MISCELLANEOUS REPORT NO 59

The Oriental Armyworm, separata (Wlk.) Distribution, Biology and Control: A Literature Review

H. C. SHARMA and J. C. DAVIES

Published by:

CENTRE FOR OVERSEAS PEST RESEARCH COLLEGE HOUSE WRIGHTS LANE LONDON W8 5SJ O Crown copyright 1983

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Printed by Hobbs the Printers Ltd of Southampton for the Centre for Oversees Pest Research H. C. SHARMA and J. C. DAVIES*

The Oriental Armyworm, (Wlk. ) Distribution, Biology and Control: A Literature Review

International Crops Research Institute for the Semi-Arid Tropics Patancheru P.O. 502 324 A.P., India

SUMMARY

The or~entalarmyworm, Mythimna separata IWlk.1, IS one of the serious cereal pests in the Asran and AustralIan conlrnents It IS widely distrrbuted bctween 45 N and 45's lat~tude and 60~Etn beyond 170' W lony~tude,covering 27 countr~es/colonialterritnr~es/~slands from humid tropical to temperate regions. Serious outbreaks have occurred In India, Ch~na, Japan, Australla, New Zcaland. FIII, Bangladesh, and Tha~lancl.Var~ous theories are quoted to cxpla~nthe outbreaks. Exact y~eld10% ~stimatcsare not ~va~lable,but heavy losses have beer1 experienced In India, Ch~na,Japan, Australla, New Zealand, and Barigladesh, development varies accord~ngto tempcratlrre and tiumid~ty.Populat~ons have been mon~toredclslng light traps, molassrs.ba~tcd traps, ariri dry sorghum leaves. Larval brhav~our varles cons~derablyIn thc sol~taryand yreyarlous phases. 111 and Japan migration s reported Forty-two parasrtes, 16 predators, and 12 pathogens have been reported from the field. Male and female pheromones have beer1 analysed. Mythirnna separ(3t.a feeds on 33 plant species belonq~ngto 8 fam~lrcs.D~fferent~al varietal suscept~b~lityoccurs Iri wheat, rlcc, malze, and sorghum. A large number of chem~calsg~ve eflect~ve control.

Introduction

The or~entalarmyworm, Mythinma () (Pseudaletia) separata (Wlk.) 1s a serlous cereal pest in Asia and (Butant, 1955; Chao and Chen, 1947: Hamhlyn, 1959; Common, 1965). It IS also comrr~or~lyknown as the southern armyworm, Chinese armyworm, paddy armyworm, sorghum armyworm, army caterpillar, ear-cutting caterpillar, and paddy cutworm. Other closely rclattx noctu~dsof economic Importance iri Asia, Australasia, Africa, and America include M. unipuncta (Haw), M. loreyi (Dup.1, M. pseudo-loreyi (Rungs), M. phaea (Hmps.), M. polyrabda (Hmps.),M. pinna (Saalm), M. convect;, (Wlk.), M. venalha (Moore),M. albistigma (Moore.), M. yu (Guen.), M, aspersa (Snell.), M. c~~rvula(Wlk.), M. compta (Moore.)t, M, inxilans (Butl.), M. irregularis (Wlk.), M. roseilinea (Wlk.), Spodoptera exempta (Wlk.), S, litlira (F.), S. frcigiperda (J. E. Smith) and S. littoralis (Boisd.) (Dav~dsonand Peairs, 1966; Nair, 1975; Avidov and Harpaz,'1969; Grist and Lever, 1969; Carneg~eand D~ck,1972; Khas~mudin,1978; ICRISAT, 1980). The reports pubitshed from India on M. ~/r,~pilnctaare now known to concern M, separata (Ramamant and Subba Rao, 1965). M. separata was considered to he a serious pest of rice and a mrnor pest of wheat by Fletcher (1917) Its f~t-stoutbreak was observed on sugarcane durlng 1953 in Bihar (Butani, 1955). Since then, M. separata outbreaks and serio~~sdamage have been reported in rice, wheat, sorghum, and millets In Asia and Australasla.

Distribution and outbreaks

The oriental armyworm is widely distributed in Asia and Australia between 45'N and 45 S latitude and 60 E to beyond 170W longitude, which includes 27 countries or islands ranglng from humld tropical to temperate reglons (Table 1 and Figure 1). It is recorded in Afghanistan, USSR, China, and Japan in the north, Western Australia and New Zealand in the south, and from Pakistan iri the west to Western Samoa In the east. Although there are no reports of tts occurrence from Malaysia, Timur. Tasman~a,and other Islands lying between the aforesa~d limits, it is likely to be present in all the countries/islands of this region. It is probably pi-esent all over lndla (Figure 2). However, it has not yet been reported from Gujarat, Sikk~ni,Pond~cherry, Goa, arid ~slaridsin the Arabian Sea and the Bay of Bengal. * Presur~taddress: Centre for Overseas Pest Research. Collpgr House. Wrights Larirt, Loridon W8 5SJ. I1.K . forrnf'rlv working at ICRISAT. 1 Now Viertenfa comota (MOOIP) (OauJoa WJON wqes 12 uel~lisa~ LZ sau~dd~llqdzz weulal~92 eau1n3 MaN pue ended 61 pue~leqlsz uelslyed 8~ ue~~elpz spuelsl YIO~JONLL tlssn sz pueleaz M~N91 (ulatsa~)eocues 22 e1uopale3 M~N Table 1. Geographical d~strlbut~onof the oriental armyworm, M separata Wlk

CountryITerr~toryilsland Rpfprpncp - .------Afqharrlstan Cott~rell(19541 Anon 119671 Grist and Lcver (19691 Australia Smith and Caldwell 119471 Common (19651 Anon (19671 Httchock (1969 and 1974) Broadley 11979a and b) Learmonth (1980) Banqladpsh Alam (1960,1962,1965 dnd 19671 Khan 11967) Anon 119671 Dean 119791 Burma Anon (19671 Chtna Chdo and Chen (19471 Chu etdl (19611 Lln eta/ (19631 Lln and Chang (19641 Anon 11967) Anon 11967~1Kao 1976) Anon (1977al Md 119791 Ytnchang (19801 FIJI Hinckley 11963) Anon (19671 Anon 119691 Lever 119691 Anon (19701 Honq Korig Anori 119671 lndla Andhra Pradesh, Assdm, Lefroy (19091 Fletchrr 11914 and 191 71 Ghosh (19241 Ramachandra Rdo (19241 Blhar Delhl, Haryana, H~machal, Hussaln 11935) Narayanan (19531 Butan) (19551 Puttardr~ahand Usrnan 119571 Kadam Jammu dnd Kashrn~r.Karnataka and Patel 119601 Kusbwaha eta/ 119641 Ayyar (19631 Avasthv and Chaudhary (1965) Kurdla, Maharashtra, Madhya Slngh and Slnha 119651 Kushwaha and Jaln 119661 Anon (19671 B~ndra119681 Sax~rid Prad~sh,Manlpu~ Orissa Punlab and Rawat 11968) Reddv 11968) Sr~vastavaand Sr~vastava(19691 Kdtlyar arid Patel Rajasthan Tdmll Nadu, Uttar (1969b dnd cl Prddhan (1971 I Nevlqund and Mdthad 11972a and b) Khan er a1 (1972) Pradesti dnd West 6enqdi Blndra arid Singh (1973) Rai (1973) Kukarn~er a1 119741 Pawar (19761 Bhatnaqar and Davles 11976 1978, and 1979a artd hi S~nghand Ral (19771 Goplnadhdn and Kushwaha 119781 Anon (19801 Chaudhary and S~ngh119801 Kushwaha eta/ (19801 Krtshnaswamy (19811 Slnqh and Manchanda 119811 Indonesia Mokrotovarov (19651 Anon 119671 Japan lwao 11956 1958a and b 1959a and b 1962,1963 1967a and b) Anon (1967) Gureva drid Kryzhanovskll (19681 Kanda ~ndNditn (1977, 1978, and 19791 Kampu~hra Anon (19671 Korea Anori 119671 Leper a1 (1970) Laos D~an(1978) New Caledonia Anon (19671 and (1978) NEWZealand Hambyn 119591 Anon (19671 Flower and Hrlsori 11974) Valunt~ne119751 Norfolk Islands Anon 119671 and Holloway (1977) Paklstdrr Anon 11967) and Baloch (1978) Pdliiia dnd New Guirird Anon 119671 and Barrett (19671 Ptiil~pp~ries Anori (19671 Cadapan dnd Sonchet 119721 Sabatt (North Bnrrirol Anori (19671 Sarnoa (West~rrl) Ariori (19671 Srl Lanka Anon (1967) USSR Klyuchko 119641 Anori 11967) Grist and Lever (19691 Talw,~n Tu dnd Lln 119661 Anon (19671 Ttidtldrid Anon 11973 arid 19741 V~rtridm Anon 11962 and 19671 Wect It ian Anon 119671

