1111 ESEARCH PAPER, SERIES NUMBER 123 JANUARY 1987

UPLANDRICE. INSECTl--1i M gm T -E!R.EOLOU t RTANCE ,AND" ONTzROL

J.A. LITSINGER, DANDI SOEKARIAAl. BARRION, and

T~einternation' Rice Research Instiute P.O.".16 93, Manila, Philippines UPLAND RICE PESTS: THEIR ECOLOGY, IMPORTANCE, AND CONTROL

J.A. Litsinger'. A.T. Barrion2. 3 and Dandi Soekarna

ABSTRACI

The diversity of upland rice environments gives rise to a more heterogeneous insect fauna compared with the more homogeneous lowlands. A wide array of soil-inhabiting pests - ants, termites, white grubs, crickets, root aphids, root mealybugs, root bugs, and wireworms -- common in upland rice cannot tolerate flooding. Seedling maggots replace the aquatic whorl maggots as vegetativc foliar pests. lnsect-vectored virus diseases arc rare in upland rice. Small upland ricefields cause concentrations of the more vagile seed pests during ripening. The less stable upland environment -- more restricted growing season, smaller area planted, greater drought stress - poses greater problems of survival to , which have overcome them by polyphagy, greater longevity, off-season dormancy, and/or dispersal. There is no one insect that specializes in upland rice. Yield losses to insects, however, are comparable to those of lowland rice. Cultural conrol methods include increased tillage, higher seeding rates, and crop rotation for soil pests. Foliar pests can be minimized by synchronous planting of early­ maturing varieties. Plant resistance as a method of insect control has iot been greatly exploited because the other breeding objectives of high yield, drought tolerance, and blast resistance take priority. Resistance to stem borers should be a high priority. A rich fauna of natural enemies exists, but they face even greater problems of survival than the pests. The normal low yields of upland rice preclude high levels of insecticide use. Seed treatment and baiting are low-cost mnethods, as is low-volume sprayi 1g.

Entomologist, Entomology Department, International Rice Research Institute (IRRI), P.O. Bo- 933, Manila, Philippines. 2 3 Scnior research assistant, Entomology Department, IRRI. Senior researcher, Pest and Disease Section, Bogor Institute for Food Crops (BORIF), Jalan Cimanggu, Kecil 2, Bogor, Indonesia. UPLAND RICE INSECT PESTS: THEIR ECOLOGY, IMPORTANCE, AND CONTROL

Upland ricefields are nonpuddled and unbunded, without Although a dryland habitat represents an extreme the expectation of impounding water. They may be located hydrological condition, upland rice is host to all but the in many settings - in isolated pockets surrounded by most aquatic insects (11, 176). Whorl maggots, case­ irrigated wetland fields, or along steep slopes of recently worms, water weevils, and bloodworms require ponding. cleared forest. Each enviionment produces a unique Many soil and scedling pests are not common in lowland composition of insect . ricefields. Deep water rice is established in dry soil and, Information on upland rice insect pests is limited and therefore, has more in common with upland and rainfed scattered among reports dealing with other rice cultures. We lowland rice than with irrigated rice, even though water found eight reports specifically treating insect problems of depths may later reach 1-3 in.Second to deep water rice, upland rice culture (11, 59, 66, 79, 116, 120, 163, 176) but upland rice represents the most unstable rice environment none worldwide in scope. Most literature focuses on one for foliar insects. But upland rice is highly stable for soil species or a group of related species. Table I lists upland rice insects. insect pests that others have identified and those we have The significance of abiotic factors in upland rice insect observed. Many citations refer to lists of pests with rice as a ecology will be apparent in a discussion of the most recorded host, or are based on insect habits. prevalent groups of insects attacking upland rice Papers concerning upland rice often fail to distinguish worldwide. between the two ecologically significant aspects ---culture and environment embodied in th': term upland rice. It Soil -inhabiting pests is dangerous to extrapolate results of studies of insect Well-drained, nonpuddled upland rice soils favor pests pests in well-drained fields where seeds of upland rice that pass at least one growth stage underground. Soil .varieties are sown into dry soil (upland rice culture) but in pests feed on underground plant parts (sown seed or an otherwise lowland environment (fields surrounded by roots), develop entirely in a subterranean habitat, and flooded rice). Insects readily fly from the lowland rice- leave only for adult mating and dispersal flights. Soil fields to the fields planted in upland rice culture. These pests include ants (Formicidae), termites (Isoptera), mole insects, therefore, are not necessarily upland rice insects, crickets (Gryllotalpidae), field crickets (), white They breed on lowland rice and colonize upland rice. We grubs and black (Scarabacidac), root aphids gain greater ecological significance from faunistic records (), root-feeding mealybugs (Pseudococcidae), and ecological studies from an upland rice environment, root-feeding bugs (Lygacidac, Cydnidae), false wire­ where lowland ricefields ar- out of the effective dispersal worms (Tenebrionidac), wireworms (Elateridae), root range of most insect species (tens of kilometers away). In weevils (), and soil-inhabiting cutworms border areas where lowlands and uplands meet, studies (Noctuidac). cannot accurately represent either upland or lowland rice A subterranean environment limits mobility, particularly (181). in locating food, and soil insects have adapted by 1)being long-lived either as individuals (beetles or orthopterans), as IPEST (iROUPS colonies olsocial insects (ants or termites), or as dependents on social insects (mealybugs and aphids), and 2) having a Upland rice insects are more influenced by physical than wide host range (all species). by biological or socioeconomic parameters: I) well- Anis. Species of -rrestrial ants, notably the ubiquitous drained soils (lack of prolonged flooding or soil puddling), fire ant Solenopsis geminata and harvester ants Pheidole 2) high probability of drought during crop growth, 3) spp., specialize in feeding on ungerminated grass seed. restricted growing season (lengthy nonrice fallow), 4) Solenopsis readily colonizes disturbed habitats, which in ricefields interspersed with other crops (diversified flora), turn initially encourage the growth of grasses (39). Colonies and 5) low use of agrochemicals (because of low and of these granivorous ants have a specialized caste that unstable yield), processes seed for food. Foraging workers bring seed to 4 IRPS No. 123, January 198/

Table 1.Insect pests of upland rice worldwide. Order Species Distribution Reference Isoptera Rhinotermitidae Coptotermesformosanus Shiraki Asia 85 Heterotermesindicola (Wasmann) Asia Termitidae 82 Anacanthotermesviarum (Koenig) Asia 190 Anacanthotermesrugifrons Mathul et Sen-Sarma Asia 190 Captiterpnesnitobei (Shiraki) Asia 82, 85 Corniterrnesstriatus (lHagen) Latin America 151 Procapriterines mnushae Oshima Asia 82, 85 Pericapritermes nigerianus Silvestri (=socialis) Africa 7 Macroternes nutalensis (Haviland) Africa 85 Macrotermes beiiicosus Smcat hman Africa 85 Macroterines gilius (Hagen) Asia 178 Macrotermnes spp. Africa 12 Odontoterniesformosanus (Shiraki) Asia 82, 85 Procornitermesarauloi Emerson Latin America 85, 195 Procornitermws triacifer (Siivestri) Latin America 195 Trinerviterniesgenzinatus Wasmann Africa 85 [= ebenerianus Sjostedt] Microceroterynesspp. Asia 193 Microterinesspp. Asia, Africa 11, 12, 157 Synternes nolestits (Burmeister) Latin America 60, 85, 195 Termites unspecified Dermaptera Forficulidae Asia 72 DiaperasticuserythrocephalusOlivier Africa 36 Don lineare (Eschscholtz) Gryllotalpidae Latin America 186 GryllotalpaafricanaPalisot de Baeuvois Africa 12, 32 (=africanu) Burmeistera Asia 59, 72, 76, 84 Neocurtilla (= Gryllotalpa)hexadacryla(Perty) Latin America 81, 150, 195, 203, 211 Scapteriscusdidact),hts(Latreille) Latin America 81 Gryllidae Grjllusassinilis(= bimaculatus)(Fabricius) Asia 76 Gryllus (=Liogryllus) bimaculatusde Geer Asia 178 Brachytrupes portentosus (Lichtenstein) Asia b Brachytnmpes mnembranaceus Drury Africa 48 Plebeiogrylhsplebejus (Saussure) Asia b Teleogryllus testaceus(Walker) Asia 76, 178 Teleogryllus occipitalis (Serville) Asia b Loxoblenimus haani Saussure Asia b Velarifictorusaspersus(Walker) Asia b Velarifictonis sp. Asia b Euscyrtus concinnus (de Haan) Asia Pyrgomorphidae b Atractomorpha burri I. Bolivar Asia 55 A tractomorphapsittacinapsittacina(de Haan) Asia Tettigoniidae 178 Conocephahslongipennis(de Haan) Asia 178 Conocephalusrnaculatus(Lethierry) Asia 178 Conocep.tahissaltator Saussure Latin America 65 Euconocephah s varius (Walker) Asia 178 Caulopsiscuspidatus (Scudder) Latin America 68, 81 Caulopsis oberthuriScudder Latin America 68 Phaneroptera furcifera (Stal) Asia 178 Tetrigidae Amphinofus spp. Asia Acrididae 178 Aiolopus thalassinustamulus (Fabricius) Asia 178 Acrida ivilleinsei Dirsch Asia 178 Gesomida mundata zonocera Navas Asia 178 Gonistabicolor (de Haan) Asia 55 Oxyajaponicajaponica (Thunberg) Asia 72, 178 Oxya hya intricata (Stal) Asia 178 Oxyafitscovittata (Marscliall) Asia 78 Oxya velox (Fabricius) Asia 178 Gas frimargus niarmoratusgrandis (Saussure) Asia 178 Xenocatantopshunilishumilis Serville Asia 55 Hierogiyphus banian (Fabricius) Asia 55, 72 ltieroglyphusdaganensis Krauss Africa 6 Hicroglyphus nigrorepletus(1. Bolivar) Asia 82 Hieroglyphus oilzii'orosCarl Asia 82 Heteropternisrespondens(Walker) Asia 55 Patanga succincta (Linnaeus) Asia 54, 55, 59, 130 Stenocatantops splendens (Thunberg) Asia 55, 178 Chondracrisrosea brunneri Uvarov Asia 55 Trigonidium cicindeloides Rambur Asia 78 Locusta migratoriamanilensis (Meyen) Asia 55 Locusta migratoriacapito Saussure Africa 82 IRPS No. 123, January 1987 5

Table I continued.

