 47% were braconids, 19% parasitoid of S. litura from Taiwan. ichneumonids, 16% tachinids, 10% eulophids, 3% Braunc (1982) found Telenomus remus Nixon to chalcids, and 2% scelionids, encrytids and muscids. be a common egg-larval parasitoid of S, lilura in In gcneral, 84% were Hymenoptera, and 16% Diptera. Western Samoa with parasitism averaging 54%. In India, 32 different species of parasitoids have Complete parasitization was observed only in small been reported as larval parasitoids ofS, litura. Among egg masses (up to 150 eggs) and the percentage of these, Apanrcles and Bracon sp. were the most parasitization decreased with an increase in size of commonly reported. In 1974, Rai surveyed vegetable egg-mass. T. renius could oviposit only in host eggs crops in the state of Kamataka and found that 10% of on the &~rfaceof the host egg mass. Thus, the larval mortality was caused by Chelonusformosanus effectiveness of T. remus was limited on the large (Sonan). Battu (1977), during a survey of castor and compact egg masses of S.lirlrra. cauliflower in the Punjab, found thatParasarcophaga misera (Walk.) and Cam/)olitis sp. also attack S. LARVAL PARASITOIDS lirura larvae. Jayanth and Nagarkatti (I 984) reponed the emergence of up to 12 tachinid parasitoids Generally, thelarval stageofs. litura is more prone [Perihaea orhata (Wideman)] from a single S. lirura to parasitism. Larval parasitoids of S. litura attack larva in Karnataka state, lndia. young to mature larvae and afew alsoattack eggsand Rao and Satyanarayana (1984), during a pest larvae, and larvae and prepupae. Fifty-eight parasitoid survey of natural enemies of S. litura in Andhra species have been reported to attack the larval stage Pradesh, India, reported Zele chlorophthalma Nees Natural enemics of Spodoprera lrrrtru 279 as a larval parasitoid and Lasiochacidia them is a larval -pupal pamitoid (Ichneumonsp.)and erythropodus Nee as a pupal parasitoid. one a prepupal pmsitoid (Chelonw sp.; Table 2). Sathe (1987) in a survey for natural enemies of S. Lasiochalcidia erythropodus Cameron was litura in Maharashtra region of lndia reported rcportcd as a pupal parasitoid of S,litura in Andhra Compoletes chloridae Uchida and Ap~trtel~~sPradcsh, lndia (Rao and Satyanamyana. 1984). colemani Viereck. During the same survey two new tfowcvcr. Bhatnagar (1981) reported this species as BraconiP species (Enicospilus sp. and a larval parasitoid. Echthromorpha sp.) were found responsible for the 5% parasitization of S. litrrru while A. colentuni and PREDATORS A. prodcniae parasitized up to 20% larvae. Laboratory tests to determine the effect of host Altogether 36 predatory insects from 14 farn~lies plants on the degree of parasitism of S. litura larvac and 12 species of spiders, representing six families by Cotesia (Apanteles) marginiventris Cresson, were reported to feed on S. litura eggs, larvae and showed least preference for larvae on tobacco (Jalali pupae in different pans of the world. Of the total et al., 1987). Although 20% of the larvac wcrc prcd;~torsrcportcd to feed on S. litura, 508 of the parasitized on tobacco leaves, female parasitoids insect predatory fauna and 83% of the spiders were became inactive after contact with the leaves and from India (Table 1). died within 1 hr suggesting that C. ntarginiventris Sitaramaiah et al. (1975) from Andhra Pradesh, would not be suitable for release against S, litrtru on India, observed for the first time the reduvid Harpactor tobacco. The most preferred host for the parasitoid costalis Stal. predating on S. litura larvae in tobacco was Khol rabi (56% parasitiytion). crops. Laboratory experiments revealed that the Application of 2% neem (Azadiracta indica) predator consumes an average of 63 S. litura larvae kernel suspension to eggs ofS. lituru before and after during its life span. parasitization by Telenomus rentus Nixon had no Nymphs and adults of Andrallus spinidens (F.) effect on the development of the parasitoid in lndia (pentatomid) were observed feeding on S. litura (Joshi et a]., 1981). Thus, neem seed kernel suspension larvac in rice in Himachal Pradesh, India (Pawar was considered suitable in the integrated control of S. 1976). Another pentatomid species, Canthoconidia litura in tobacco nurseries. Jurcellatu (Wolf), was observed feeding on larvae of Six parasitoid species, Apanteles rNJicrus (Hal.), S. litura in tobacco nurseries in Andhra Pradesh, C. marginiventris (Cresson), A. kazak Telenga, lndia (Kapoor et al., 1975). The biology of this Compoletes clrloridae Uchida, Hyposoter didymator predator was studied in the laboratory with a view to Thumb and Telenomus remus Nixon were use C,furcellata in an integrated pest management introduced to Western Australia from overseas in programme for tobacco pests. 1978- 1983and released against S,litura and 1 I other Nakasuji et al. (1976) observed a predatory wasp, economically important pests. The highest level of preferentially selecting fifth and sixth instar larvae parasitism by A. ruficrus was noticed in Mythimna over early instars. The wasps were more active and sp. (80% and above; Michael et a]., 1984). attacked more larvae in fields with high larval density In Western Samoa, Stechmannand Semisi (1984) than those with low larval density. However, the collected information on S,litura damage levels in percentage of predation was lower in the field with relation to natural populations ofApanteles sp. in the highest density of S. lirura larvae. tam fields. They found that this pest is more severe on Chu (1979), from a survey in Jawa Timqr, "Taro" crop where insecticides and herbicides were Indonesia, reported a carabid beetle, Casnoidia widely used, which perhaps created imbalance indica (Thnb.) and a staphylinid beetle Paederus between the pest and its natural enemies. Barrion and fuscipes Curt, feeding on S. litura and other Litsinger (1987) reported the presence of Peribaea economically important lepidopterous insects. Chu orbata (Wideman) as a gregarious larval parasitoid and Chu (1 975) studied the effects of temperature on on S. litura. the growth of C.furcellata and found that 7 1.2 16and 134 C degree days were required for egg, nymph and PUPAL PARASITOIDS adult stages, respectively. It wasconcludedthat there are five to six generations per year of this predator in Relatively few pupal parasitoids have been northern Taiwan. reported from S. litura when compared to parasitoids Deng and Jim (1985) reported Corrocephalus sp. of other life stages. Eight parasitoid species have (Tettigometridae) as new predator on egg masses of been reported from the pupal stage of S,litura, one of S. litura in Guangxi, China. This katydid was 280 G. V. RANGA RAO et al. successfully reared on artificial diet. Field releases of in the second fortnight of June, was highest in mid- nymphs and adults of Conocephalus sp. were August, and decreased by November. attempted against the control of Scirpophaga incertulus (Walk.). Virus

DISEASES Viral diseases of this species have been reported from China, Japan, India and New Zealand. Among Protozoa the viruses, nuclear polyhedrosis viruses are the most common and potent. Nosema carpocupse Paillot was found to infect S. Krishnaiah et al. (1985) conducted field trials litura larvae in New Zealand (Malone and Wigley, with a nuclear polyhedrosis virus against S. lirura 1980), India (Narayanan and Jayaraj, 1979), Japan damage in black gram (Vigna mungo) fields in (Watanabe, 1976)and China (Tsai et at., 1978; Li and Andhra Pradesh, India. Two spraysof virus suspension Wenn, 1987). containing 1.96 x 109polyhedral inclusion bodieslml at the rate of 1500 mlha, gave effective control Bacteria similar to chemical insecticides tested. Chari et al. (1985) evaluated the effectiveness of Seven bacteriaare known to infect S. litura at larval integrated management