Sfr~ousoutbreaks of orlcntal armyworm hdd beer1 reported rn Ind~a(Ghosh, 1924 but an^, 1955, T~rumala Rao, 1956, Puttar~idriahand Usman, 1957 Slngh dnd Slnha 1965 Chaudhary dnd Ram7dn, 1967, Sarup eta/, 1969, Kalode et dl, 1971 Verma arid Khurana. 1971, Katlydr eta/,1972 Khan eta1 , 1972, B~ndraand Singh 1973. Butter eta/, 1979, Chaudhary and S~nqh,1980, Pdtel, 1980, S~righand Manchanda, 1981) Jdpan (Nagano ef a/,1971, 0ku and Kobayashl, 1974 dnd 1977), Clilna (Chao and Chen, 1947, Lln, 1963, Chrn eta/, 1965, Ch~n eta/, 1965, LI eta1 , 1963 and 1965, Chln, 1979), Australid (Sm~tharid Caldwell, 1947 and 1948, Common, 1965, Broadley, 1979a and b), and New ZedldIId (Hamblyn, 1959) It has also been reported In serlous numbers from Bangladesh (Alam, 1960 and 1965, Khan, 1967 Dedn 1979). FIJI(Anon , 1969, Lever 1969), arid Tha~land(Anon . 1973and 1974)

Factors result~ngIn outbreaks

The alrlval of mlgrant populat~ons1s one ~mpo~tantfact01 lead~nqto outbredks, they credte an rmbalancr between natural enpmlrs and host populat~ons,a? well ds br~rigirigmore ~ndlv~dualsInto d rpglort

Varlous causes of armyworm outbreaks have been suggected In Indla, heavy ralnfall followed by drouqht (Ghosh, 1924). floodinq (Butant, 1955, Puttarudr~ahand Usman, 1957), and trash mulch~rigof flplds have been My thrmna separata known to occur State or union boundartes- - - - Nat~onalboundar~es -. -

suggested as contributory fdctors In China, the temperdture In December affects the size and t~meof occurence of the infestat~ons Dry weather during January-Apr~l and h~ghhum~d~ty dur~ng July-August often results In serious outbreaks of subsequent generations (Anon, 1976~)The populat~onslze is ~nfluer~cedby relat~vehumidity (r h ) and ra~nfall(Ch~n, 1979) In Vietnam outbreaks co~nc~dewith heavy monsoon ralris and floods In Bangladesh the outbreaks have been preceded by drought, while In FIJIheavy ralns lead to outbreaks (Gr~stand Lever, 1969) It has been suggested that heavy manuring favours armyworm outbreaks In Japan (Koyama, 1966) In monsoon reglons, the dry spells possibly restrict the larval pardsites and pathogens The h~gherhum~dity and temperature there result In w~despreadovlposltlon and qu~ckdevelopment Crop losses

Crop loss estimates are not available from most of the countries llsted In Table 1 However, heavy losses have been experienced In Indla, Bangladesh, Chlna, Japan, Australla, and New Zealand In outbreak sltuatlons there may b? heavy to complete loss of the crop over extensive dreas The yleld losses are Influenced largely by the stage at whlch damage occurs and the gregarious behavlour of the larvae Durlng 1956-57 the armyworm damaged over 48000 hectares of pasture In New Zealand (Hamblyn, 1959) Alam (1965) reported 100% damage on rlce In Banqladesh Durlng 1955 armyworm damage occurred on 22000 hectares of rlce wlth a loss of over 40000 tonnes (Alam, 1960) Up to 36 6% sorghum heads were reported to be infested In Thalland durlng 1973 (Anon , 1973) Balasubraman~anet a1 (1975) reported 47-53% lnfestatlon In finger millet and S~nghand Manchanda (1981) reported 35-82 larvaelm' In rye Yleld losses of 1641 kglha have been reported In paddy (Katlyar and Patel, 1969b) and 10-30% In wheat (Chdudhary and Slngh, 1980) On different paddy cultlvars graln losses have been observed to vary from 200 to 500 kglha, I e 3 6 to 22 0% of the total yield (Pophall etal , 1980) Alam et a1 (1980a) reported detailed stud~~son the populat~onlddmdgelevels and yleld loss relatlonshlps in paddy They concluded that 15 larvaelh~llat boot and panlcle stages reduced the yleld by 37 98 and 92 92% respectlvely Wlth an Increase of 1 Idrvdlhlll the yleld decreased by 1 22 and 0 88 glhlll at thc boot and pdnlcle stdges respect~vely Wlth an Increase of 1% In ledf area consurncd \he yleld decreased by 0 07 and 0 88 glhill at boot and panicle stages respectlvely

B~ologyand seasonal actlvlty

Biology

The females lay 500-900 eqgs, with a recorded maxlmum of 1943 (Hamblyn, 1959 Hs~aet a1 , 1963) The eqg stage lasts for 2-7 days (Avasthy and Chaudhary, 1965) Larval development 1s completed In 14-22 days (Puttarudr~ahand Usman, 1957, Avasthy and Chaudhary, 1965, Cadapan and Sanchez, 1972, Slngh and Ral, 1977) The pupal perlod lasts for 8-9 days (Avasthy and Chaudhary, 1965, Slngh and Ral 1977) Total development take 26-38 days (Avasthy and Chaudhary. 1965, Cadapan and Sdnchez, 1972 Singh and Ral, 1977) Adults emerge at 20 00-23 00 h (Anon, 1976b) and survlve for 4-5 ddys (Avasthy and Chaudhary, 1965) Matlng takes place on the thlrd and ovlposltion on fourth ddy after emergence (Kanda and Na~to,1979) To be able to pair, some adults need to feed on sugar whlle others do so wlthout any feedlng (Mlyahara, 1978) Studles of the pest's nutrltlon have been conducted by Chin (1964), Chln et a1 (1964) and Quo and LIU (19641, and on hlstochemistry of sperrnatophore by Mu and Sun (1980)

The ~nsect'sdevelopment IS Influenced by temperature, humldlty, and host plants (LI etal , 1963, Wu etal , 1964, Chln etal , 1964 and 1965, Slnchaisrl and Sogawa, 1969, Hlrai, 1975, Dhallwal and Balns, 1978, Alam and Khatrl, 1980) The lower threshold temperatures for development of eggs, larvae, and pupde have been found to be 10 2-10 4, 7 4-7 6, and 9 9-10 1'C respectively (Slnchalsr~and Sogawa, 1969, and Wu etal, 1964) Chin etal, (1964) tound that at 32 C, egg hatchlng decreased at 20% r h , for proper development 100% r h was optlmum Th larvae d~dnot survlve at r h < 60% The rate of survlval of the flrst lnstar larvae was Influenced by the r h durlng egg stage At temperatures between 25 and 32 C, 90'16 of the eggs hatched at r h 22% None of the eggs hatched at 32-33 C (Chen etal , 19651 Survlval of pupae and prepupae Increased wlth r h (Chin eral, 1965) Fully grown larvae prefer high humldlty (Blndra and Slngh, 1973) The survlval rate is higher on hlghly manured crops (Koyama 1966) Tanaka (1976) and Patel (1979a) studled the feedlng rhythm of the larvae, feedlng normally occurs durlng the nlqht, the larvae hlde In cracks during the day Hamblyn (1959) observed that dur~ngan outbreak the larvae remalned on sunny slopes durlng the day time and fed malnly at nlqht Patel (1980) observed some unusual behavlour on a warm, dry, and sunny day The larvae (third to flfth Instar) marched towards walls and shady places lnslde the house at 14 00 h, some congregated on the mouth of earthen pots fllled wlth water, at 19 00 h they moved back to the orlglnal flelds

Adult populations can be monltored wlth llght traps (Spltzer, 1970, Persson, 1977) or molasses ba~tedtraps (Koyama, 1968) The rate of ovlposltlon can be determined by using dry leaves of sorghum for ovlpositlon (Tanaka et a/,1971) Spltzer (1970) observed actlvity during wlnters In New Zealand and concluded that there 1s no wlnter dlapause In the llfe cycle of thls Insect The larvae are present ~nthe fleld throughout the year (Blndra and Singh, 1973, and Anon , 1974) Atwal (1976) reported that the larval perlod may be prolonged to 88-100 days rn Punjab durlng winter The armyworm has been reared on an art~flcialdlet by Sato (1965), Hlral (19761, Kojlma and Nakayama (1979). Hattori and Atsusawa (1980). and on haycubes of Dactylisglomerata by Kanda and Nalto (1977) Phase var~atlon