Order Family Species Distribution Reference Locusta migratoria migratorioidesReische Africa 33 et Fairmaire Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82, 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68 lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186, 246 Pseudococcidae "'taennococcus spp. Asia 120 Lhorizococcus i/u Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia, Africa, 245 Latin America Dysmicocc:us brevipes (Cockcrell)c Asia, Africa, 117, 203,245 Latin America Dysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245 Brevennia (= tererococcus)rehi (Lindinger) Asia 72, 245 Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp. Latin America 60 Saccharicoccus sacchari (Cockerell) Africa, Asia, 245 Latin America Trionymus ceres Williamsc Asia 119,245 j rionymus sp. Asia b Mealybug unspecified Asia 83, 84 Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji) Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia, Africa 12, 97, 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia, Africa, 66, 76, 120, 195,203, Latin America 221,229,240, 249 Tetraneura akinire Sasaki Asia 221,249 Tetraneurabasui ltillcRisLambers Asia 249 Terraneura nigriabdominalis (Sasaki) Asia, Africa 7,12, 30, 76, 227,229 (=Dryopeia hirsuta) Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249 [= Forda harukawai Tanaka] Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America, 76, 195, 229, 249 Asia Geoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221,229, 249 Hysteroneura sctariae (Thomas) Africa, Asia 7 Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72, 120 Cercopidae Deois schach (Fabricius) Latin America 61, 68, 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61, 68, 194 Tomaspis ( Dcois) flai'opicta (Stal) Latin America 61, 61, 194 To naspis ( Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68, 194 Tomaspis ( Aeneohn.'a) spectabilis (Distant) Latin America 219, 233 Aeneolamia varia Fciinai Latin America 81, 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54 Delphacidae Sogatella furcifera (Horvath) Asia 59, 241 Ni'apanatalugens (Stal) Asia 30, 59, 72, 84, 241 Sogatodes oryzicola (Muir) Latin America 61,67,150 Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia b Peregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18 Cicadellidae Recilia dorsalis (Motschulsky) Asia 59, 241 Nephotettiv nigropictus (Stal) Asia 59,241 6 IRPS No. 123, January 1987

Table I continued, Order Family Species Distribution Reference Nephotettix virescens (Distant) Asia 59, 241 Nephotettix modulatus Melichar Africa 217 Cofana spectra(Distant) Asia 59, 241 Graphocephala spp. Latin America 61,66 Hortensia simnilis (Walker) Latin America 61, 66, 81, 150 Exitianusobscurinervis (Stal) Latin America 61,66 Balclutha spp. Latin America 66 Draeculacephalaclypeata Osborn Latin America 81,150 Cicadulinabipunctella(Matsumura) Lygaeidae Asia b Cymoninus turaensis(Paiva) Asia 178 Ninus insigni"Stal Asia 178 Pachybrachiusnenosus Horvath Asia 30 Caenoblissuspilosus Barber Asia Blissusleucoptens 83 (Say) Latin America 42, 81,150, 203 Blissus spp. Asia 30 Dlmnorphoptentscornutus novaeguineae Ghauri Asia 80 Dimorphopterus similis Slater Africa 62 Paroniuspiratoides (Costa) Cydnidae Asia 178 Pangaeus spp. Latin America 195 Aethus indicus (Westwood) Asia 178 Geotomnus pygmaeus (Dallas) Asia 178 Stibaropus inolginus Schiodte Asia 178 C'yrornenusbergi Froeschner Latin America 203 Cyrtomnenus ciliatus (Palisot de Beauvois) Latin America 203 Cyrtonenuscrassus Walker Latin America 203 Tominetus spp. Latin America 203 Alydldae Riptortuslinearis (Fabricius) Asia 55, 178 Riptorts spp. Africa 12 Leptocorisa acuta (Thunberg) Asia 72, 78, 120, 214 Leptocorisabiguttata (Walker) Asia 214 LeptocorisapalawanensisAhmad Asia b Leptocorisaoratorius(Fabricius) Asia 59, 120,178 Leptocorisachinensis Dallas Asia 114 LeprocorisasolomonensisAhmad Asia 178 Leptocorisaspp. Asia 54 Stenocoris southiwoodi Ahmrad Africa 6,36, 82 Stenocoris claviformis Ahmad Africa Mirperus 12 torridusWestwood Africa 6, 12, 82 Coreidae aetus trigonus (Thunberg) Asia Corimelaenidae Alkindus 178 atratusDistant Latin America 81, 150 ScaptocorisdivergensFroescher Latin America 60 Scaptocoris castaneusPerty Pentatomidae Latin America is, 195 Scotinopharatarsalis (Vollenhoven) Asia 59 Scotinopharascotti Horvath Asia 120 Eysarcoris (=Stollia) ventralis (Westwood) Asia 178 Dolycorisindicus Stal Asia 78 Macrinajuvenca Burmeister Africa 36 Tanda antiguensis (Westwood) Latin America 186 Tantiageifi Schout Africa 36 Tantia perditor(Fabricius) Latin America 186 Pygotnenidavaripennis (Westwood) Asia b Menida spp. Nezara viridula Asia 55 (Linnaeus) Asia, Africa 12, 30, 54, 55, 78, Oebalus 82, 120 poecilus (Dallas) Latin America 61, 66, 81,195 Oebalusypsilon-gnseus (de Geer) Latin America Oebalus 61, 65, 66, 195, 211 grisescens (Sailer) Latin America 150, 195 Tibracea limbativentris (Stal) Latin America 61, 66, 81, 150, 219, Aspavia 233 armigera(Fabricius) Africa 6, 12, 82 Diplo.'ysfissa Erichson Africa 6, 82 Acrosternum marginatum(Palisot de Beauvois) Latin America 186 Acrosternum spp. Thysanoptera Thripidae Latin America 186 Frankliniellarodeos? Pergande Latin America 61, 66 Breginatothrips 'enustos Watson Latin America 68 Stenchaetothrips(=Baliothrips)biformis Asia 213,235 (Bagnall) (=(Jlloethripsoryzae (Williams) HaplothripspriesnerianusBagnall Asia 178 IRPS No. 123, January 1987 7

Table I continued.

Order Family Species Distribution Reference t.oleoptera Scarabacidae !leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusme!es robustui, Prell) Alissonotum pauper Burnieis'er Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28, 195, 233 Devscinetusgagates Burmnc:ster Latin America 20. 195 Dyscinctt" spp. Latin America 60 Eutheola Inimilis Bturmcist.:r Latin America 20, 28, 8 1, 195, 203, 233 Eutheola ',identarr(Biurnci,ter) Latin America 121, 203 Heteron)ciwv aw 'rson"Jack Africa 82 Hcreron),chusbtu.,terculatus Kolbe Africa 82 tleteroni'chtslicas (Klug) Africa 82 fh'teron),chus mosamhicus Peringuey Africa 4, 6, 48, 82 (= Veteronychus or',zae Britton) leterony'chts lioderes Redt,.nbacher Asia 77, 82, 207 ltcteron',chusarator (Fabricius) Africa 82 tleteront'chusplebepus (Klug) A frica 57, 82 Ileteron 'chus pseu..'ocongonnsis Ferreire Africa 32 Ileteronvchns rogiJ'rons Fairmaire Africa 82 Heteron 'chus nrsticus niger (K lug) Africa 82 Heteronvchus spp. Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp. Asia 120 Ligvros fossator B urmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20, 195 Dynastinac Ligtvros huonilis Burmeister Latin America 60, 82 A/aladera castanea (Arrow) Asia 82 Maladera orientalis .Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phv/lognathusdicynsius (Fabricius) Asia 82 Melolonthinae E.xopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna [= lolotrichia] consanguinea Asia 122 (Blanchard) Leucopholisrorida (Fabricius) Asia 82, 117 Lepidiota blanchardi Dalla Torre Asia 76, 82 Leucopholis irrorata (Chevrolat) Asia 76, 134 Scrica internpta Walker Asia 76 Holotrichialon.ipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77, 78 Holotrichia pnindanaoana Brenske Asia b Stenoer,,es spp. Latin America 71, 195 Atoenius spp. Latin America, Asia 203 Ph. lupl:.!g. aequata(Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas),poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pam'isetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Ph'llophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82, 117, 159 Phvllophagaspp. Latin America 81, 150 White grub unspecified Asia 84 Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var. barhata Burmeister Asia 77, 78, 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala pal/ida (Fabricius) Asia 82 8 IRPS No. 123, January 1987

Table 1 continued. Order Family Species Distribution Reference Anoma'a volita (Blanchard) Asia 82 Adoretus caliginosus Burmeister Asia 82 Adoretus comprcssus (Weber) Asia 82 Papuana huhneri (Fairmaire) Asia 82 Papuana incrmnis Prell Asia Popillia 82 capricollis Hlope Asia 78 Tenebrionidae Gonocephalu: depressu;r (Fabricius) Asia 82, 117, 178 Gonocephahm acuiangulum (Fairmaire) Asia Gonocephalum 82, 117 sinph'x (Fabricius) Africa 82 Epitragus sallei Champion Latin America 203 Anaedus ptnctatissimnts Champion Latin America Elateridae Aeolus spp. 203 ,4griotes Latin America 195 spp. Latin America 195 Agriotes mancus (Say) Latin America 203 Conoderts spp. Latin America 195 Aeoloderma brachmana (Candeze) Asia h Wireworni unspecified Languriidae Asia, Latin America 60, 73, 117 Anadastus filijf rmis (Fabricius) Asia Chrysomelidae Dicladispa 200 viridic',anca(Kraat z) Africa 36 Trichispasericea (Guerin) Africa Oediopalpastervalis (Weise) II Latin America 61,66, 68 Oediopalpa guerini (Italy) Latin America Chaetocnema 61,66, 68 cvlindrica(Baly) Asia 225 Chactocnema denticulata S tephens Latin America 61, 66, 68, 81 Chaetocnemabasalis (Baly) Asia 178 Aulacophora sp. nr. similis Olivier Asia Monolepta 55 car'ipemne Ialy Asia 55 Monolepta bilasciata (Ilornstedt) Asia h Chrysomelidae Monolepta signata Olivier Asia 55 Cerotoma airojcsciata J acoby Latin America 203 Diabrotica adelpha Ilarold Latin Amefi,:a 203 Diabrotica balteata Leconte Latin America 203 Diabrotica graminLa ltaly Latin America 203 Diabrotica speciosa (Germar) Latin America 61,66, 68 Diabrotica limitafa quindecinmunctata(Germar) Latin Amcrie: 195 Diabrotica melanocephala (Fabricius) Latin Amw , 195 Diabrotica spp. Latin Aierwca Altica (= 61, 81 Fondia)madagascariensis(Allard) Africa 57 Altica spp. Asia Epitrix cuctmeris 178 (Harris) Latin Americ 203 Epitrix spp. Latin America 81 Oulenjo or 'pae (Kuwayama) Asia 82 Disonchq (= Donacia) spp. Latin America 195 Flea unspecified Asia 120 Coccinellidae Chnootriba (= h'tIilachna) sinilis assimilis Mulsant Africa 6, 11 Curculionidae Atactogaster iduriens Walker Asia 185 Hypomeces squtamosus (Fabricius) Asia 120 Tan vinecus discoidalis Gyllenhal Africa 82 Phaulosoinusmusculinus (= mnus) Csiki Africa 82 Neobaridia anplitarsis Casey Latin America Lymantriidae Psalis pennatula 66, 68, 195, 233 (Fabricius) Asia 117 Laelia sufihsa (Walker) Asia 117 Euproctis 'irgtncula (Walker) Asia 117 Euproctis minor (Snellen) Asia Euproctis 117 xanthorrhoea (Kollar) Asia 117 Amatidae Amata sp. Asia A. S. Pradhan, Arctiidae Diacrisia obliqua unpubl. data (Walker) Asia 82 Creatonotusgangis Linnacus Asia Limacodidae 82 Latoia (= Parasa) bicolor (Walker) Asia Eupterotidac Nisaga simplex 82 Walker Asia 182 Noctuidac Sesamia botanephaga Tams & Bowden Africa 12 Sesamnia nonagroides botanephagaTfums et Bowden Africa Sesamia calamistis 6, 36 llampson Africa 6, 8, 11, Sesamia inferens 12, 36 (Walker) Asia 59, 82, 84,240 Spodoptera eridania (Cramer) Latin America Spodoptera mauritia acron),ctoides 150 Guenee Asia -0, 59, 196 Spodoptera frugiperda (. E. Smith) Latin America 28, 61, 66, 68, 81, 150, 152,195,203,219,233 IRPS No. 123, January 1987 9

Table I continued.