of natural enemies and viral stage in India (Table 2). Ansari et al. (1987) reported diseases to control S. litura on tobacco seedlings in Serratia marcescens Bizio from Karnataka, India, Gujarat, India. They concluded that acombination of attacking larvae of the noctuids Helicoverpa biological control agents, insect growth regulators, arntigera Hiibner and S, litura. In labortory tests, S. antifeedants and a trap crop on all sides of a nursery litura was found more susceptible to the bacterium is an ecologically sound procedure for the control of than H. armigera. The bacterium was equally S. litura. pathogenic when ingested through artificial diet or Narayanan (1985) from Karnataka, reported the the natural food plant, but pathogenicity by contact occurrence of a granulosis virus in dead S. Iitura application to the body of larvae was poor. larvae. Eggs and all six larval instars were highly Zaz and Kushwaha (1983) found Bacillrrs susceptible to the virus, the mortality was 100% in thuringiensis Berliner (B.t.) to be an effective eggs and first to fifth instar larvae and 50% in the last microbial insecticide against S. litlira larvae in larval instar. The disease killed older larvae more cauliflower fields in Rajasthan, India. Application of rapidly than the younger ones. the B.1. in combination with endosulfan resulted in 85% larval mortality in the field. Nematodes

Fungi Four nematode species have been reported parasitizings. litura in Indiaand oneof them has also So far four fungi have been reported to infect S. been reported to be parasitizing S. lirura in Japan. litura and cause physiological disorders in larval Bhatnagar et at. (1985), found S. lirura larvae growth and development. Asayama and Ohoishi parasitized by the mermithid nematodes, Ovomermis ( 1980) from Japan and Phadke and Rao ( 1978) from alhicans (Siebold), Hexamermis sp., and India, investigated the pathogenicity of a grem Pentatomermis sp. They observed more nematode muscardine fungus Nomuraea rileyi (Farlow) activity on alfisols than on vertisols. They also Samson. Laboratory studies in India indicated that discussed the population dynamics and distribution this fungus was harmless to eggs of an egg parasitoid, of nematodes and the arthropod hosts. Kondo and Telonemusprodiror Nixon, on Achea janata L., and Ishibashi (1984) explained the infectivity and recommended the combined use of the fungus and propagation of entomogenous nematodes the egg parasite in biocontrol programmes against A. Steinernemu sp. on S. litura from Japan. janara. This may also apply to S, litura management. Zaz and Kushwaha (1983) reported Beauveria CONCLUSIONS bassiana (Balsarno) Vuillemin. infecting S. lirura in cauliflower crops in Rajasthan. Siddaramaiah et al. Although 71 parasitoids and 48 predators are (1986) reported an incidence of larval infection with known to attack S,lirura at different stages, most of Metarhiziwn anisopliae (Metchnikoff) Sorokin in these associations were incidcnta1,either some species groundnut in Karnataka. The infection first appeared were rare or more closely associated with other hosts. Natural cnernies of Spodoprera liruro 28 1

Among the pathogens, five protozoans, four fungi, Japan) and morphology of the fungus grown on seven bacteria, four viruses and four nematodes are the larvae. Japanese J. Appl. Entomol. Zool. 24, known to infect the larval stage of S. litura. All 105- 107. reports of bacterial infections are from India. Asayama T. and Osaki N. A. (1970) Cytoplasmic Among the 10 species of egg parasitoids reported polyhedrosis of the cotton leaf worm, Prodenia the Tl'ichogramma was the most widely lirura (F.). J. Invertebr. Pathol. 