Phase varlatlon IS an Important character~stlcof armyworm larvae Larvae reared under crowded cond~t~ons become darker than those reared In ~solat~on(Iwao, 1956, 1958a and b, 1962, 1967a and b) The factor respons~ble for the darken~ngmay result from the mutual st~mulat~onof larvae (Iwao, 1962) The black plqment (~ndolemelan~n) IS produced In the Integument when larvae are reared In crowded condlt~onsTransect~on of the oesophageal connectlves 21 h before the f~fthmoult led to the development of pale larvae (from larvae reared In crowded condltlons) (Ikemoto, 1971b) Larvae reared under crowded cond~t~onsassume darker colourat~onIndependent of temperature, or r h Such ldrvae Pat more food and develop more rap~dly,glvlng rlse to smaller pupae compared w~th those reared In lsolatlon The darken~ngIS Induced by a llmlted food supply and teduced by subsequent ~soldtlon Crowded larvae are more tolerant of an inf favour able food supply and are more ~rr~table(Iwao, 1956, 1958a and b, 1959a and b, and 1962) CI-owd~ngdoes not affect ovlposltlon, fecund~ty,or adult colourat~onbut adult longevity IS extended The emerglng from crowded larvae have a lower water content (Iwao, 1959b) The larvae of M loreyi develop at a slower rate under crowded cond~tionsand do not become very dark and the adults from crowded cultures are short l~ved(Iwao, 1962) The black larvae of M separata are more responsive In an illuminated held (Iwao, 1963) The respiratory rate of the bldck larva? 1s hlgher than that of the pale larva? (Sh1ba7ak1and Ito, 1969) and black larvae w~thstandstarvation much longer because they havp a hlgher fat content than pale larvae (Iwao, 1967b) The pale larvae hdve more urlr ac~din the integument and less ~nthe hdernolymph than the black larvde lmmediately after moult~nqto the s~xthlnstar (Ikpmoto, 1973) At the time of pupation, the phenolox~dasp actlv~tybecomes higher in bldck larvae thdn the pale larvde (Ikemoto, 1972) lkemoto (1965) found that populat~on dens~tydffects catalase actlvlty

Plgmentatlon of the larvae IS controlled by a hormorial factor (Ogura 1975a) orlglnatlnq flom the brdln and sub oesophageal gariql~on(Ogura, eta1 , 1971, Oqurd, 1976) Some st~miilat~on1s transm~tted postrrlorly through the oesophageal connectlves to promote the release of a hormone or hormones from thp sub oesophageal gangl~on causlng melanlzatlon (Ogura et a/, 1971) Sub oesophageal gangha cultured in vttro for 10 days cont~nuedto secrete a mrlanlzat~onand reddlsh colourat~orihormone (Ogura and M~tsuhoshi 1978) Implantation of sub oesophageal ganglla of Bomhyx mor~L pupae Into Isolated dbdomens of armyworm larvae Induced cuticular melanlzat~onGanglld of pupae destined to lay d~apausregqc produce more lritensive melan17at1on(Oyura and Salto, 1973) However, larval oesophageal ganql~a,braln, and 1st thorac~cqdnglla of B mori do not Induce rnelan~zdt~on A llgature behind the thorax of crowded or Isolated larvae causes black or redd~shbrown plgmentdtion In the driterlor part after larval ecdys~s Extlrpatlon of braln, corpus allatum, or sub oesophageal ganql~onreduce the degree of melanizat~onIn crowded larvae (Ogura, 1975b),wh~le the~r ~mpldntat~on rrsults In black plgmentatlon Thorac~c and abdom~nalgangl~a ~mplantatlon Induce l~ghtcolouratlon (Ogura and Sarto, 1972)

The ~njectlonof adenos~nemonophosphate (AMP) d~butyl,cycllc AMP, cycllc gudnoslne monophosphate (GMP), or d~butylcycllc GMP ~ntoIsolated abdomens of flfth lnstar ldrvaecause darken~ngof body ~olorlnject~on of melanlzat~onand reddish brown coloratlorl hormone (MRCH), together w~ththeophyll~ne, causes more Intense melan~zatlonthan MRCH alone Cycl~cAMP may play a major role as an intercellular messenger of MRCH (Matsumoto eta/, 1979) Integuments cultured /n vivo show melan~zationwh~n host moult Into the flnal lnstar Cultured lrlteguments are respons~blefor thr MRCH se~retedfrom the co cultured sub oesophageal ganglla (Ogura and M~tsuhosh~,1979) lkemoto (1971a) studred the non proteln fohn posltlve components In haemolymph In relat~onto melan~nsynthesis and LIUand Fenq (1965) studled the blood sugar nletabol~sm

Extensive studlps carr~edout In Chlna and Japan suggest that moths of thls armyworm mlgrate, In Chlnd most research was devoted to th~saspect dur~ngthe per~od1961-65 (Ma, 1979) Moths move to a more su~tablehabltat from unfavourable env~ronmrnts,drid ~t has been concluded that the larvae do not undergo d~apauseThe nupt~al fl~ghthas been suqqested as the ~n~tialcause of m~gratlon(Quo era/, 1963) Hwdng and How (1966) suggested that the per~odbefore sexudl maturlty IS the optlmum tlme for long d~stancemlgratlon Lln (Chdng Shan) (1963) sugges tad that the moths follow asc~nd~ng,trans, and descending movements In Ch~na Lln and Chang (1964) proposed a model of the regular~t~esof outbreaks and long d~stanceseasonal m~grat~onSuch movements have been attr~buted to cyclonlc centres, cold front areas, and thunderstorms LI et a/ (1963 and 1965) considered that moths from South Chlnd are asourceof early sprlng populations The date of flrst appearance of moths co~ncldesw~th southerly wlnds w~tha ,70'% coinr~dence(Lln et a1 , 1963) Increases In the number of moths occur on the same day at all locations (Lln et dl, 1963 Koydma 1970) Outbreaks begln In the south and then appear In the antral and north eastern reglons (Chen era1 , 1965), return mlgratlon occurs In summer or autumn Adults migrate to Chaoyang In May and In the opposite dlrectlon in October (Anon., 1976~).Mlgrant moths have also been Intercepted on ships voyaging between the Gulf of Chili and the Yellow Sea (Hsia eta/., 1963) and 500 km away from the mainland of Japan during 1968 (Asahlna, 1968). Captures of moths wlth dyes have Indicated that moths can fly 600-1400 km (Grist and Lever, 1969).

In Japan outbreaks have been attributed to migrating moths carrled by south-easterly wlnds (Oku and Kobayashl, 1974 and 1978; Oku and Koyama, 1976). Oku and Kobayashl (1977) suggested the possibility of armyworm migration from Chlna. Adults may migrate eastwards wlth the alr currents (Oku and Kobayashl, 1977). Nagano et al. (19721 reported a method of forecasting outbreaks In Japan. In Fijl, all but three of the eleven outbreaks over the perlod 1939-65 occurred during the February-March period when rarnfall exceeded 8 75 mm (Lever, 1969). There are no reports on the pest's posslble migration In India, Australia, New Zealand, and other countries.

Natural enemies

A number of natural enemips have been rtbcorded attacking the larvae and pupae (Table 2). In nature, up to 89% parasltlsm of the larvae has been reported (Katlyar and Gargav, 1971, Cadapan and Sanchez, 1972). Apanteles roficrus (Hal.) 1s the most important paraslte and ~t has been found to parasitise up to 50% of larvae In Chlna (Anon. 1976c) and up to 86.7% In lndla (Butter, 1978). A. ruficrus has been successfully used in biological control in New Zealand (Simmonds, 1976; Simmonds and Bennet, 1977). Katlyar and Gargav (1971) reported up to 32% pupal parasitism.

Nuclear polyhedrosts virus (NPV) has been reported to cause extensive mortality of the larvae in nature (Neelgund and Marhad, 1972b, Tsai, 1965, Battu et a/., 1977). The NPV from this armyworm does not Infect the silkworm, Anthereae mylitta (Dru ) (Dhaduti and Mathad, 1979), and Indian honey bee Apis cerand indica F. (Dhadutl and Mathad, 1980) The gastric juices of albtno rats inactivate the NPV (Kumar and Mathad, 1979).