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Africa 32, 217 Spodoptera exem pta (Walker) Africa 32, 33, 36,217 Spodoptera ornithogalli (Guenee) Latin America 150 Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Asia 178 Mythimna (- Pseudaletia)latifascia[= adultera Latin America 195 (Schaus)] (Walker) Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30, 117 Mocis latipes (Guenee) Latin America 66, 68, 149, 150, 203, 219,233 Platysenta (=Spodoptera) compta (Walker) Asia A. Barrion, unpubl. data Agrotis ipsilon (Hufnagel) Latin America, 33, 68, 81, 150, 1)5, Africa 203 Agrotis spp. Latin America 60 Achaeajanata (Linnaeus) Asia 117 EldanasaccharinaWalker Africa 36 Armyworm unspecified Asia 72, 84 Pyralidae Cnaphalocrocis medinalis (Guenee) Asia 30, 45,59, 72, 84,233 Marasmia bilinealisHampson Africa 36 Marasmia trapezalis(Guenee) Africa 32, 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Asia b Marasiniaspp. Asia b Leaf roller unspecified Asia 72 M2liarphaseparatellaRagont Africa 12, 57 Maliarphasp. Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Africa 12, 92 Chilo polychrysus (Meyrick) Asia 59, 72 Qhilo diffusilineus J. de Joannis Africa 6, 12,29 Chilosuppressalis(Walker) Asia 59, 72 Cibozacconius Blezzinski Africa 6, 11, 12 Cravubus spp. Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar) Asia 82 Scirpophagaincertulas(Walker) Asia 59, 72 Scirpophagainnotata (Walker) Asia 59, 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28, 60, 61, 66, 81, 150, 162,195,203,219,233 Diatraeasaccharalis(Fabricius) Latin America 28, 66. 81, 111, 150, 195 Rupela albincllaCramer Latin America 111 Stem borer unspecified Asia 72 Undetermined stem borer nr. Maliarpha Asia b Gelechlidae BrachniaarctraeaMayr Asia 55 Brachmi. spp. Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Asia 164 Pelopidasmathias (Fabricius) Asia, Africa 36, 155,217 Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6 Satyridac Melanitisleda ismene Cramer Asia 84,213 Mycalesis asophisHew Asia 30 Mycalesis spp. Asia b Diptera Cecldomyiidae Orseoliaoryzae (Wood-Masn.) Asia 72, 88, 131, 181 Orseoliaoryzivora Harris et Gagne Africa 6,11,32,218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson) Asia 205 Oscinellaspp. Asia 205 Gaurax spp. Asia 205 Chloropsoryzae Matsuntura (= kuwonae Aldrich) Asia 82 Mucldae Atherigonaoryzae Malloch Asia 30, 82, 117,130 Atherigona exigua Stein Asia 82, 117, 178 10 IRPS No. 123, January 1987

Table I continued. Order Family Species Distribution Reference Atherigona oritentalis Schiner Asia 82 Atherigonaindica Malloch Diopsidae Diopsis longicornis(= Asia, Africa 82,205 thoracica)Macqurt Africa 11, 12,48 Diopsis spp. Agronyzidae Africa 217 Pseudonap.nyza asiatica Spencer Asia Pseudonapomyza 213 spicata (Malloch) Asia h Agrono'za orvzae Munakata lHymenoptera Formicidae Acromyren" landoltibalzani Asia b (Emery) Latin America 61,68, 195 Acromyrmex landolti fracticornis(Forel) Latin America 61, 195 Acromyrnex heyeri 'Forel) Latin America 61, 68, Atta bisphaericaFore! 195 Latin America 61, 68, 195 Atta capiguaraGoncalves Latin America Atta laeLigata(F. Smi 14, 61, 68, 195 h1) Latin America 61, 68, 195 Atta opacicepsBorgmeier Latin America Solenopsisgeminata 61, 195 ('abricius) Asia b Pheidole sp. Asia Pheidolegeton b diversus Jerdon Asia 1) Monomorium pharaonis (Linnaeus) Asia b Ants unspecified Asia 120 aG. orlentalls formerly recorded in Asia as G. africana. thPhilippines, reported in this study. CSubterranean root feeders.

subterranean nests. Foraging ants will feed on germinated young rice plants (47, 68). They take leaves to fungal gardens seeds only when the supply of ungerminated seed is low. in underground nests where fungi predigest the However, plant these ant specialists feed on a wide variety of material. plants and thus help control grassy weeds (39). This dual Termites. Subterranean termites of the family Termitidae, pest/ benefit role extends from tending ants, which protect which lack symbiotic protozoa to help digest plants, are the honeydew-excreting pest species including root aphids and most frequently mentioned termite pests of upland rice (11, mealybugs (workers repel potential predators and even 85, 120). This family cultures fungi in special underground selectively kill parasitized aphids), to predators of leaf- and cells --- fungal combs made of half-digested plant material stem-feeding rice pests such as leaffolders, armyworms, (117). The fungi, inoculated onto the combs, break white grubs, and stem borers (1 17). down plant material into food that termits can digest. The The upland rice environment is particularly well suited to termites then consume the fungal combs. These grassland ants because of the lack of flooding. Frequent tillage termites build nests below the plow layer in upland perpetuates grasses and granivorous species that are highly ricefields. The colonies attack living plants only when dead adapted to constructing new nests in recently tilled fields, plant material is not available. They will attack a drought­ Ant nests, however, are relatively shallow, and tilling with a stressed crop but prefer older plants having greater cellulose moldboard plow can destroy them. Also, upland ricefields content. They will damage newly planted crops where clean are u';ually small and isolated, and a given area in a small culture has removed vegetation. field has more field borders than it has in a larger field, Infestations are worse in deep, light-textured soils with allowing high rates of infiltration from more sedentary low moisture content. The first sign of damage is yellowing species such as Plieidole that do not colonize tilled fields but of older leaves (195). Termites feed on roots (the plant nest in soil under shrubs and trees along field borders, yellows, then wilts, and finally collapses) and germinating Pheidole workers forage into fields from borders and store seed (loss of stand), or move above ground at night to cut seed in their nests. In slash-and-burn or other no-tillage rice seedlings at ground level, which they cover with soil for later cultures, nests would be preserved in the field and possibly consumption. Soft-bodied termites are highly sensitive to lead to high rates of seed . However, granivorous desiccation. They live within a self-contained system of species do not prefer forest habitats, airtight chambers and build surface tunnels linei with Damage is characterized by reduced, usually patchy plant nud and body secretions (carton) to maintain more than 90% stand (120). Seed-thieving ants greatly prefer rice, but they relative humidity. Termites can be located by their tunnels. can be selective regarding plants they collect and store. They Termites apparently can withstand limited submergence harvest more seed than they can consume, so patches of (157) and can be a pest of dry lowland seedbeds (117), germip ing surplus seed (such as Rottboellia) will sprout perhaps by having nests below perched water from their nests during the rainy season. tables. Termites appear to do more damage in Latin America In Latin America, leaf-cutting ants (Aita spp. and where large ricefields are plowed with tractors depriving Acromyrmex spp.) occur only in upland fields and defoliate termites of preferred vegetation and forcing them to live on IRPS No. 123, January 1987 11

rice plants. In Asia, upland ricefields are typically small with pupae and adults and eventually dig their way to the soil Iprge field border areas (ecotones) where termites can forage surface and fly to nearby trees to seek food and mates. The for preferred food other than growing rice plants. All adults of most white grub species feed only on trees, but in grassland termites do not feed on rice. In an upland area in some species, notably Eutheolahumilis, the adults defoliate Batangas Province, Philippines, ricefields are commonly and cut off stems at ground level (28). Putheola hidentata infested by Hosiit/iternesluzonensis (Oshima); however, damages sugarcane fields in Venezuela (121). Floods kill this termite does not damage rice. In the same province, white grubs (208). Grasslands can support large populations though, Mrroternesgi/vus(Hagen)feeds on rice seedlings and, therefore, white grubs can be more abundant in newly (178). planted upland ricefields that were previously fallow (81). In Africa, termites may damage rice even though fields Black beetles live up to a year; adults can attack a crop at are small and surrounded by perennial vegetation. These any stage by burrowing into the soil (127) and feeding at the termite species perhaps prefer rice over other hosts, base of stems to cause whiteheads (207). Adults emerge with Mole crickets. Nymphs and adults are nocturnal and the early rains and are normally more abundant and do burrow through soil!, feeding on roots of a wide variety of more damage on a young rice crop with small root systems plants, or forage above ground as predators of insects. (38, 81). Adults live 3-5 mo and are even cannibalistic. Mole crickets The larvae feed only on dead organic matter in dryland prefer low-lyin!,, moist upland soils with high organic fields and do not attack rice. Adults arc highly mobile and, matter (II, 81). In the lowlands they inhabit rice bunds or although sensitive to flooding, can invade a field soon after nonflooded fields. Adults are highly mobile and can leave a it drains (115). flooded field to locate a more suitable habitat. Losses show Root aplhids. Several genera of migratory aphids feed on as wilted plants (from root feeding) or reduced stands (120). the roots of upland rice (249). Populations build up more in Damage is more common near field borders, where mole light-textured soils with high percolation rates. Damage has crickets relocate after tillage (126). also been reported from dry seed heds of lowland rice (52). Fiehl crickets. Nymphs and adults of field crickets have Tending ants -- Pheidh/le, Cren ;'aster, Tetramoriuw, similar nocturnal habits and damage rice as much as mole lsius, Tapinoma -- are necessary for root aphids to crickets do. Piles of weeds removed from fields attract them. multiply. Aiits harbor aphids in their nests over winter or They make subterranean nests and tunnel through the soil during unfavorable periods and relocate them on rice plants, to feed on roots. Some species prefer seed to roots. Others digging tunnels along the root systems to allow these soft­ feed at the base of stems, causing deadhearts. bodied insects to penetrate the soil. Most intensive studies Tunnel entrances surrounded by excavated soil are easily on aphid life histories have been done in temperate seen. Young plants are cut at ground level and stored in environments (158, 222, 229). Root aphids fly to rice plants underground cell". Field crickets tre highly polyphagous, as at the beginning of the rice season and pass through several are all soil pests (117). Sachan et al (198) reported that field generations, continually developing winged forms to crickets prefer maize to upland rice. Like mole crickets, field relocate on alternate grassy hosts until winter, when they crickets cannot survive in standing water and are, therefore, move to perennial hosts, usually trees. Yield loss occurs more prevalent in upland fields, mainly through reduced tillering, but infested plants become Whitegrubsandh/ackbeetles. Scarab beetles that feed on yellow and stunted, and, in extreme cases, wilt (227). Aphid living roots as larvae but not as adults are called white grubs populations build up gradually, so damage usually begins in or chafers (Melolonthinae, Rutellinae), and those that feed the late vegetative and reproductive growth stages. as adults but not as larvae on plant crowns are called black Root-feeding ineal tugs. More than half of the mealybug beetles (Dynastinac). In the tropics, chafers or white grubs species or. rice feed on roots (245). Being soft bodied, have a I-yr life cycle starting with adults emerging from the mealybugs are not adapted to living underground, but they soil about I mo after the first downpour of the rainy season survive because tending ants (the same species that tend root (134, 159). They lay eggs at the same time farmers sow aphids) dig burrows for them and move them from plant to upland rice. The rice passes its most susceptible stage, and plant. Well-drained soils ihelp mcalybugs survive. Although damage is mostly avoided when the white grubs are small. six species are recorded as subterranean root pests of rice, After several months, the long-lived larvae are large enough we found no report citing damage. Root-feeding mealybugs for two or three to denude the root system of mature rice are probably more prevalent on perennial grasses than on (223). This intensity of damage is rare, but wilting occurs short-lived annual rice. when root loss is combined with drought stress. The most common rice mealyb ig, Brevennia rehi, is a The larvae need damp (not saturated or dry) soil to foliar feeder favored by lack ol rain,; apparently it is survive and pass the unfavorable dry season 0.5-2 m responsible for high yield loss in Northeast India and underground in aestivation in the moist soil. The first heavy Bangladesh (N. Panda, pers. comm., as stated by IRRI rains of the season (20-30 am/d) stimulate the grubs to [1061). This report needs confirmation. Mealybug damage resume activity, and after several weeks they develop into was only briefly mentioned in another report (156). 12 IRPS No. 123, January 1987