16,292-294. distributed in India, Indonesia andchina. More larval Ayyanna T., Arjuna Rao P.. Subba Ratnam G. V., parasitoids have been reported from S. litura, 83% of Krishna Murthy B. H. and Narayana K. L. (1982) the parasitoids are hymenopterous and the remaining Chemical control of Spodoprera Iirura (F.) on 17% were dipterous. The parasitoids that attack the groundnut crop. Pesticides 16, 19-20. pupal stage are relatively fewer in number. Only Barrion A. T. and Litsinger J. A. (1987) A larval eight species have been reported, one of which was a parasite of swanning caterpillar and common larval pupal parasitoid and another prepupal cutworm in the Philippines. Inr. Rice Res. Newsl. parasitoid. 12.34-35. ' Altogether 48 species of insects and spiders were Battu G. S. (1977) Occurrence of Parasarcopha~a observed to predate on eggs, larvae and pupae of S. miseru (Walker) and Campoletis sp. as parasites lirura. Spiders account for 25% of the total predators. of Spodoptera lirura (F.) from India. Current Sci. In the past, the mass releases of egg and larval 46,568-569. parasitoids forthecontrol ofS. litura,indifferentcrops Battu G. S., Bindra 0. S. and Rangarajan M. (1972) in different geographical regions had achieved only Investigalions on microbial infections of insect partial success (Patel et al., 1979; Michael et al., pests in the Punjab. lndian J. Enrontol. 33,317- 1984). Our personal observations in ICRISAT 325. groundnut fields revealed more leaves with defoliator Battu G. S., Dilawari V. K. and Bindra 0. S. (1978) damage in insecticide applied fields than unsprayed Investigations on microbial infections of insect areas (Wightman et al., 1990). Similar observations pests in the Punjab--11. Indian J. Entomol. 39, were also made during farmers' field surveys in the 27 1-280. post-rainy season in Coastal Andhra Pradesh, India Bhanotar R. K. and Srivastava Y. N. (1985) Impact (Ranga Rao and Shanower, 1988). Stechmann and of reptiles on food production and environment. Semisi (1984) also shared the same opinion after Non insect pests and predators. Proc, of the surveying Taro fields in Western Samoa. In view of Narional Symposium on Impact of Non Insect the development of insecticidal resistance and the Pests and Predators Food Production and destmctionofthenaturalenemiesandthe polyphagous Enr~iro~~nletlt.March 12-14, 1984 India. All nature of this species, there is need to give morc lndia Scientific Writers Society. pp. 116-120, consideration to the role of natural enemies as a 265-266. component in integrated approaches to manage this Bhatnagar V. S. (1981) Arthropod parasitoids of species. insect pests (excluding Heliorhissp.) recorded in Andhra Pmdesh, India. Cropping Systems REFERENCES (Et~tomology)Progress Report, ICRISAT. BhatnagarV. S., PawarC. S., Jadhav D. R. andDavis Anonymous (1977) Report of the Deparrmenr of J. C. (1985) Mennithid nematodes as parasites of Scientific and industrial Research for the year Heliorhis sp, and other crop pests in Andhra ended 3 1 March 1977. New Zealand. 70 pp. Pradesh, India. Proc. of Indian Academy of Anonymous (1988) Annual Report-lnternariottal Sciences. Sci. 94,509-5 15. Crops Research lnsrirure for the Semi-Arid Braune H. J. (1980) Field studies on taro cluster Tropics(lCRISAT),Patancheru, Andhra Pradesh, caterpillar. Alafua Agric. Bull. 5,42-46. India. Braune H. J. (1982) Effect of structure of host egg Ansari M. A,, Pawar A. D. and Ahmed S. N. (1987) mass on the effectiveness of egg parasite of A note on pathogenicity of naturally occurring Spodoprera lirura (F.) (Lepidoptera: ). bacterium Serratia marcescens Bizio on some Western Samoa, Drosera 1,7-16. lepidopterouspests. Plant Pror. Bull. lndia 39,27- Braune H.J. and Kan S. K. (198 1)Speciescomposition 28. and biology of the predator-parasite complex of AsayamaT. and Ohoishi K. (198'0) Field infection of the taro cluster caterpillar, Spodoptera lirura (F.) Spodoprera litura (F.) larvae by the in Western Samoa. Alfua Agric. BUN. 6,650. entomogenous fungus, Nomuraea rileyi (in Braune H. I., Kan S. K. and Tnviranus H. (1981) 282 G. V. RANGA RAO et al.

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