Pheromones

The males of M separata have a scent brush on the flrst abdominal sternite, females have sacklike membranes between abdominal segments VII and VIII (Quo eta/., 1963). In males, the major male scent component 1s benzaldehyde, which functions as an arrestant (Clearwater, 1972). Hirai (1977 and 1980a and b) found benzyl alcohol In large amounts in male scents He also reported the presence of benzaldehyde, benzolc acld, and butyl alcohol. Benzyl and phenylether have been reported as the storage materials whlch release the aromatic pheromone (almond odour) upon hydrolysis (Clearwater, 1975). The metabolism probably occura In two types of vesicles (Clearwater and Sarafis, 1973). Hlrai (1980b) could not detect any chemtcals from the halrs on male genitalla. Males stimulated by the presence of females produced smaller amounts of benzaldehyde and larger amounts of benzyl alcohol than the virgin males. Durlng courtship, only benzaldehyde was detected In the alr. In females, (2)-1I-hexadecanol and (2) 1l.hexadecenyl acetate have been isolated from the abdominal tip. These components in 9:l and 1 :l ratlos have shown phermonal actlvity In the laboratory (Takahashi et 81.. 1979; Sato et a/., 1980). Sato et a/. (1980) obtained hlgh catches of males in a greenhouse with a mixture of acetate and alcohol components in 9.1 or 4.1 ratlos with amounts in the range of 0.1 to 1.0 mg. The attractlveness of the mixture was equal to ten females.

Host range and host plant resistance

Host range

The oriental armyworm ~sapest of Gramineae, it has also been reported to feed on plants belonging to such dlverse botanical groups as rape, sugarbeet, hemp, nut grass, flax, and pea. It has been recorded feedlng on 33 plant species and some unspecif~edgraminaceous grasses (Table 3). The major hosts include important cereal crops; flnger millet (E, coracana), barley (H. vulgare), rice (0. sativa), pearl millet (P, arnericanc~rn),sorghum (S.bicolor), wheat (T. aestivurn), and (Z.mays). Of the 33 reported host plants there are 1 Chenopod~aceae,3 Cruciferae, 1 Cyperaceae, 23 Gramlneae, 2 Legumlnosae, 1 Linaceae, 1 Malvaceae, and 1 Solanaceae. Table 2 Natural enemles of the or~entalarmvworm. M seaarata (Wlk ) --- - Natural enemy Reference

Bethylldae Hymenoptera Parasterola sp Avasthy and Chaudhary (1963 1965 and 19661 Bombyllldae D~ptera Anthrax sp Saxena (19651 Bracon~dae Hymenoptera Apan teles flavrpes I Cam I Rao 119691 Apanteles glomera rus ( L 1 Khan et a1 119721 Apanteles parbhanr Rao Rao (1969) Apanteles ruftcrus (Hal I Khan (19461 Bhatnagat (1948) Rao (19531 Rao (1969) Kat~yarand Rawat (1972) Mohyudd~nand Shah (1977) B~ndraand Slngh (1973) Anon (1975 1976a and 1977bl H~ll(19771 Cumber eta1 (19771 Butte (19781 Roberts 119791 Apan teles sp Chao and Chen (19471 Gop~nadhanand Kushwaha (19781 Reddy and Dav~~s(1979) Drsophrys sp Katlyar and Rawdt (1972) Meteorus sp Chao and Chen (19471 Rogas (Rhogas) fuscomacu/atus Ashm Chao and Chcn 11947) Rogas sp Kat~yarand Rawdt 11972) Bhdtnagar and Davles (1979bl Chalc~d~daeHymenopterd Saxena 11965) Katlyar and Gargav 11971) D~rmest~daeColeoptrra An threnus flavr~esLPC Kushwaha and Gop~nadhan(1972) Gop~nadhanand Kushwaha (19781 Encyrtldae Hymenopt~ra Oencyrrus sp nr major Ferr~e~e Goplnddhan and Kushwahd (19781 Eurytoma sp Kushwaha and Goplnadhdn (1972) Gop~rladharland Kushwaha (19781 Euloph~dac Hymenoptera Tetrdstrchus sp Kat~yarand Gargav (19711 lchneumon~dae Hym~noptpra Barrchneumon solrtarrus Mar I Katlyar dnd Gargav (1971) Cratrchneumon sp Rao et a1 11969) Enrcosp~lussp Avasthy and Chaudhdry 119651 Saxena 119651 ltoplectts narangae Ahm Rao et a1 (1976) Melobor/s leucanra n sp Kus~gemafr 11972) Metoprus rufus Cam Bhatnagar and Davles (1979b) Metoprus pulchrrpes Cam Katlyar and Gargav (1971)

Drchrocheles sp Phor~dae Dlptera Meqaselta sp Gop~nadhanand Kushwaha 119781 SarcophdgldaP Dlptera Sarcophaqo orrentolotdes Senlor Wh~te Blndra and Slngh (19731

Telenomus gudngdong~ns~sn sp

Actra montrcold Mall Cherlan and Ananthdr~drayanan(1941 ) Grcd~aprrma Baranov Kdtlyar and Garqav (19711 Compstlura sp Broadley 11979b1 Conephalrd (Pseudogonra) crrtnerascens Rond Avasthy and Chdudharv (19651 Conephalra (Pseudogonral rufrfrons (W~ud1 Kdt~yarand Gargdv (1971) Cuphocera varrd (F I Cherlan and Andnttianarayanan (1941) Cuphocera sp Broadlcy (1979b) Dolrcholon pdrddoxum Brdnd & Berg Avdsthy and Chaudhary (1965) Exor~stdfdllax IMq I Khan et a/ (1972) Katlyar and Rawat (19721 Gonrophthalmus ausrralrs (Baranov i Broadley (1979bl Lrnnaemya vulptnordes Baranov Avasrhy and Chaudhary (19651 Perrbaea sp Broadley (1979b) Table 2-cont~nued

Natural enemy - Reference Pseudopenchaeta anomala (V~llen) Kat~yarand Gargav 11971 1 Sturrnla roconsp~co~des(Baranov Cher~anand Ananthdnarayanan (1941 I Thecocarcel~aoculeta (Baranov ) Kat~yarand Garyav (1971)

2 Predators (a) B~rds Acrldotheres tr~sr~sL Avasthy and Chaudhary (1956) Chaudhary dnd S~ngh119801 Acndorheres fuscus, L Gr~stand LPver (1969) Bubulcus rb~sL Avasthy and Chaudhdry (19651 Chaudhary dnd S~ngh(1980) Coraclas benghalens~s L Chaudhary and Slngh 11980) Corvus splcndens L Blndra and S~ngh(1973) Dlcrurus mdcrocercus L Chaudhary and S~nqh11980) Passer domest~cusL Blndra arid Slngh 119701 Tandka 11975) Ih) Cdrdtjldae Colcoptera Calosorna ausfrdlls Hop Smlth and Caldwrll (1948) Carabus sp Chao and Cherl (1947) Form~c~daeHyrnenopt~ra Khan and Sharma (1972)

Anedrallus splnldens F Tenebr~onldae Coleoptera Kat~yarand Patel 11969) Vpsp~dae Hym~nopterd Hlrose and Kakdg~11980) H~roseand Takay~(19801 Gr~stand LPVQ~(1969) Ic) Toads Bufo sp Avasthy dnd Chaudhary (1965)

3 Pathogens (a) Bdcter~a Alarr 11967) Battu era1 (1971 1 Bindra and Srngh 119731 Shlng era1 (1979) Gop~nddhanand Kushwaha 119781 Bac~lluscereus Kushwaha and Gop~nadhari11972) Serratla rnarcescens Rangaraldn et dl (1968) Str~prococcussp Kushwdha and Gopinadhan (1972) it11 Fung~ Enfomophrhora sp Cadapan and Sanche7 11972) M~tarrhlr~urnan~sopllae G~lstand Luvcr (19691 Nomuraea rlley~ Broddlry (1979b) Sp~car~afurnosoroseus Tscng shenq et dl 11 965) Sp~canasp Cadapan drid Sanchu (1972) (c) V~ruses Vlrus dlseds~ Alam (1962) Nuclear polyhedros~s(NPVI Neelgund dnd Marhdd 11972b) Neelqund (19751 Manlunath and Mathad (1918 and 1979) Nrrlgund and Mathad (1978) Tsal (1965) Battu eta1 11977) NPV of Hereronychus arator IF ) Longworth and Arch~bald(19751 NPV of Prodenla lrturd IF Hwanq arid Dinq 119751 Table 3. Host range of the oriental armyworm, M. separata