Root weevils. A few species of upland weevils as it decays. Plants die under heavy attack, whereas lighter (IH'porneces, Donacia, Atactogaster) attack rice roots as infestations cause stunting, delayed maturity, larvae and ragged and feed on stems and leaves as adults. Damage is leaves (236). patchy. The larvae are not adapted to submergence and are Eggs are laid only on actively growing plants (49), aid true upland soil weevils isopposed to the so-called water damage occurs wiihin the first week after crop emergence weevils. They rarely become sufficiently abundant to cause (170). Larvae pupate more in stems than in soil. Seedling damage. Adults on rice foliage or the soil surface show that maggot attack is highly seasonal (206), normally peaking 2-3 the roots are being attacked by the grubs. Root weevils are mo after the onset of the rainy season. These pests possibly highly polyphagous (185). aestivatc over the dry season. Rice isonly one of their many Root heeth's. Soil-inhabiting false wireworms Gono- graminaceous crop and weed hosts. cephalumt spp. are reported in upland ricefields. The adults The rice stem maggot (hlorops or.'zae is adapted to and larvae normally feed on decaying organic matter, but temperate upland and lowland rice growing regions of Asia. land preparation for upland rice removes these sources. The In Japan and China upland rice grows in mountainous larvae feed on seedling roots, and nocturnal adults roam on regions, a habitat of this chloropid fly. The larvae tunnel the soil surface to cut off seedlings at ground level (82). This into tilestems. The stem maggot is more damaging during group of ground beetles is highly omnivorous; adults even the seedling stage. l.arvac hibernate in grasses. prey on stem borer larvae (183). True wireworms (Elate- File several dipterans reported as stem maggots in dry­ ridae) feed on living roots, but rice as a host has been seeded lowland seedlings inIndia (205) could presumably reported Infrequently (117). In Latin America, the highly attack upland rio,:. pclyphagous larvae of' Diabrotica rootworms are also Leaf urin'rs. I.caf miners are common on wheat recorded on rice. and maize but are rarely reported on upland rice (213). Three Root htu,,s. Upland rice seedlings can be attacked by the leaf miners are more prevalent on upland than lowland rice adults anI nlViphs of chinch bugs Blissusand Caenoblissus inthe Philippines. Pvudonalomy a asicanh', '. Vficata, and bro n bug Scaptocoris, which feed on the roots with and lIAgromrza or'zae attack at the seedling stage. The theirsucking nouthparts (80, 83). Three genera of burrower larvae develop inside the tunnels they construct in the bugs A tms. (,'eotonhus, and Stiharopus - are common parenchyma tissue. P. Vpicata and A.orvzac prefer maize, iil grasslands in the Philippines and may attack the first crop E/ehtsine, and other grasses to rice. . spicatacan become (rice or maize) planted after the dry season. Chinch bugs are abundant on wheat. most abundant after a dry spell. These insects are large and Leaf beetles. Chrysomelid and coccinellid !eaf bcetles can kill seedlings or stunt and reduce tillering on older scrape or otherwise remove leaf tissue as larvae or adults or plants. I hey prefer other hosts and rarely damage rice. both. This damage accelerates desiccation of plants. Tile Adults can also feed oii developing rice grains (81). larvae of sonic species are root feeders. Flea beetles Epitrix ('utworms. Soil-dwelling noctuid larvae hide under- and (lraetocnetna and rootworms l)iahroticaspp. feed on ground during the day to avoid predation by and birds rice foliage only as adults. Their root-feeding larvae have bccone active at night above grour, cutting off you ing hosts other than rice. The adults and larvae of the hispa-like plants at ground level (hence tile naiie cvtworn). .arvae Oediolmlpa spp. beetles of' Brazil and (hmootriha drag severed plants into hurrows. l'lie larvae have (=EPfi­ wide host /ach'ra) beetle of Africa feed on upland rice leaves. Oealio­ ranges arid pupatc in the soil..lgrotis ipsilon occurs pa/pa spp. larvae are leaf miners. (/mootriha is also p:'­ worldwide and appears to be the most common cutworm valent in lowland environments (11). Leaf beetles prefer pest of upland environments but is more prevalent in vegetative rice (81). lowland rice, particularly in dry seedbeds. The rice hispa Dicladi.paarnigera is not reported on upland rice and prcers IFoliar pests more aquatic habitats. In Central Africa, however, I)cladispm viridicyaiea attacks upland Other groups of insects are adapted to upland environments rice in the vegetative stage as well as lowland seedbeds. because they pass one growth stage underground, either as 7richi.pasericea is reported on upland rice in West Africa egg (in heinrietabolous species) or (in holomnetabolous and Madagascar (57). groups). The rice leaf beetle 1d1ea 01i'zae, riiuech like Chiorops Rice seedlibig maggots. AitJeriota flies occur in non- or.|rzae, is adapted to temperate regions of Ashi and is fhoodcd environments in Asia and Africa (180). Feeding of usually a pest on lowland rice but seldom oi inpland rice in larvae, one per iller, causes deadhearts. Flies, as opposed to the mountains (82). The adult hibernates on grasses. , only secondarily evolved as pliytophagous pests Arm.mworms. Species of grassland-adapted Spoptloera, froni a saprophagous Origin. ,Athr'nonalarvae secondarihv P'latysemta, and A i'thimna larvae create widespread epi­ feed onidecaying tillers, whereas larvae of stem moths borer demics. They can' defoliate ricefields, generally in patches. actively Iced on living tillers to cause deadlicarts. Seedling from early vegetative growth to harvest (41). Armywornms maggots will die ifthey cannot sever a tiller on which to feed pupate inthe soil and highly favor upland rice (34). Adults IRPS No. 123, January 1987 13

can disperse long distances to colonize even remote upland the end of the crop growth cycle. Therefore, late-maturing areas. They become more abundant than their natural varieties, staggered planting, and lack offloodingfavorstem enemies, particularly after favorable rains following a borer buildup. Many stem borer species can lie dormant in prolonged drought (196). The drought kills the armyworm the stUbbleafter harvest, and theirsurvival is encouraged by and natural enemies (mainly parasites), but the armyworm panicle harvesting (leaving tall stems) and lack of tillage can recolonize rapidly. after harvest. The natural increase insoil fertility from mineralization Stem borers disperse and lay many eggs. Many species are of soil N over the drought period promotes luxuriant plant polyphagous; however, the monophagous Maliarpiasepa­ growth to foster a rapid buildup of these highly fecund ratella,Diopsis Ivmgicornis, Scirpophaga incert/las, and S. species. Weeding rice increases tilelikelihood of damage. innotata can also be abundant on upland rice. ('hilo The parasites normally return to control the armyworms, Suplressalisdevelops fasterand becomes la rger when reared but not until after serious damage. on upland than on lowland rice (226). Outbreaks on upland rice have been reported in Ghana The lesser corn stalk borer :lasmopal tt /Itto.selh/s is (3). inzibar (38), Central Africa (36), lPanama (165), Brazil semisubterranean and is perhaps the stern borer most (28), Malaysia (196), and India (167). Defoliation may be adapted to upland rice(129). Highly polyphagous, it infests severe, often leaving only the base of stems. Armyworm maize, peanut, and cowpea. It attacks seedling rice (one larvae hide during the day under litter or in soil cracks. At can kill tip to four plants before maximum tillering) night they ascend plants to feed. L.arvae pupate in the soil. (195). (hiv and Maliarphastem borers prefer older plants. Thrips. Thrips frequent upland rice, but damage is less But Scirpophagaspp. will attack young plants even in the than in the lowlands. Thrips larvae and adults feed on leaf nursery. lasn opalls larvae tunnel into the stems at or blades by rasping, causing leaf rolling and stippling. Thrips below the soil level, mausing dcadhcarts. Larvae are not prefer vegetative stage rice, and their survival increases found in their turnnels inside sterns beca use, \vhei disturbed, during drought (heavy rain washes them from foliage), they retreat into cases made from soil particles bound by although drought stress usually outweighs thrips damage. silk. The cases areattached to the tunnel entrance (143). This Drought and thrips cause leaf rolling in upland rice (235), behavior probably evolved as protection from natural but thrips have been overlooked perhaps becaLse of their enemies. Larvae pupate in the soil. The lesser corn stalk small size. Thrips are most numerous indry season irrigated borer is more abundant during drought; damage is often rice. The combination of favorable plant growth and lack of confused with that of drought (233). This species prefers rainfall is ideal for thrips. In contrast, the lack of rainfall in sandy soils, arid its distribution within a field may relate to upland rice hurts the crop and eventually the thrips, whose soil texture (143). numbers must decline. Diatraeasaccharalis, aside from the lesser corn stalk In [.atin America, Frank/inilla rodeos attacks panicles borer, is frequently a stem borer pest in Latin America. before they emerge from the boot, calusing sterile grains. According to Teran (233) rice and maize are more preferred Stem hug~s. Black bugs are the most common group of hosts than sugarcaine, its namesake. rice stem-feeding -lemiptera in Asia. The most frequently Atci'tonac/rvsograpl//a is a less known upland rice stem mentioned species ----Scotinophara coarctaia and S.hrida borer in Asia with habits similar to E/asinopall s (82) - prefer aquatic habitats and therefore could not occur in because it is not found in lowlands. Ac#u,'tra has grassy weed upland rice as sometimes reported. Upland species in the alternate hosts. As a it is often confused with the same gerurs - notably S. tarsalis and S. scotti -- are dark-headed stern borer(hi/ polur(.standstriped stern morphologically similar and remove sap from stems of borer ('hi/ sup)lpsrcais. grasses and upland rice. The upland species are notably Other common upland rice stem borers are also reported more abundant in upland rice planted near forests, in the lowlands. The dominant species in upland rice vary In Latin America, tilelarge pentatomid Tibraca /im/'a- greatly with location. In Uttar lPradesh, India (78), and in tiventris causes deadhearts and whitehcads on upland rice. Japan (123), the pink stem borer Sesamia htfi,rens is more Both adults and nymphs remove sap from internodes of prevalent. plants more than 3 wk old. Tillers are killed by mechanical hIKenya, the principal rice stem borer is Chi partelus damage and by the entry of secondary microbial infection. (92). The lowland African sten borers Aialiarpha sepa­ This damage is often confused with that of lepidopterous ratella,Diopsis /vngicornis, and Sesamia calatnisnis are less stem borers (233). Eggs are laid on the leaves and stems, abundant Oimupland than on lowland rice(217) but perhaps Nymphs are gregarious. This species can also be found on cause greater yield loss because plant injury is greater if irrigated rice. Drought reduces population buildup. combined with drought ,',css (92). The stalk-eyed stern Stiem borers. Lepidopterous and dipterous stem borers borer D)ivpsis is more prevalent in wet habitats (6, 1I,217). are widely recorded on upland rice, but cultural practices In Central and West Africa, Maliarpha and Sesamia and alternate hosts determine abtndance. Except for calanistis are the dominant stern borers of upland rice (8, 9. Elasmolmll)uS Iinosellts, numbers usually build up toward 10, II). (lilo di/h/'lfilineui" and Chi/o partellus inhabit the 14 IRPS No. 123, January 1987

upland savannas, whereas Chilo zacconio prefers the showed that Pelopidas mathias prefers an upland habitat lowlands (II). A'aliarpha is most abundant in the mangrove (155). Parnaragutata seasonally migrates between lowland swamp habitats (91), coastal regions (16), or rain forest and upland areas and is equally abundant in both environ­ zones (II) of Africa and appears to be the homologue of ments (164). Scirlpophaga incertulas. The yellow stem bor,:r can also be The green horned caterpillar Melanitis leda ismene is prevalent in upland rice (120, 167). The sugarcane borer more abundant in lowlands (212), but our experience in the Eldanasaccharinais so far only a potential pest of upland Philippines suggests that Pelopidas niathiasand 11elanifis rice (91) despite Grist and .ever's (82) warning. leda ismene are often more abundant in uplands than in Grasshotl)ers andI locitas. Short-horned grasshoppers lowlands. Bamboo iaunbuiva spp. annually sustains P. and locusts -- Locv'ta, Patanrga, Schistocerca - layeggs in mathiaspopulations in the Philippine uplands. Adults feed the soil, inhabit grasslands, and develop into swarms to on nectar from flowers and migrate to upland seriously damage rice during the upland rice from time to time (173). Large late vegetative stage. numbers can destroy a field, leaving only the stubble In low 1 iPi' erlled; h i-1 " ;';d dill L CtI i 7-.0 ,l(, c'muoroer s, giassnoppei.rs an cut panicles (173). Farmers may become numerous, probably because of high egg predation. not plant if locust swarms are imminent. Prolonged dry Egg and larval parasitization are often low. Butterflies can weather followed by favorable rain favors development of readily disperse long distances to seek renlote upland large swarms (173). Usually breeding in di) grasslands, they , ricefields. However, they have low biotic potential, and even disperse toattack crops including upland rice(191). Eggsare it predation rates are low the\ rarely become abundant. susceptible to desiccation d uring drough t and toinundation Upland environments, with their diverse microhabitats, can during seasonal rains. With favorable soil moisture over provide more favorable sites than lowland rice plains. consecutive seasons, natural enemies ca nnot prevent a large tutterfly pests of rice have broad host ranges and become population increase, leading to migratory swarms (64). dormant Juring urnfavorable times of the year. Most grasshoppers on rice are adapted to uplands (107), I'ol.'*hagrotslaidopwra.The larvae of many poly­ but lowland rice grasshoppers 'iervgl(rlhusspp. and O.':ra phagous moths are reported to defoliate upland rice: spp. occur occasionally on upland rice (2, 82). They lay eggs tussock moths l'sa/ispelnnatula, Iawlia sUi.fita, on rice foliage. Nymphs are scmiaquatic. . luproctis 'irgtwlta,E. mnor,and P vathorrhoea;slug caterpillar The gryllid L'sctrtus coucinnus is a pest of lowland and litotiahicolor,hairy caterpillars Nisaga sinp/' .vandA mata deep water rice in Bangladesh and Thailand. In the spp.; and wooly bears l)iacrisiaohliqtia and Crvatonott.s Philippines, it damages upland rice grown in isolated gangis. Many of these polyphgous larvac migrate and are pockets near lakes. The gryllid nymphs and ad ults feed on transients in rice, including Helh*Whis arm iera and 1t. zea. the central portions of folrage, leasing only the midrib and highly pol phagous species leaf. nargins. (82). Gall nidge.s. The Asian rice gall midge Orseoliaorrzae Leal //ler. Pvralid moths whose larvae fold leaves to (88, 131) and its African counterpart 0. ortzivora(6, 11, 12, make a feeding shelter occur worldwide (217) but arc 32,217) prelerlowland to upland rice. Gall midges in rainfed appareritly most common in Asia. Leaffolde rsattacking rice areas are prevalent only during tle rainy season and disperse in Asia are a complex of species (26). (Ctalhahcrci s poorly. Their alternate hosts (wild rices, Leersia,Ischaetnuan, inedinalis, the best-known rice Ia fl'olde r in Asia, may prefer and Pasa/hmt)are mostlyaquatic grasses and are less likely grassy weeds over rice (86). It is a late colonizer of upland to be near upland rice areas. In an area of mixed upland, rice. U1arasmja e.viua, a lowland species (220), nreflers rice floating, and rainfed lowland rice cultures in northern to grass specres (86) and colonizes upland rice during il: Thailand, upland rice, which is ilanted a month earlier, was early growth stages. .llarasnria patnalis is the most corn- attacked first,but subsequcent populations were much lower rnonly encountered species in upland rice in the Philippines. than in rainfed lowla id or floating rice. Low numbers of gall Perennial grasses are their alternate hosts, and they can midges passed the dry season orr perennial weeds such survive year-round as in upland rice environments. The leaf- Paspahim (88). folders 11arasmia ruralisand Alarasmia spp. are least S/mot (hid(s. Reports of aphids attacking upland rice aburndant. Dormancy is unknown in leaffolders. Cnaplha- plants are few (249). Without maize, the maize leaf aphid /ocrocis medhnalis, however, is known as a migrant (90). Rho alosihhunm maidis attacks upland rice in the Philip­ Leaffolders remove photosynthetic tissue; attack during the pines. The plum peach aphid IIrsteroneuraselariae was flag leaf stage is particularly injurious. Popurlations are reported in Sierra Leerne toinfest leavesand tnriperied grain normally held in check by natural enemies, but leaffolders (7). Siphaglrctriaecaused transverse linrea r necrotic striae become parlicularly abundant in conditions of high plant iir rice leaves in Italy (172). I-lea v rains held fertility and shade, , populations in check. Rice is not a preferred host for shoot aphids. Butterflies. Skippers PelopidasInathiasand Borhofanta Leafhoper" andI /)anthol)pe:rs. Cicadellids and del­ are more prevalent in African lowland than in upland phacidsdispersecasilyand readilycolonie upland ricefrom habitats (6), but extensive studies in Japan and China nearby lowland areas. A study in Sarawak (241) showed IRPS No. 123, January 1987 15

similar species in upland and lowland rice. Nephoietti.x" The principal vectors of rice virus diseases generally have species dominated. The most prevalent species, accounting narrow host ranges, and alternate hosts in the grass family for 90-95% of the sweep net samples, w:re N. virescens. N. are mostly annuals, dying in the dr3 season. Some virus umropictus, Nilaparvia hgens, So'atellafiircifera, Cfana vectors such as BPH can disperse lontg distances, but few do. s.ectra,and Recilia dorsalis.Lowlands had higher popula- Th,: restriction of upland rice and weed hosts to the rainy tions. Katanyukul and Chandaratat (120) monitored hopper season, the small area planted to upland rice, and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results. A comparative study of plant- diseases in upland rice. hoppers in Fiji showed more brown planthoppe; 'BPH) and In Latin America, Sogctdh,s o(rzicola and S. cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease, but reports on upland lowland rice, and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia, areas with standing water (89). Peru, and Venezuela (35, 42). S. ,rricolao,:curs in Brazil Hopperburn in upland ricefields in Asia is rare (84). In on upland rice, but hojaI blanca has not been reported (67). Batangas, Philippines, isolated fields of upland rice become S. or'zicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH. A report from Fiji Asia. WBPH, however, is not known to transmit any v'irus cites hopperburn from BPH and WBPH (250). In these two disease. situations, upland rice was grown on highly ferti!c soil, and In Africa, pale yellow mosaic virus is endemic to swampy N fertilizer was applied. Hoppers respond to better nutrition, lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles. A minor fertilizer. vector, a flea beetle Chaeiocnema spp., is an upland rice In Asia, upland rice ,ras are normally not far from pest, and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers. Otherwise hopper contract the virus. species with greater polyphagy than BPH or Nelhotetti.v Spittle bugs. Cercopid nymphs producc a protective virescens would be dominant, frothy covering that looks like saliva. In Latin America, In Africa, leathoppers and planthoppers do not become spittle bugs damage upland rice. In parts of Brazil where abundant on upland rice (6, II) probably because lowland upland ricefields are surrounded by pastures, adults rice is not widely rlanted nearby. The only reported migrating from grasses can damage rice severely and cause hopperburn ..as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98). Hoppers, colonization (194, 195). Dispersal to rice occurs even if however, are a gre'ter problem in Latin America, where pastures are verdant, and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168). Nymphs are rarely abundant enough to phocephala spp., Hortensia spp., lEvitianus ohscurunervis, damage rice. Regular rainfall favors their development Balclutha spp., and Draeculacephala spp.), 'vhich then (233). disperse to cause hopperburn on seedling rice (68). Rosetto et al (195) reported Graphocep7hala spp. and Hortensia spp. Seed pests to be equally abundant on upland and lowland rice. Alydid and pentatomid bugs are seed pests of upland rice The leafliopper and planthopper epidemics in Asia that worldwide. In Asia, Leptocorisa sp. prevail. Their habits caused severe losses from hopperburn and vijus di:;cases make them particularly suitable for upland rice: 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice. Except in a few isolated cases (160), upland season before rice sets its grain); 2)adults are long-i,,'ed and rice has been spared from tungro, grassy stunt, and ragged mobile, allowing them to find isolated plantings 0' rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs, only birds and rats appear to have ;"­ and BPI] (123). Virus diseases have occurred in upland rice capability), and 3) adults aestivate in forested areas or where lowland rice was nearby. Rice dwarf disease, however, sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187). grassy weeds are present (I199). Rice seed bugs concentrate Virus diseases can be perpetuated only on livir, plants or on small-scale upland riceficlds because they can actively insect vectors, and no living rice - plan's, '.tooned, or search them out. The prevalence of groves of trees charac­ volunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall. sites, keeping the seed bugs close by. The smaller brown planthopper Laodcelphax striatellu.,; Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18). It spreads black- because: 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice. Early planting of lowland rice rice during the milk to hard dough stages), 2) they lack nurseries spreads these viruses, but upland ,ice has remained aestivation sites, and 3) weedy fallow areas are limited. uninfected. The low'ands of Asia are dominated by L. oratorius, 16 IRPS No. 123, January 1987

which lays its eggs hign on tile foliage (199). L. acuta and L. are more abundant in lowland culture. They can specialize solonlonlesis oviposit at ground level, the former on litter in rice because they are highly vagile and can attack all and the latter loose on the ground. These egg laying habis growth stages. By specializing in rice, monophagous rice explain theirtenvironmental prefcrencecs(214). In Japan, the pests can outcompete related but polyphagous species. postdiapause adults of L. chinensis migrate from moun- Intensive andI extensive lowland rice culture has favored tainous areas to upland ricefields during floweiing (1 14). monophagous species. Riqwla alhinella (Cramer), the only Pentatornid bugs prevail in Latin America (8 1,82, 195). monophagous species in Latin America, is rarely reported and in Africa, pentatomid and coreid bugs arc equally cited on upland i.ce (I1). (6. 82). l)amage is often characterstic of the species (5). Because of its limited temporal and spatial existence, 'rcoris ventralis and M'nida are spp. the common upland rice has not favored the evolution of specialized pentatomid seed bugs of Philippine upland rice, but they species. Clhang(44)shows evidence that upland rice has only prefer lEchinoch/va spp. grass seed to rice. ,Ve:araviritda recently been cultured by man from the lowlands: thus, pests occurs worldwide and h,often more abundant in the uplands have had little time to fully adapt to the crop. All but the (54). We saw few reports of seed bugs on upland rice, but lowland-adapted insect pest species have alternate plant see(d bugs apparently opelrate cqually between by, land and hosts to rice. Root pests with limited mobility are highly upland rice. Good dispersal is necessary ior species that call polyphagous. Pests that specialize in one growth stage feed for only 2-3 x,.on a crop and mu,st first find those crops e.g., secdling maggots, seed bugs are oligophagous or at ripening stages. polyphagous. Other pc:.t species feed "nflowers. In Mato Grosso, Brazil Most alternat, hosts of rice pests are annual grasses, (194), and Santa Cruz, Bolivia. the panicle weevil A'eo- which also are seasonal. Upla.,_'rice is also a host, but not hariuia aml/litarsis attacks rice during flowering (68, 233). the preferred one, of the most highly polypliagous The spccies: larva e rice sten horers: adutlts feed at the bases of soil pests, grass hoppers, armvwarrs, and ,Vearaviirielda. spikelets, caIusing empty grains. Also inL.atin America, the The d ry season poses igreat cydnid ,-lkiituttatralus obstacle to ani upland rice isa seed bug as anad tilt and nymph insect's survival, and only sonic perennial plant (194). hosts are suitable food during the off-seasoin. Plolyphagy allows A number of beetles ecd on poilen - coccinellids. upland rice insects to suLrx-ixe during thC rainy season when lialr'oti'a, Aulac lhora, and :Monoh'pl-a (55) - but plants- both annual and perennial -areactiv'elygrowing becatise riCe is self-pollinating and fcrtiliation occurs before and therefore more nutritious. ,:pikelets open, they pose no threat to yield. Loiugevit.r. The ability of a;.i insect to live 2-3 mo without undergoing dormancy helps species sich as root-feeding 'I!ST ECOLOGY white grbs (as larvae) and secd-feeding bugs (as adtilt,) to survive unfavorable periods. White grubs arc not highly We now look at characteiistics of the life cycles of upland fecund and their life strategy is to improve survivorship of rice insect pests to lea rn hew each plays an adaptive role in limited progeny. Their subterranean habitat hides them upland rice ecology. from manv natural enemies. lBtut root tissue is not highly nutritious, so they ltilst eat great Life history strategies quantities and need a large insect biomass to digest it. Other soil-inhabiting, root­ Or'zasativadoes not tolerate drodgh: well, so upland rice is feeding insects have short life cycles but are more fecund. highly seasonal, normally grown in the wettest months of Seed pests need greater longcvi vas adults to locate ahost the year. Usually. rice is present for less than half of the year. at early grain dcclopment arnd give sedentary offspring The rice-fiee fallow poses serious problems of survival to time to develop. Few groups of upland rice insects live long upland rice insects, which have evolved att least four in active development. Polyphagy and longevity mechanisms only to ov'rcolne tle cvcl ic-l lack of a host: increase their ability I) polyphagy, 2) to survive during the rainy season. longevity, 3) dormancy, and 4) vagility Other mechanisms are needed to survive the dry season. (dispersal). orantc. In the tropics, the nonrice season is the dry Pol'dt/)agy. l'olyphagy is defined as having hosts of at season, while intemperate areas. tile off-+season is winter. least two botanical families; oligophagy, of more than one Many upland rice insects undergo dormancy inlsummer gents: and ionophagy, of only one ventus (40) - inour (aestivation) or winter (hibernation). Dormancy is simply case, Orvza. Most rices, wild however, are illi tiC and inactivity: during winters with prolonged temperatures theiei'irc are normally far removed from upland rice below 10 C, insects find shelter arnd cease movement. hahitat,: othewise more would be alternate hosts for Activity is resu med with warmer t empcera tu res. A deep state uplanJ iice insects. Thte highly mornophagous rice pests ­ of irnactivity, termed diapauise, occurs in sole species that Scir.'iol)haita hiceritlas, Nilal)arvaiahiens, Iand Nealo-- prepare physiologically, (havxing low metabolism and tetti.y vircsctw,:s in Asia and Aalirpha secparatella and convcrting gonad s to fatty fuel reserves) for the unfavorable 1ioj)sis lngicornis inAfrica - occur on upland rice but season (148). They enter inanticipation of un­ IRPS No. 123, January 1987 17