Common name Scientific name Reference

Chenopodiaceaa Sugarbeet Beta vulgaris Khan and Sharrna (1971). Crucifereae Rape Brassica campestris Blndra and S~ngh(19731 Chinese cabbage Brassica campestris var. capitata Grist and Lever 11 9691. Turnip Brassica rapa Grlst and Lever 11969). Cyperaceae Nutgrass Cyperus rotundus KalodeetaL (1971 I Gramineae Oat A vena sativa Blndra and Singh (19731. - Brachiaria muta Grist and Lever (1969). Bermuda grass Cynodon dactylon Kalode et a/. (1971 I; Blndra and S~rrgh(19731 Jungle rlce Echinochloa colonum Gargav et a/. (1972). Japanese barnyard m~liet Echinochloa crusgalli KalodeetaL (1971). Finger millet Eleusine coracana Rai (19731; Balasubraman~anetal. (1975). Goose grass Eleusine indica Kalode er a/. (1971) Eriocaulon sexangulare Gargav et a/. (19721. Grasses Nagano eta/. (19721; Anon. (19781; Kanda and Na~to11978); Broadley 11979a). Barley Hordeum vulgare B~ndraand S~ngh(19731; Anon. 11976~1. R~ce Hlnckley (19631; Wu 119631; Pschorn-Walcher (19641; Mokrotovarov (1965): Anon. (19691; Kulshreshtha etal. 119701; Chakrabarty eta/. (1971I; Kalode el a/. (1971); Verma er a/. 11971 1; Q~wakar(1972 and 1975); Patel (19721; Anon. (1976dl: Pawar (1976); Slngh and Ral 119771; Dean 11978); Rawat and Slngh (1979); Chu 119791:Gangradeetal. (19801. L~ttlemillet Panicum miliare GIIS~and Lever (19691. Panicum proliferum Kalodc erab (1971 I. Panicum scrobiculatum Kalode et a/. (1971 I. Panicum serigerum Grlst and Lever (19691. Pearl m~llet Pennisetum americanum Kalode et a/. 11971 1; B~ndraand S~ngh(19731; Anon. 11976d). Sugarcane Saccharum officinarum Butani (1955); Saxena (1965); Chaudhary and Ramzan (1967). Rye Secale cereale Slogh and Manchanda (1981 1 Johnson grass Sorghum halepense B~ndraand Slngh (1973); Atwal (19761 Sorghum Kalode etal. (1971 I, Neelgund and Mathad (1972b); Anon. (1973): Doughton (19741; Blndra and Rathore (19651, lronslde 11979). Foxtall mlllet Setaria italica Gr~stand Lever 119691. - Setaria glauca Kalodeeral. (1971). Wheat Triticum aesttvum Chu era/. (1961 1; S~nghand S~nha11965); Saxena and Rawat (19681; Blndra and Singh (19701; Bhattacharjee and Gupta (1971); Verrna and Khurana (1971 1; Blndra and S~ngh119731; Anon. (1976~1;Chiang (19771. Zea mays Anon. (1962 and 1976d); Helson (19701; Rajagopal arid Channa~ Basavanna (1975 and 19771: Buxton (19761. Leguminosae Pea Pisum sativum Sharma etal. (19701. Beans Grlst and Lever (1969). Linaceae Linum usitatissimum Tu and Lln 119661 Malvaceae Hemp Cannabis sativa Solanaceee Tobacco Nicotiana rabacum Grist and Lever (1969). Host plant reststance

As stated above, M separata is polyphagous, dnd crops such as sorghum, millets, rice, and maize are potential hosts (Kalode eta1 , 1971) It has been reported to prefer triple gene dwarf wheat varieties (Verma and Khurand, 1971) The larval population is Influenced by pldnt characteristics such as plant helght, number of tillers, and canopy Significantly lower numbers of larvae have been observed In the wheat genotypes UP-2002, Raj 1381, and WL-1533 (Butter eta1 , 1979) In rice the development IS slower on seedlings than on older plants (Kdlode eta/, 1979) Lines R35-2750, R35-2752, R-2384, RP-9-4, RPW6-12, W-13400, RPW6-17, and Surekha have been reported to suffer < 10% yleld loss compared with 22% in BKN-680-52 (Pophali eta/, 1980) The differential susceptibil~tyof maize qenotpes has been observed In Thailand (Anon , 1973) In sorghum, Rangarajan etdl (1974) reported genotypes R-16, R-24, 604, and CS-3541 to be less susceptible ( < 16 3% damage) The hybrlds were more susceptlble than the varieties Kulkarni and Ramakrishna (1975) found that the genotypes line-141, E-303,

NJ-1953, SB-803, 1-744,296,604, and E-302 manifested L 6% damage compared with 47 9% In Type 302 Hybrids 648 AX1075, SPH-4, 2077XCS 3541, and CSH-1 were also less susceptible in comparisori with SPH-3 Lines SB-412, SB-803, SB-461, and CS 3541 have also been reported to be less susceptlble (Kulkarni era1 , 19781 The relative concentrations of cyanide in ledves do not affect th~cxtent of armyworm damage (Woodhead et a1 , 1980)

Chem~calcontrol

In fleld trials a number of single ~nsecticidesand mixtures have been report~dto control this insect They include BHC and aldrln (Butani, 1955), 1 1 or 1 2 duct mixture ot DDT (5%,)and BHC (0 5%) (Wei, 19591, dieldrin and DDT (Hamblyn, 19591, trichlorpon (2 5% dust) (Chariy and Li, 19641, BHC and DDT dusts (Btndra and Rathore, 1965), BHC dustlng (Saxena and Rawat, 1968) enclrin (0 02%). phosphamidon (0 03%) and endosulfan (0 05%) (Purohit et a1 , 1971 ), chlorpyrifos (0 05%) (Kalode rt dl , 1972), rthyl parathion (005%) (Gargav and Katiyar, 1972), ultra low volume sprays of 96% malathiori (0 85 Iitreiha) (Slnqh drld Maw, 1972), trichlorphon, fenthion, and methyl parathion (Balasubramanian, 19731, trichlorphon, endosulfan, dnd DDT (Hitchock, 1974). I~ptophos(Mackay and O'Connor, 1976). BHC (lo0?)carbaryl (5'XI, and endosulfan (4%) dusts (Kishore and Jotwanl, 1976), quinalphos and methyl pardthlon (0 5 kqiha) (Patel eta/,1979) and carharyl, BHC, and dichlorvos (Singh er di , 19801

Methyl parathlon, mevinphos, nalcd, EPN d~ct~lorvosethyl parathion, isodr~n,phenthoate, trlchlorphon. fenitrothion, diazinon, and dzinphoc ethyl have been found to be mow toxlc than DDT ur~derlaboratory conditions (Sarup et a/, 19691 Thr addition of benzyl dlcohol and cresol to gammd-BHC as a carrler have been found to increase the rate of penetration but not the final mortality of sixth instdr larvae Urethdrlr, iodo acetic acid, and acetophenone synerglse the action of qamma-BHC Excltatlon of ldrvae wlth qamma BHC results in a rapld kill by DDT The qredter reslstdricr of sixth lnstar larvae to BHC has been attr~butedto a detoxification mechanism (Chanq, 1964) Slnchaisrl er a/ (1977) reported the LDqovalir~s of four organophosphorus insect~cidesto fifth lnstar larvae Methyl parathion was the most toxic, however, phenthoate, which was least toxlc, showed five times hlgher anti cholinesterase activity than methyl parathion Mlxtures of Bartllr~sth~tnnqtensis var gdllenae and DDT d~dnot produce appreciable etfects on the ldrvae (Tsyan ~t a/, 1963)

In China the antibiotic Blastidin has been found to br toxic wh~nappllrd to the leaves of food plants (Anon, 1977a) Admlnistratlon of catharidin producrd pathological chanqes (Lu, 1964) Cartap (neurotoxin der~vedfrom marlneworm, Luhr~conereisheteropoda) is toxic to the larvae (Sakal rtai , 19671 Cheng ~t a/ (1964) reported the effects of gamma BHC on haemolymph Treatment of adults wlth thiotrpa (5%) reduced egg laying by half, and none of the eggs hatched, only a small perrentaqe of egqs tailed to hdtch whrii treated males were mated with untreated females (Chang and Chiang, 1963) Trlphenyl tinacetdte hds been teported as an dntlfeedant and IS toxic to the larvae (Math~lrand Saxena, 1972) Patel (1979b) studied the performance of sprayprs and water volumes uslng fenitrothlon at 1 kg allha He found that a hand compression sprdyer was significantly superior fol the control of larvae (95%) than a mist blower (56%) Knapsack and foot sprayers alqo performed well The population reduction at low, medlum, and hlgh volumes was 83, 83 dnd 68% resprrt~vrly