favorable weather and resume activity after favorable Armyworms, skippers, and locusts actively travel hundreds weather begins, usually in response to photoperiod or of kilometers in air fronts such as intertropical convergence through a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237). Not all individuals of a population have the breeding grounds (31). Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is planted. terminating diapause -- this is seen as a mechanism to These large insects are active fliers. Smaller insects such as prevent total mortality if a false cue appears, such as a thrips, aphids, and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind. season (154). Otherspecies ean readily travel tens of kilometers - stem Probably, more upland rice insects undL :go dormancy borers, leaffolders, leafhoppes, cutworms. spittle bugs, leaf than we realize. Candidates are Atherigona spp., root beetles, seed bugs, ants, and termites. aphids, root weevils, crickets, leaf beetles, thrips, and seed The least dispersive species are white grabs, mealybugS, bugs. Insects known to entervarious states of aestivation are seedling maggots, crickets, weevils, and gall midges. leptocorisaspp. adults (199), Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9), Scirpophaga innotata larvae (117), Chilo partelhtsv enable it to survive and adapt to the changing upland larvae (21), ScirpopThagainccetulas(179), white grub larvae environments. Several evolutionary' avenues lead to fitness (134), Patangasuccincia adults (6, 82), and Schistocerca ('Table 2), but there is no reason to believe one set of gregariaeggs (82). attributes is better than another (224). Insects known to enter hibernation are LodeIpha.D striatellus nymphs (124), Chti/o 'Tvlpre.ssalis larivae (125), Drought Sesaia inferens larvae (146) .... but not S. calavisis(2), Well-drained rainfed upland soils are subject to drought IDiaraea saccharalis larvae (75);- Scirpophagatcerlttas if rains do not fall within 2-3 wk. Prolonged drought larae(5 lacchrli s lae followed (75);sa,SlO.hagasshopers by favorable rains stimulates armyworm eggs (82);and felo pidas athias pOpae (232). breaks, but mealybtigs, root aphids, and andthrips locust become out­

Lcpmotorisa oratorisand L. acuta adults aggregate on numerous onlyafter dry spells, for two reasons. First, heavy trees, sugarcane, or other shady, moist sites. L. paluwanmsis., rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan, Philippines, lives on the mealybugs, and soil-inhabiting root aphids and root mealy­ grass Brachiariamutica (Forsk.) Stapf in the absence of bugs. Drought eliminates this source of mortality. Second, rice. The adults can live for 5 mo. When aestivating, their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed. They will, proteins into soluble N compounds, which enter the phloem however, puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245). Greater search of water. This behavior has led to errors of the host nutrition, therefore, leads to greater survivorship and range of rice bugs (I). Aestivation quickly terminates with fecundity. rain, probably meaning aestivation is not deep. Small rice area Some stem borers aestivate as mature larvae in rice Except in the highly mechanized, large landholdings of stubble or in straw stacked as livestock feed. However, Latin America, upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora. The small fields result in higher ratios dry season even if the stubble remains undisturbed. If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice, few stem borers ricefields. Upland ricefields themselves may be intercropped survive. Rainfall terminates dormancy, but it normally with cereals, legumes, or root crops. Mixed intercropping takes several weeks for moths to develop and emerge. with a wide variety of species also occ", in tribal slash-and­ Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50, 74). to construct pupal chambers in which they aestivate. The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness, so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture. Aestivation terminates to minimize weeds. Rice requires more tillage than does with the first heavy rains, but as in stem borer.,, 3-4 wk pass maize because rice competes less well with weeds. Upland before adults develop, rice is more a subsistence than a cash crop, and small areas Insects may respond differently in each region to cold (I ha) can feed a family. temperature or drought. Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56). Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders. Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions. well as aestivation sites for LetTocorisa spp. (199). Field i/agiliti. The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat. colonizers and a habitat for natural enemies. z 0

Table 2. Life history patterns of upland rice insects or insect groups. 00 Host range Longevity Dormancy Vagility Fertility Monophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High (Oryza spp.) (grasses, sedges) (angiosperms) (<3 wk) (>2 mo) (<1 kin) (<50 eggs) (>500 eggs) Ants Termites x x x x x Mole crickets x x x x x x x Field crickets x x x x x White grubs x x Root aphids x x x x x x x Mealybugs x x x x x x Seedling maggots x x x x x Leaf beetles x x Armyworms x x x x x x Thrips x x x x x Chilo spp. x x x x x x Maliarpha x x x Locusts x x x x x X Gall midges x x x x x x iVephotettL" i'irescens x x x x x x Leprocorisaspp. x x Leaftolders x x x x x Sogatella x x furcifera x x x x x x IRPS No. 123. January 1Q87 19

All this appeals to favor insect pest buildup. but the notwithstanding the release of stored nutrients in response contrary is true. Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders. populations respond exponentially with the proportion of land devoted to rice up to about 75% of the area, then the M POR ANCE I IP1A\ 1) RWI P1St S rate of response declines. In most upland rico areas, tie rice crop occupies less than 50'i of the area- therefore, the Visits to upland ricefields normally reveal 1kw insect pests. potential for population buildup is low. This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields. insect abundance using annual light trap atchcs in upland. Another outcome of the highly diverse flora and ph..,,ical rainfed lowland. and irrigated lowland sites in the Philip­ environment of upland rice isthateach upland ricearca has pines. Although only one upland site was studied, data a unique composition of insect pests. showed equal or fewer insects at the upland site than at the For example, in Tanauan, Batangas, Philippines, the rainfcd lowland sites, and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF). armvworm (Table 3). M Lltiple regression analysis showed that the key Aythntnaseparata, WBPH. ants So/li psis gentitata,and factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata. Sixty kilometers away in intensity variables were the number of rice crops grown per Real. Quezon, the main pests are seedling maggot Athe- year, followed by area devoted to rice. Table 3 shows that rigona or.'"ae. flea beetle (Thac'itcm'nra hasalis, thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurn'mis, and rice bug Leptocorisa was planted in only 20Vi. of the area. Fertiliter and oratorius. In a third Philippine upland rice site in Claveria, insecticide use had little bea ring on the trend. The armters at Misamis Oriental, Mindanao, the main pests are ants the upland site applied 60 kg N ha, higher than at most Solenopsis gemtinata; seedling maggot Alieriona orvzae; irrigated sites. white grubs Holotrichia in)iU/aiuoan t and Lucopholi.v Insect occurrence at the upland site was :omparable to irrorata: stem borers Sesamia ifiA're'lr.. (hi/o auricilius,C. that at the Cagayan rainfed lowland site, parlicularly in 1982 stp'essallis, Scirpophaga incertulas, S. innoiata, and when drought prevented many farmers fron planting(only ,4c ,ona chr'rsurapella; root aphids Tetraneura niriah- 40(,%of the rice area eventually was planted that year). M ore dunrinalisand Rhupa/osiphunm r./iahbdminalis; lcaffolders favorable weather occurred in 1980 and 1981. which M'arasniapataltis and Chaphalocrocis Medinalis; army- supported greater rice areas and consequently more insects. worm MIthinna separata: and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato, oratorius.The great diversity of insect problems is typical Mindanao. where farmers plant 2.3-2.5 rice crops per year fer upland rice and makes control efforts more difficult, with irrigation. The Zarago/a. Nu'tva Ecija, site also had particularly regarding breeding for insect resistance. high pest abundance, and at this site the fields were highly asynchronous. Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance. Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils, which, when combined with eiratic rainfall, remain low. Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year. (Farmers in Claveria, Misamis Oriental, Mindanao. in costly soil amendments and land management. Often Philippines, tried asecond upland ricecrop i 1984,a ycarof upland rice isgrown far from markets, making inputs even favorable rains, but no harvest was possible b-cause of more expensive. Farmers growing a subsistence crop hosi- drought, blast, and birds.) Rice area isa!so bound to remain tate to invest in costly inputs to raise yields. low because of the high labor and power rcquirelmenlts to Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop. Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons, however.are unless tile risk of crop failure were reduced such as it was planted asynchronously, such as the Cla\cra site. with irrigation systems in lowland rice. The seasonal abundance of Upland rice insect pests The Tanauan, Batangas, site is atypical for upland rice. attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable, and because of Tanauan, FBatangas (Fig. I. 2). W13P11 was ,he most its nearness to Manila, farmers have cash resources f't'm common rice hopper, and its population peaked at the sales of vegetables to purchase tertilier. They applied an midgrowth stage as it does in lowland rice. D-Vac suction average of 60 kg N/ha after particle initiation and obtained samples showed an earlier peak than that from a light trap, 2.5-3.5 t/ha yields. The high fertility isone reason why RLF. indicating emigration. W13PI usually emigrate after the armyworms, and WBPH developed into large numbers, vegetative stage. Populations in 1980 reached levels that Farmers. however, used no insecticide on upland rice caused patches of slight yellowing in some fields. WBPH although they did on vegetables. probably cause hopperburn in Batangas. They immigrate The poor growing conditions fOr rice generally mean year-round. as shown by low levels in the light trap. BPH. poor growth and low fecundity for insect pests fIeding on it, however, were scarce in the light trap and not collected with Table 3. Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland, rainfed lowland, and irrigated lowland locations where light traps were operated daily for least I yr. at ,-I 0 Rainfed Rainfed lowland Irrigated lowland upland Pangasinan Cagayan lloilo Santa Maria, Cale. Nueva Ecija South Cotabato Tanauan. Caaringayan. Laguna Caaringayan Bangag. Bangag Bangag Buray, Santa Motiica. Batangas Manaoag 1982-83 Cabanatuan Zaragoza Avancena, Naniaman, 1980-81 Solana 1981-82 1982-83 Oton Oton 1980-81 1981-82 1981-82 Koronadal Koronadal 1979-80 1980-81 1979-80 1979-80 1983 1983 Insect (kerosene light trap catch Nephotcttixno.lyr)virescens 650 1,240 500 1,800 330 210 3,400 N. nigropictus 210 1,600 160 360 18,400 5,500 2,900 - - 140 90 20 - Reciliadorsalis ­ 1.700 - - 3.200 1,700 710 180 890 630 140 170 2,400 Nilaparratalugens 800 690 - - 4,200 4,600 130 300 520 750 180 60 3,300 Sogatellafiircifera 1,700 1.600 740 310 31,000 13.000 8,200 310 210 3,300 610 270 980 Cyrrorhinus 1,600 6,300 - - 12,000 6.700 600 540 3,200 290 160 330 Scirpophaga spp. 120 410 620 - - 21,000 50 410 150 180 220 270 12.000 Cnaphalocrocis 30 1.000 540 610 270 11,000 medinalis 70 70 200 140 100 6,100 Chilo 6 60 250 60 60 1.900 suppressalis 0 - - 30 20 300 200 0 6 3 160 - C auricilius 0 - - 0 0 - 20 3 60 20 3 0 Sesania inferens 0 - . .. 80 60 3 80 10 Cropping intensity, 51 - _ - Rice crops (no./yr) 1.0 1.0 1.2 1.0 1.0 1.0 1.3 Rice area (7c) 1.7 1.9 2.0 2.0 2.5 2.3 20 85 85 60 70 40 Fertilizer (kg N/ha) 85 85 90 85 80 70 60 20 20 0 0 0 70 Insecticide (no. of 30 30 40 60 40 30 0 0.4 0.4 0 0.3 0 30 1.3 1.3 3.0 5.0 4.0 3.3 applications/crop) 3.3 IRPS No. 123, January 1987 21

Light trapped Insects Drytnd __/ 2 (no/trap-nght) Suctioned Insects (no/m ) 50 -)o 40- SOW9 ,jO ffurciWO Light trp

D-Vc suction 20 4 to. 20

Egg porasitizahion (M) 75., 300 60 SOO&Re/Oftwciffr 45 30­ 15. - insects (notrop-night)

4 Nileporvata Iuen5

E'gg parastization(%)

Light trapped insects 2 ) (no /trop-nght) Suchoned insects (no /m 8 Cytorh~nuls hvidipennii 0 6 4 05 2

50" 0 40-uiInsects (noltrop-night) Ipanh pe g

Moohase 1. Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper 60)-t Nilaparvata lugens, and lea ffolder Cnaphalocrocis mnedinalis andplthpe g 20 L I II . . ______I I I . parasites and mtrid predator O',rtorhjntus livi- Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern, daily 1 9 80o 19 8 1rainfall, and moon phase. Tanauan, Batangas, 1980-81. the D- Vac. Cyrtorhinus build up on the crop coterminous are collected inlight traps during full moon (109). with WBPH buildup. Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pests rainy season. RLF, not readily collected in light traps, was and those pest species are generally less impotant to upland prevalent only during the rice crop. rice than lowland rice, there is a general belief that insect Nephotettix virescens was more abundant than N.nigro- pests are less important on upland than lowland rice. picLus in the light trap collections, and on the crop, GLH rcview of limited data on upland rice shows an averag ofA population peaked at the reproductive stage, as did zigzag 10-21% yield loss from insect pests, on a par with that leafhopper (ZLH). Crtorhinus tracked the populations of lowland uee determined by the same methodology (132)of planthoppers and leafhoppers. Both GLH species were (Table 4) . McGuire and Crandall (144) made a similar collected year-round in the light trap, indicating immigra- estimate for Central America. Cramer (51), however, tion from nearby irrigated rice areas. Possibly low because of the estimated 3.5% yield loss in South America based on a populations, insect number did not correlate with similar pest complex - stem borers and seed bugs ­ moon phase, although reports indicate that more hoppers attacking irrigated rice in North America. This estimate did 22 IRPS No. 123, January 1987

Light trapped insects/ D~~nd (no /trap-night ) 2 St plht Suctined insects (no/m ) D-Voc suction 1520

1o 0 Lih trap 5 ~105 o5 00 Eggparositizotion(%) 50 40 NephOtgtha virneens 30 20 10I0 A,- Insects (no./trop -night) 4 Nephatelh, nigropc¢ i

2 o I vV'^J -. -- "VA____ ^ f _

Light trapped insects (no /trap-night) Suctioned insects (no /rnE,

108 ReCdO doRsol-120 6 15

4 10 2 05 iwxALAi& 0 Light trapped insects (no/trap-night) 2 Suc! oned ,nsects (no/m )

8 Cyrtorhmnus 10 I6o, nn] i00.75

4 05

2 025 0 0

2. Seasonal abundance measured by light trap Roinfall (mm/d) Moon phase and D-Vac suction sampler of green leafhoppers Nephotettix virescens and N. ngropicrus and zigzag I leafhopper Recilia dorsalis, their egg 60 parasites, and micid predator Cyrtorhinus livi- 40 dipennis set against cropping pattern, daily 5 Lf rainfall, and - moon phase. Tanauan, Batangas, Apr May Jun Jul Aug Sp Oct Nov 1980-81. I aMar 1980 1981 not account for soil pests and was not based on field data. in the absence of pests. Carbofuran granules are phytotonic Akinsola (I1) cited mainly data from irrigated rice areas in to rice (239). The prevailing hypothesis isthat carbofuran Africa but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer. The phyto­ and 30% from insects. However, losses in Thailand over a tonic effect isthrough greater availability of fertilizer to the 4-yr period ranged from I to 13%, averaging 5%(106). plant. Denitrifying bacteria are known to be more active in Yield loss data, however, should be interpreted with aerobic than anaerobic soils; therefore it may not be mere caution (184). The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69%) trials on farmers' fields under farmers' normal agronomic occurred in those trials with 12-30 kg basal N/ha. On the management practices, except for Bukidnon and Capiz. other hand, contro! with carbofuran granules may not be as where researchers used fertilizer. Batangas farmers are good as a seed treatment against seed pests, and perhaps atypical because ofhigh levels of fertilizer used. The trials in yields would have been higher if a systemic seed treatment Brazil were carried out at an experiment station, but insecticide had been used. conditions were similar to those of local farmers. The 1979 and 1980 trials in Batangas had two full- Carbofuran granules were used in every trial as a broad- protection treatments, one using carbofuran granules spectrum and insecticide applied basally with the seed. An the other using bendiocarb WP as a seed treatment. Plots objection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those pro­ insecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules. The other trials in the Table 4. Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines, Brazil, and Upper Volta.

Yield (t/ha) Yield loss (5) Location Year Cultivar Protected Protected Check Basal N Total Vegetative Reproductive Ripening (kg/ha) Reference with without carbofuran carbofuran Protected No with carbofuran carbofuran Philippines Pili, Camarines Sur 1975 Bursiging puti 1.4 - 1.0 Tanauan, 29 - - Batangas 1974 Kinanda 3.3 - 2.8 15 - __0 99 1974 Dagge 1.1 - 0.5 55 - - 0 99 1975 Kinanda 3.6 3.4 2.7 25 21 6 0 -7 0 100 1975 Dagge 3.5 2.4 2.7 23 -11 - - 0 100 1976 Dagge 3.1 2.4 3.0 3 -20 - - 14 0 101 1977 Dagge 3.0 2.7 2.6 13 4 - - 0 101 1978 Dagge 2.7 - 2.9 -7 - 0 102 UPL Ri5 3.7 - 3.3 11 - - -- 0 102 1979 Dagge 2.6 2.9 2.8 -7 3 23 18 -1 0 102 UPL Ri5 4.0 4.3 3.3 18 23 14 11 6 0 102 1980 Dagge 3.0 3.1 2.9 3 7 5 3 10 0 103 UPL RiS 4.0 4.2 4.3 -7 -2 0 -10 -5 0 Pangantucan, Bukidnon 1979 UPL Ri5 4.2 - 2.3 103 45 - 8 4 8 30 104 1980 UPL Ri5 4.6 4.7 3.6 22 23 2 10 9 30 104 Dumarao, t-apiz 1979 UPL RiS 3.5 2.5 1.1 69 56 22 7 7 30 104 Brazil Goias, Minas Ge~ais 1977 IAC47 1.9 - 1.5 24 - - 12 66 1978 IAC47 2.1 - 1.5 28 - - - 12 66 1979 IAC47 0.9 - 0.6 35 Upper Volta...6 - - 12 66 Farako-Ba 1977 IRAT10 3.8 - 4.0 -5 - - - - 242 1978 IRATIO 6.6 - 4.3 35 - - - - 243 Mean 21 10 Z

tJ

- w 24 IRPS No. 123, January 1987

Philippines, which had two treatments offering protection In the 9 Philippine trials where yield losses were measured with and without carbofuran granules, relied on further by growth stage, 10% of the yield loss (21%) occurred at the control with foliar spray, and not on seed dressings. The vegetative stage, and 51,7 occurred at each of the repro­ carbofuran-protected plots had an 18%yield lo,s compared ductive and ripening stages. Ibis result is similar to the with only 10% with foliar sprayS. This yield difference is pattern of yield loss in lowland rice measured by successive probably due to differences in protection against sown-seed treatments where the crop is unprotected at each growth pests (foliar spray; do not control sown-seed pests) rather stage but proticted during the other growth stages. than phytotonic effects became basal fertilizer wa:, used in However, the insect complexes r sponsible for vegetative only 2 of the 10 comparisons, stage yield losses in upland rice are entirely diftfrent than in The recorded yield losses were not related to yield lowland rice. The key vegetative stage pests in lowland rice potential (Fig. 3). One might expect higheryielding crops to arc aquatic. whereas those in upland rice are sown--seed have higher percentage of yield losses, but this w;s not tile (sown in soil), root, and seedling pests. This dift' rence may case. One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportancc injury from drought stress. l.ow yielding fields from d ought of upland rice insect pests. Many reports of upland rice pests stress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46, 108). insect-protected and -unprotected treatments. Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses, becatusese\eral studies drought stress than sown-scd pests. A second reason have shown that upland rice can readily recover from foliage concerns seed pests, which would have a greater impact on reinoval during that stage (1,17, 152, 165, 188). Upland rice low tillering, low yielding varieties such as )aggc. Low poscsses a hitgh obilitV to compensate for ca rly' loss of tillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal to of seeds; owe\er, 1)P1 R i5 is high yielding because it tillers produce a higher viel than plants \%ithout any foliage actively, which can close the canopy to provide greater weed removed (63. 192. 231). [ie reason for this apparent control. discrepancy, hovc\e;, is that yield loss at the vegetative stage does not octUr frol leat arca loss but fron pests Yield loss (/) removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for. 70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect 60 - as control. =-09 Insecticide seed treatments res;uled in denser plant stands, (n=21) which in turn created a selectively more favorable environ­ ment for 50 blast (69). This resuIt was parlticurla rlx'exacerbated when cirbofuran granules were used in seed furrows. Aside * ffrom a greater plant stand. ca rbofuran iay have auigmented the N le\cl and stimulated blast. If so, ca rbofuran should 40 - be replaced with another insecticide, or the seeding or fertilizer I * rates decreased. Other methods to determine the impact 30 ­ of upland rice pests wvcrc to interview farners or to develop single species • • correlations of population levels 2 and yield loss. Feneira (66) did both in Brazil. NIM)st (75(1) Golas farmers reported 20- insect losses in ufpland rice, and 25"1 said those losses S av'cragcd 37(1. [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4). For every I0 4 of seedlings removC d by leaf-cutting ants, a I ,7',yield * loss is predicted. Similarly, Ferreira (66) found that ai 0 - average of 5.3% thrips per panicle before panicle initiation meant 3 times more unfillh d grains. -IO0 I 1 -101 1 I I 0 2 3 4 5 6 7 Yield (t/ho) C;)N [ROt IM FI IODS 3. Correlation between upland rice yield and yield lols caused by Upland rice insect insects pests may be controlled by cultural, measured in 21 trials in the Philippines, Ivory Coast, and Brazil. genetic, biological, and chemical methods. IRPS No. 123, January 1987 25