Acknowledgements

Our thanks dre due to Drs W Reed, K Leuschner, A B Mohammed, and R P Thdkur for their vdluable comments and to Mr I Krlshna Murthy for typing the manusct ipt Bibliography

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CHENG (Chen-heng), LIU (Zhl-y~ng),and ZHONG (Deng-qulng) (1964). Effects of l~ndane(gamma-BHC) on some crlterla of the haemolymph In In Leucanla (Pseudaletla) separata Act Entomologrca Srnrca 13 897-899 CHERIAN, M. C., and ANANTHANARAYANAN, K. P (1941) Clrphls alb~str~gmaH., a pest of grasses In South Ind~a.Journal of the Bombay Natural Hlstory Soclety 42 61 1-616 CHIANG, H. C. (1977). Pest management In the People's Republlc of Ch~namonltorlng and forecastrng Insect populat~onsIn rlce, wheat, cctton and malze. FA0 Plant Protection Bullet~n25 1-8. CHIN, C. (1964). Studles on the nutrltlon of the armyworm (Pseudaletla (Leucanla) separata) (Walk.).The ~nfluencc of d~etaryand env~ronmentalwater on the larval growth Act Entomolog~caSlnrca 13. 659-670.

CHIN (Chun-teh), WE1 (D~ng-ye),and WANG (Tsong-Shen) (1964) Stud~eson the nutrltlon of armyworm (Pseuda- letra (Leucanla) separata) Wlk.). The feed~ngand utlllzat~onof sugars by adult moths. Acta Entornolog~caSlnrca 13. 773-784.

CHIN, S , HO, C., and MA, S. (1964). The relat~onsbetween hurnld~t~esrn the env~ronmentand the rates of survlval and development of the armyworm, Leucania separata Walker. I. Eggs and frrst lnstar larvae. Acta Entotnolog~ca Slnrca 13. 835-843.

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DHADUTI, S G , and MATHAD S B (1979) Effect of nuclear polyhedrosls vlrus of armyworm. Mythirnna (Pseudaletia) separntd on the tasar s~lkwo~m,Antheraea rr~yl~ttaCurrent Science 48 750-751

DHADUTI, S G , and MATHAD, S B (1980) Effect of nuclear polyhedrosls vlru of th~armyworm, Mythinina (Pseudaletia) separata on colonlrs of Apic cerana indica Journdl of Agriclllturdl Research 19 77-78

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Internat~onalCrops Research Institute for the ~ern~-~r?dTroplcs Dat~nrhnraa A D lnrla= IKEMOTO, H (19651, Larval dens~tyand catalase actlvlty In the armyworm Leucania (Pseudaletia) separata Japanese Journal of Applied Entomology and Zoology 9 60-62 IKEMOTO, H (1971a) On the non proteln fol~n(Lowry) posltlve components In the larval haemolymph of the armyworm, Leucanla (Mythimna)separata In relatlon to melanln synthesis Botyu Kagaku 36 59-65 IKEMOTO, H (1971b) On the black plgment of larval Integument of the armyworm, Leucanla separata Botyu Kagaku 36 128-131 IKEMOTO, H (1972) Phenoloxldase In the armyworm, Leucania separata Walker Botyu Kagaku 37 43-48 IKEMOTO, H (1973) The relat~onshlpof the larval denslty to the urlc ac~dcontent In the armyworm, Leucania separdta Walker Botyil Kagaku 38 40-42 I RONSI DE, D A (1979) l nsect pests of graln sorghum Part 2 Oueensland Agr~culturalJournal 105 25-28 IWAO, S (1956) Phase varlatlon In true armyworm, Leucania unipuncta Haworth (I!Researches ln Population Ecology 3 60-78 IWAO, S (1958a) Effect of populatlon densitv on the larval growth pattern Japanese Journal of Applied Entomology and Zoology 2 237-243 IWAO, S (1958b) Some analysls on the effect of populatlon denslty on larval coloration and growth In armyworm, Leucanla unipuncta Haworth Physiology and Ecology 8 107-1 16 IWAO, S (1959a) Phase difference In the range of food tolerance of the flnal lnstar larvae Japanese Journalof Applied Entomology and Zoology 3 164- 17 1 IWAO, S (1959b) Comparison of the manrfestatlon of the denslty dependent varlabllltles In Leucania unipuncta Haworth, L loreyl Duponchel dnd L placida Butter (Lepldoptera Noctuldae) Japanese Journal of Ecology 9 32-38 IWAO S (1962) Studles on the phase varlatlon and related phenomenon In some Lep~dopteraInsects Memoir of the College of Agriculti~re,No 84 Kyoto, Japan pp 80 IWAO, S (1963) Some behav~ouralchanges associated wlth phase varlatlon In the armyworm Leucania separata Walker I Reactlon of larvae to mechanrcal stlmull Japanese Journal of Appl~edEntomology andZoology 7 125-131

IWAO, S (1967a) Differences In the llght reactions of the armyworm. Leucanlaseparara Walker, In relatlon to thelr phase status Nature, London 21 3 941 -942 IWAO, S (1967b) Resistance to starvatron of pale and black larvae of the armyworm, Leucania separata Walker (Lepldoptera Noctuldae) Botyu Kagakil 32 44-46 KADAM, M V , and PATEL G A (1960) Pests of cereals (1 ) pp 25-39 In Crop pests and how to fight them Drrectorate of Publlclty, Government of Maharashtra Bombay, lndla

KALODE, M B , RAO, P S P , and VARMA, A (1972) Toxlclty of some modern lnsectrcldes to paddy cutworm, Pselldaletia separata Walker Indian Journal of Entomology 34 84-85 KALODE, M B , VARMA, A, YADAV, C P , and ISRAEL, P (1971 1 Feedlng reactlon of armyworm (C~rphis compta Moore) larvae fed on some cultivated and wrld plants lildian Journal of Agricultural Sciences 41 568-572 KALODE, M B , VARMA, A, YADAV C P , and ISRAEL, P (1979) Feedlng behavlour of paddy cutworm, Pseudaletia separata Walker In relatlon to drfferent aqes of rlce plant Indian Journal of Entomology 39 291- 292

KANDA, K , and NAITO, A (1977) Rearlng of the armyworm, Leucania separata Walker on haycubes Applied Entomology and Zoology 12 75-76

KANDA, K , and NAITO, A (1978) Larval denslty of the orlental armyworm, Leucaniaseparata Walker (Mythrrnnd separata Walker (Mythimna separata (Wlk )) caustng the complete destruction of grasses (In Japan) Japanese Journal of Appl~edEntomology and Zoology 22 205-207

KANDA, K , and NAITO, A (1979) Behav~ourof orlental armyworm moth, Leucaniaseparata Walker, from emergence to ovlposltlon Japanese Journal of Applied Entomology andzoology 23 69-77

KAO, W T (1976) Studies on the phototact~cbehavlour of noctuld moths A speclal lnterrelat~onshlpbetween the response of compound eyes and the behavrour Acta Entomologica Sintca 19 59-62 KATIYAR, 0 P , and GARGAV, V P (1971) New record of some parasrtes of rice armyworms JNKVV Research Journal 5 56

KATIYAR, 0 P , and PATEL, R K (1969a) Trrbolrum sp An enemy of pupae of rlce armyworms Screnceand Culture 35 482 KATIYAR, 0 P , and PATEL, R K (1969b) A note on est~matlonof grain losses to paddy by armyworm lnternat~onalRrce Commun~catronNewsletter 18 26-27 KATIYAR, 0 P , and PATEL, R K (1969~)Rrce armyworms In Chhar~sgarh Pesticrdes 3 39-40 KATIYAR, 0 P , and RAWAT, R R (1972) Larval parasltes of Psei~daletraseparata (Walker) Journal of the Bombay Natural History So~rety69 212-213 KATIYAR, 0 P , LAL, L , REDDY, A R ,and MUKHARJI, S P (1972) Outbreak of rrce armywormsat Varanasr, Current Science 41 579 KHAN, D U (1967) Use of a~rcraftIn agr~culture pp 130-132 In Proceedings of the Agr~culturdlSymposlurn, 13-14, May. 1966 Dacca Atom~cEnergy Ceritre KHAN, M Q (1946) Lrfe hrstory of cdstor sem~loopersIn Hyderabad (Deccan) Indian Joi~rnalofEntomology 8 111-115 KHAN, R M , and SHARMA, S K (1971) Some new pest records of sugarbeet In lndra Inclran Journal of Entoinology 33 105-1 06

KHAN, R M , and SHARMA, S K (1972) Cataglypsrs Dicolor Fdb (Hym Forrnlc~dae)as a predator on few Noctu~dlarvae Madras Agrrcc~/rura/Journal 59 192

KHAN, R M , SHARMA S K , and GUPTA, B M (1972) Appedrarlces of armyworms on wheat crop In outbreak form and the~rcontrol Madras Agrrcultural Journal 59 663-665 KHASIMUDIN, S (1978) Courtsh~pand Matlng behav~ourof the Afr~canarmyworm, Spodoptera exempta (Walker' (Lep~doptera Noctu~dae)Bulletrn of Entomologrcal Research 68 195-202 KISHORE, P , and JOTWANI, M G (1976) Chem~calcontrol of earhead worms on sorqhum Entomologrst's Newsletter 6 65 KLY UCH KO, Z F (1964) Qrphis unrponcta In the Ukra~neZasch~ta Rdstenli ot Vredrteler Boleznei 10 44

KOJIMA, L , and NAKAYAMA, 1 (1979) Rear~ngof varlous lepldopterous larvde on artrflc~aldret of rdentrcal composltlon Japanese Journal of Applred Entomology and Zoology 23 261 -263

KOYAMA, J (1966) The relat~onbetween the outbreak of armyworms (Leucanla separata Walker) and the r~chly nitrogenous manured cult~vat~onof rlce plants Journal of Appl~edEntomology andzooloqy 10 123-128

KOYAMA, J (1968) On the capture records of armyworm moths, Leucanra separatd Walker by molasses traps Japanese Journal of Applred Entomology and Zoology 12 123-1 28

KOYAMA, J (1970) Some consrderat~onson the chronolog,cal records of outbreaks of the armyworm (Leucanra separata Walker) Japanese Journal of Applred Entomology and Zoology 14 57-63

KRISHNASWAMY, N (1981) aeware of outbreak of earcuttrng caterp~llaron paddy In Assam (Mythrrnna (Pseudaletra) separata) (Wdlk ) (Nocturdae Lep~doptera)Central Plant Protectron Stat~on,Gauhatl, Assam (D~rectorateof Plant Protect~on,Quarant~ne and Storage), M~n~stryof Aqrlculture and Irr~gat~on,Government of lndla pp 10

KULKARNI. K A, and RAMAKRISHNA, V C (1975) Stud~eson the suscept~b~l~tyof sorghum lrnes and hybr~ds to the armyworms (Pseudaletra separata Walker) Sorghum Newsletter 18 33

KULKARNI, K A, NAIDU, B N B , and THIMMAIAH, G (1974) Seasonal ~nc~denceof sorghum armyworm (Pseudaletra separata Walker Sorghum Newsletter 17 46

KULKARNI, K A, PARAMESHWARAPPA, R , and KAJJARI, N B (1978) Screening of h~ghy~eld~ng sorghum varletres to armyworm (Mythrmna separdta Walker) Mysore Journal of Agr~culturalScrences 12 572-574

KULSHRESHTHA, J P , KALODE, M B , and VARMA, A (1970) Paddy cutworm (Pseudaletraseparata Walker) and armyworm (Clrphrs compta Moore) as serlous pests of h~ghyleld~ng var~etles of rlce Oryza 7 143-145

KUMAR, M Y , and MATHAD, S B (1979) Effect of alb~norat gastrrc julce on the virulence of fithrmna (Pseuddletra) separata nuclear polyhedros~svlrus Current Scrence 48 872-873 KUSHWAHA, K S , and GOPINADHAN, P B (1972) A note on blotlc agents regulating field populations of the armyworm, Pseudaletia separata Walker ( Lepldoptera) University of Udalpur Research Journal 10 78-79 KUSHWAHA, K S , and JAIN, S K (1966) On some forage lnsect pests of Rajasthan Proceedings of the 2nd All Ind~aCongress of Zoology, 1962 2 404-410 KUSHWAHA, K S , NOOR, A, and RATHORE, R R S (1980) lnsect pest management In arid agriculture Annals of Arld Zone 19 503-509 KUSHWAHA, K S , SHARMA, L S , and BHARDWAJ, S C (1964) Common Lep~dopteranfauna of Udaipur (Ra~asthan)Part ll (Moths) Un~vers~tyof Udaipijr Research Studles 2 109-127 KUSIGEMATI. K (1972) On the specles of Melohoris Holmgren from Japan (Hymenoptera, Ichneumonldae) Memoirs of the Faculty of Agricultore, Kagoshlrna Untverslty 8 1 15-1 18 LIN (Chanq Shanl, SUN (Ch~nIU), CHEN (RUILo), and CHANG (J Tsun plng) (1963) Studies on the regular~tyof the outbreak of the or~entalarmyworm, Leucanla separatd Walker I The early sprlng orlentatlon of the orlental armyworm moths dnd ~tsrelation to wlnds Acta Entornolog~caSinlca 12 243-261 LIU (Fanq), and FENG (Hui) (1965) The metdbollsm of blood sugars of the armyworm Pseudaletia (Leucania) separata (Wlk ) Actd Entomologica Sin~ca14 432-440 LONGWORTH, J F , and ARCHIBALD, R D (1975) A vlrus of black beetle, Heterony~husarator(F ) (Coleoptera Scarabde~dae)New Zealand Journal of Zoology 2 233-236 LU (Bao Ilen) (1964) Pathological changes follow~ngadm~nlstration of catharldln to Leucanla (Pseudalet~a)separata (Walker) Acta Enromologica Slnlca 13 824-834 MA, S C (1979) lnsect ecology In the People's Rppubl~cof Ch~naActa Entomolog~caS~nlca 22 257-266

MACKAY, A G , and O'CONNOR, B P (1976) Army caterpillar control in North Isldnd maize and barley w~th leptophos ICI New Zfaland Ltd Hamilton, New Zealand pp 194-197 MANJUNATH, D , and MATHAD, S B (1978) Temperature tolerance, thermal Inaction and ultrav~oletl~ght resistance of nuclear polyhedros~svlrus of the armyworm, Mythlmna separata (Walk ) (Lepid Noctu~dae) Zeltschrl f te fur Angewandte Entornologie 87 82-90

MANJUNATH, D , and MATHAD, S B (1979) lnfect~vltystudles on sunlight exposed nuclear polyhedros~svlrus of armyworm Myth~rnna(Pseudaletia) separata Compdrative Phys~ologyand Ecology 4 15-16 MATHUR, Y K , and SAXENA, R C (1972) Note on the posslble use of trlphenyl tinacetate as a crop protectant agalnst some lepldopterous pests Indian Journal of Agr~culturalSciences 42 427-428 MATSUMOTO, S , SUZUKI, A, YAMAOTO, H , OGURA, N , and IKEMOTO, H (1979) Cycllc nucleotldes and hormonal control of cuticular melanlzat~onIn the armyworm larvae, Leucanla separata (Lep~doptera Noctu~dae) Applled Entomology and Zoology 14 159-1 63 MIYAHARA, Y (1978) Effect of the Intake of sugar at adult stage on the matlng of Leucania separata and Agrot~s yipsilon Proceedings of the Assoc~ationof Plant Protection, Kyushi~24 101-1 04 MOHYUDDIN, A I , and SHAH, S (1977) 81olog1calcontrol of Mythimna separata (Lep Noctuidae) In New Zealand and its bearlng on biological control strategy Entomophaga 22 331-333

MOKROTOVAROV, S (1965) Pests and dlseases of rice In Indonesia Zashchita ot Rastenii Vreditelei Bolernei 11 49-50

MU, C V , and SUN, S S (1980) A study on the copulation and spermatophore formation In Leucania separata (Walker) Acta Entomolog~caSin~ca 23 16-24

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NAGANO, M , TOMIO, T , and YOSHIOKA, T (1972) Forecast~ngof the armyworm, Leucanla (Mythlmna) separata Walker In grassland 2 Outbreaks In Ikl IS (Japan) In 1971 Proceedings of the Assoc~atlonfor Plant Protection, Kyushu 18 9-10

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OGURA, N. (1975b). Induction of cuticular melan~zationIn the larvae of armyworm, Leucania separata Walker (Lepldoptera: Noctu~dae),by lmplantat~onof ganglla of the s~lkworm,Bombyx mori L. (Lepldoptera' Bombycldae) Appl~edEntomology and Zoology 10' 216-219. OGURA. N. (1976). Role of oesophageal connectives In cuticular melan~zat~onIn the larvae of armyworm, Leucanla separata Walker (Lepldoptera. Noctu~dae).Applied Entornology and Zoology 11 : 70-74.

OGURA, N., and MITSUHOSHI, J (1978). In vltro cultures of the endocrrne organs secreting melan~zatlorrand reddish colorat~onhormone In the common armyworm, Leucania separata (Lep~doptera.Noctuldae). Applied Entomology and Zoology 13: 224-277.

OGURA, N., and MITSUHOSHI, J. (1979) Melanlzat~onof Integuments cultured In v~voand In v~tro.AppliedEnto mology and Zoology 14, 118-120.

OGURA. N., and SAITO, T (1972). Hormonal funct~oncontrolllr>g plgmentatlon of the Integument In th~common armyworm larvae, Leucania separata Walker. Applied Entomology and Zoology 7: 239-242.

OGURA, N , and SAITO, T. (1973). lnductlon of embryonic dlapause In the s~lkworm,Bomhyx mori L. (Lepidop- tera. Bombyc~dae),by ~mplantat~onof the gangha of the common armyworm larvae, Leucania (Myth~mna) separata Walker (Lep~doptera.Noctuidae). Applied Entomology and Zoology 8: 46-48.

L OGURA, N., YAGI, S., and FUKAYA, M. (1971). Hormonal control of larval colorat~onIn the common armyworm, Leucania separara Walker. Applied Entomology and Zoology 6. 93-95. OKU, T , and KOBAYASHI, T (1974) Early summer outbreaks of the oriental armyworm, Mythlmna separata Walker, In the Tohoku District and posslble causative factors (Lepldoptera Noctuldae) Appl~edEntomology and ZOO~O~Y9 238-246 OKU, T , and KOBAYASHI, T (1977) The orlental armyworm outbreaks In Tohoku District, 1960 wlth speclal reference to the poss~brlrtyof mass immigration from Chlna Bullettn of the Tohoku Nat~onalAgr~cultural Exper~mentStation 55 105-1 25

OKU, T , and KOBAYASHI, T (1978) M~gratorybehavlour and Ilfe cycle of noctuld moths (Insects, Lep~doptera) with notes on recent status of mlgrant specles in Northern Japan Bullettn of the Tohoku Nat~onalAgricultural Experiment Station 58 97-209

OKU, T , and KOYAMA, J (1976) Long range migration as a posslble factor causlng the late summer outbreak of the orlentdl armyworm, Mythlmna separata Walker, in Tohoku Dlstrlct, 1969 Japanese Journal of Appl~ed Entomology and Zoology 20 184-1 90 PATEL, R K (1972) Pest complex of paddy In Madhya Pradesh Indidn Journal of Entomology 34 81 PATEL, R K (1979a) Some new ohservatlons on the hablt and behavlour of rlce earcuttlng caterplllar Oryza 16 71-72 PATEL, R K (1979b) Performance of sprayers and Influence of water volumes In lnsectlcrdal control of rice ear cutting caterpillar Internatlonal Rlce Research Newsletter 4 21 -22 PATEL, R K (1980) Note on the Incidence and marchlng behaviour of rlce earcuttlng caterplllar, Mythtmna separata (Walk ) Science and Culture 46 199-200

PATEL, R K , VERMA, R , PATITUNDA, A,and RAWAT, R R (1979) Chemlcal control of rlce earcutting caterprllar, Mythlinna separata (Walker) under fleld cond~tlons Oryza 16 57-59 PAWAR, A D (1976) Anedrallus spinidens (Fabriclus) (Asoplnae Pentatomldae Hem~ptera)as a predator of Insect pests of rlce In H~machalPradesh, lndla Rlce Entomologists Newsletter 4 24 PERSSON, B (1977) Dlstributlon of catch In relation to emergence of adults rn some noctu~dpest specles In south coastal Queensland Australian Journal of Zoology 25 95-102 POPHALI, D J , DESHMUKH, P D , KAUSHIK, U K , SRIVASTAVA, S K , PATIDAR, G L ,and GANGRADE, G A (1980) Assessment of yleld losses caused bk the earhead cuttlng caterpillar, Myth~mnaseparata WalKer Internatlonal Rtce Research Newsletter 5 8-9

PRADHAN, S (1971) lnvestigatlons on Insect pests of sorghum and millets (1965-1970) Flnal Technical Report, Drvislon of Entomology New Delhi, India Indian Agr~culturalResearch lnstltute

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PUROHIT, M L , KHATRI, A K , VERMA, S K , and THAKUR, G S (1971) Note on the chemtcal control of armyworm of rice, Pseudaletia separata (Walk ) (Noctuldae Lepldopteral lndlan Journal of Agricultural Sciences 41 69-70

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QUO (Fu) and LIU, (Chlng lung) (1964) Stud~eson the reproduct~onof the armyworm, Leucanla (Pseudaletta) separata Walker (Lepidoptera Noctuldae) I1 Effect of supplemerltary nutrltlon on fecundity Acta Entomo logica Sintca 13 785-794

QUO (Fu),WU (Tsiu ngun), TSAl (HUI leo), and LIU (Ching lung) (1963) Studles on the reproduct~onof the armyworm, Leucanla separata Walker (Lep~doptera Noctuldae) I The biological characterist~csof adults Acta Entomolog~caSinrca 13 785-794

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RANGARAJAN, A V , MAHADEVAN, N R , KANDASWAMY, G , and IYEMPERUMAL, S (1974) Some observations on the suscept~bll~tyof sorghum types and hybr~dsto the cutworm, Pseudaletia unipilncta How Sorghum Newsletter 17 48 RANGARAJAN, M , SRIVASTAVA, R P , and CHAKRAVARTI, B P (1968) A note on thc different Insect hosts of Serratla marcescens Blz~oLabdev Journal oi Zclence and Technology 6(B) 159-160

RAO, P S P , MISRA, B C , DAS, P K , and PADHI, G (1976) Effectiveness of Itoplectisnarangae Ashmead (Ichneumon~daeHymenoptera), an exotic pupal paraslte agalnst rlcp cutworms and stembalers and short notes on ~tsbehav~our Oryza 11 89-91 RAO, S N (1953) Notes on some parasltlc Hymenoptera from lnd~aw~th descr~pt~on of a new specles, Apantelrs ebilarbi Ind~anJournal of Entomology 15 25-28 RAO, S N (1969) Stud~eson Indian Ichneumonrdae (Hymenoptera, Paras~t~cd)Jocjrnalof the Bonrbdy Natr~ral History Society 66 222-226 RAO, V P , CHACKO, M J , PHALAK, V R , and DINESH RAO, H (1969) Leaf feed~ngcaterprllars of pdddy dnd the~rnatural enemles In lndla Journal of the Bombay Ndtural History Society 66 455-477

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SAKAI, M , SATO, Y , and KATO, M (1967) lnsect~c~dalactlvlty of 1,3 bis (carbdmoylthio) 2 (N,N d~methyl amino) propane hydrochlor~de,cartap, w~thspecial references to the cffect~ven~ssfor controll~ngrice stemborpr Japanese Journal of Applied Entomology and Zoology 11 125-1 34 SARUP, P , PESHWANI, K M , SINGH, D S , SRIVASTAVA, V S , and RATTEN LAL (1969) An unusual out break of armyworm, Pseudaletid separata (Walker) at Delhi in 1968 and evaluat~onof the prstic~desagalnst thr larvae ~nfest~ngwheat Indian Journal of Entomology 31 265-272

SATO, Y (1965) The artlfic~ald~et for mass rearlng of the tobacco cutworm, Prodenia litura Fahrir~usand the common armyworm, Leucania separata Walker Japanese Journal of Applied Enton~o/ogydnd Zoology 9 99- 106 SATO, Y , TAKAHASHI, S , SAKAI, M , and KODAMA, T (1980) Attractiveness of synthet~csrx pheromone to the males of the armyworm, Leucania sepdrata Walker and lorey leafworm, Leucania lorey~Dupor~ch~l (Lepidop tera Noctu~dae)Applied Entomology and Zoology 15 334-340 SAXENA, A P (1965) Blonomlcs and control of armyworm on suqarcarle Indian Sugarcane 15 143-149

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