Cultural control dry season plowing will have little effect on them. Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level. Cultural birds, chickens. dogs, and even man (field crickets, nole controls should come first in a pest management program. crickets, and white grubs a re delicacies in man tra ditional They have broad, stable effects, as pests have little pos- diets). sibility of overcoming them through selection of biotyp's The greatest degree i1 inIscct control fron tillage comes (138). On the other hand, some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs, and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters. stuble. Whitegrubsare mature, and tillage exposesthem to P/antin, time. The luxury of being able to shift the time of predators. Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years. Plowing prolonged rainifall pa ttern, lI copping intensity or underthe ice stulble helps it decompose, killing sten borer rnech.ani/ed land preparatiot. Gcnerally. planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites. insect pests. Insect populationsarelo\ aftcra dlv\'seoll or Tillage and clean culture, howes er, rcliove food for winter fallow delayed plaiting lets them build tip on termites. which then mtay attack a young rice crop. Scarab alIternate hosts that gi' vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields. No­ ratiris. This first flush i"of' weds cro\s lItxuttriartlv \with tillace 'iceCultClre Such is dibbling fa\ors the survival of' rineralied N released during the dr , season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil. available to the plants with the rains. It iliso ;igrtmitcally Zero tilla., hio eerl, conserves predators of'the GI.H (37). advanttgeous for the rice ciop to tap this trlLItlaI fertility /'/nt it, mct/toi. 'lo lessen the impact o0 soil insects, and grow vigorotsly to compete with w"rus and to tolerate brotdcasting seed or planting in f'tltrrows is prefc ired over pest infestation, dihlirig in hills. Crickets are large enotlgh to eat seedlings in Larly plantngs terd to escape seedling maggolts. Upland a hill. atrd tillering normnally ca iriot fill in tlhe missing space. rice in Bitangas is planted in Ma and .ine \\ itli the Iir'st Oftcn. dibbll ,rd is not well covered and iseasily found by rains. Fairners are highly mtoti\ated to plant early tocscape foraging insets,, ioIdents, and birds. drlitght and typlotns rica.r harc,,t. Arn Atrgtst planting Plan u/it sii'. Inc'rcatsini sced density protects tile crop would be .,evcely attacked by ..It/ rii,'m ori'vzo. At t\ti againrst seed and seelling pests. especially ants, particularly other Philippiie tipland sites Rea l, Qte/ot, antd (la\elit, if the crop will germinate and ermerge quickly. M isainis Oriental seedlig mnaggots arnrige ri'c boe t se hr trft/inll it,'I India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of' the rainy season. liaaiangas ftrmners rice alone (211). N'failC arid Urplald i:r_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines. he chatngC in pest status f.olii iniercropping is to lliarvet before the typhoor sea son peaks. ltighly locatioll-specific (133) arid the net el'fc cm.ianbe either Early plantings cornbind with early-iatiuring varictics nil. bcnelficial, ortdetriential (189). For example, BIatangias provide higher crp tolerartce for white grubs. Aestiatior farmers intercrip Itro in their ipland ricefields. A liorn­ terminates with the first rains, but 5-6 moare ineeded for the \. ..Ahis, n'rii,.' i,'otd-/ i (I ,iiiacius). which norlially feeds resting larvae to develop into beetles, emerge, and lay eggs. on taro. \%ill feed or miarbv rice plants. Intercropping and for larvae to reach the danriigii g "ird-instar stage probably would not afft0 seed pests. ,A crluanion crop (134. In lataigas, this normually iseiough tiite for harvest. planted \\ith rice and ha\ing gleater tolerarnce for root It Brazil, howecver, delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests. Acial-fei i ig pests \voultId be most affected by pests favored l,,, drought (66. 195). Cropping internsity in intrer ropping. Shoot aphids, leaf beetles, sten borers, and Bra'il is low. and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts. On the niost stable rainfall period, reducing likelihood ol'd rought. othlt halnd, the corlipalior crop stch~ sitniai/e Iiiv be more In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137). before rice, and althNgh it woultl be desirabh to avoid 11'ec/hi'. Weeuling luring the first month after clop infestations by planting rice cuatl., l'ariners lack the labor cstablishent will force pc,.ts such ;is anyvworris. which and time to do so ol a low priority crp. prefer grassy \cetds. ontor tle vorling rice. Soil i&secs will 77//age. Uplild rice Stils ae light textured and Call be mvnye to the rice crop. Cleani citrltirc also ,lay lore territes plowed duiring the d rv seasor, notl only to grin time for to attack rice. I .u cetincti . "in\ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13). soil insects arid weeds such is ntsedge that foim rhizomes. I''rt'liz,r. Itneatsing N Iciiliier oin hlwland rice: has Htowever. becta use most soil insects lie below die plow layer, fAtwvored Irigher populains of pliithoppeiv. Imlhoppers, 26 IRPS No. 123, January 1987

and leaffolde rs. However. because of the need to temper plantings up to 3 mo apart and use a set of \atieties that f'ertilizer use on upland rice *.o prevent blast. fIrtilizer rates mature in 4-7 mo. Insect pest danage, particularly from will be lower than is comonia r for lowlad rice, causing stem borers, is greater in Misarmis Oriental. Also, seedling relatixely less hopper and IaIfTfoide r population buildup. maggots damage rice there because rainfall favors frequent However. inupland rice areas without a history of blast, planting of mai/c. In Batangas, rice escapes seedling such Is Batangas. farmers apply high rates of N and maggots bCca usC of' distinct drs season and synchron0us conscqtrently the crop has high infcstation levels of WBPI] early planting with the onset of ain.s. and RI.F. The fhrmrcrs leec insect buildup by splitting the Therefore. combinations of cultural practices ea rly fertili'11r in0to two to three applicatiois bit do not applV planting, synchronous planing, crop rotation, and early­ fertilizer at the vegetati estage because this prodrces a tall maturing vaneties protect the rice crop against most crop which ",illeasily lodge during typhoons. Greater crop insect pests. fertility leads to greater insect survival (better nutrition). larger insects (to cat no re foliage), arid greater fec titd t (to Ilmant resistance lav nmre eggs) (128. 145, 171). Not all insect species respond Beca tse upland rice has few specialized insect pests and equally. hov,'r. Increasing rates (if N, l, anrid K increased many insect types attack the crop worldwide, regional Dialra',a s-acchralis and ('/iaeiur.,e/ma flea beetle but breeding and strong internatiornal cooperation are required. reduced thrips (66). Zn rcdLuced deadheart density from the Sown-sced pests arc economically controlled by insecti­ lesser cornIstalk borer. cide seed treatment and should the relore receive lowxpriori ty Fnldingi. Although most Upland access rice areas do nIot hawxc inbreeding objectives. to wa ter floodiig the fields is rR..rrnriierdcd in areas Root pests are normally cost ly to control with insecticides, of Brazil 'or root bug and white gib control ( 195). but finding resista t sonres mayvbe di fficult. Tanaka (229), Crop rotatioti. Tillage before planting a crop after rc for exa -ple, fi led to find \varieties resistant to root aphids. harvest will control stern boreis and other pests renain, .n Selecting for large root bioniass perhaps should be the the stubble arid will unearth soil pests. Planting a non- strategy fo,r pests sLuch as Whlite grubs that renliove roots. gratriiaceous crop after rice is recommncrded for termite Fainter(174), howcvcr, cites varietiesof crops tolerant of'or control (66). Crop rotation is less nttdcd in upland rice resistant t(i root pests. Some sorghun varieties are high in culture than in lowlands because atcrop-free period is cyanide, so rice sorghun might incorporate broad­ assured even after harvest of crops following rice. spcctruri chemical resistance. .t1ulchilt' \'iryl plastic irli irig designed for weed coitrol ii upland rice inJapan resulted Seedling vigor and drought tolerance are high priority inanri ntr liyhigh breeding objectives, so t,..upland rices should have large r infestation of' (ihilo sulytresvais (93, 9-1, 95). NIulched root systernsand be able to tolerate ligherlevels of root loss. Upland rice grew more luxuriantlv and attracted more Sonie root loss inolder plants is beneficial if it stimulates (Avipositing mothis than u l ched rice. Also, the rice plant new root developnrut and therefore crihances intake of soil and stein borers imatoured 10rC quickly and a partial nutrients (53). generatiori af vouing ista iNoverwintcred in Lotili--malitr.t, varieti.%. the stubble. The rice seedling inaggot rti/'L'otia spp. may be Quickly maturing crops reduce controlled thirough resistant aittics. A hi rgc breeding the tmbCr of pCst generations that Calt build up. Rapid effort on sorghniu has had fair success ini developing crop establishment ol'an ca rl v-natu ring va,itvty if)pa 11icLa r \'arreties esrstait to A.soccata Rodani (251). A field trial will lessen white gib and stem borer damiage. White grub comparing 10 upland ricc varieties showed differences larvae intle last iistar are highly destIruCtive, so aii early- aniong varieties va r.ing from II to 4("- damaged tilleis iiatri ng variety will escape serious root loss Stem borers (215). Hovver, resistarce to .1I/ill-0.r gtia species may build 1ip slowlyx in rice to ,ecormic abundant in a late- rot cross ox-rti otier. Maturing Shiraki (210) reported that upland i-ice stens wcrc crop anrd caiuse whi tlieads. Numibers of some harder species increase exponeitially with each succeeding genera- ti. so a xarietyexhat atures tr than lows-land rice stems,perhaps becaIusc (f a higher silicori Iwh o earier thian another will content, and thus were more resistant to yellow stern borer. ha\c less insect damage. Early-naturing vaiicties, however. hc tortia Iow tillering (f uplaid iice xarneties prodtres will not lessen seed pest daiage, a either ii the soil or oina higher percentage of' infested tillers than would occor in standing crop. higher tillering lowland iice under a siiilar egg density. The .St'rwhoi,'/ hAIJtt.am '" Inrs.c.'ts tcd irig on thlie aerial corollary, therefore, is to select for high tilleriig upland portions of rice plats disperse from carly to late planted rices. fields. A.\Sinsect iiiibCirs increase exponentially with each Stemii bor amcstheh main gtscnrauilttm, upp e wh rc eig'lbrig f rmc~s1( 1 ,204 )but nticl oifth e sctarges reen irig isbrecders ddo inAfrica (57, generation. otie unde r lo wland pests build tIP where neighboring fihrTe9,r1sC ' stagger their plantings. II Batangas, farmers co d t osrm s wih A h r~ o plant upland corditions (11, 96, 218). As with ,.lth'rinrnw,l speciespe i s mayn a rice within I iro, use 120-d s-arieties, and plant at the start of have to be dealt with independently: however, initial results the rainy season. Ini Misarnis Oriental, laricrs stagger their show cross resistance between African and Asian species (92). IRPS No. 123, JattuIny 1987I 27

In Brazil, screening Ior insect resistance focttses oil lou I parasites - scelionids on eggsand sphecids on ilymphs and pest problems. Varieties tolerant of l/asmolwalpus ligi.- adults. Sphecid Larra carutmaria (SImitI hI. L. sellus and resistant to )iatraea saccharalis have been huzwuensis Rohwer, and l.sal.,ti/uh'ta \Willi ams speciali, in identified (70, 153). Current work also emphasi,es 7"iraca mole crickets, pa ralyzing them nd dragging the ninto thcir lim/ativetntris and Thmialspi.s (=Iosis) flavou'icta (66). nests as food lot their voting. ()tier sphCcid species Lins These insect pests Ilt onlV citise IIigh yield losses, tihey aurulenoa (Iabricius), .lot'. ma/i/ac (.Ashmead ). .1/. lh­ also ;ire difficult to control chemically ofr by other ilealls. l.s'se'llaus (Smith). and Ml. /la/l)orio.su. (Snithf speciali,,e ManV upland rice va/ieties aic early maturing and are in field clickets. resistant to pests such as gall midge (I Xl ). Early mIturity is White grUbs,O the other hand, ha.\e Icti scriuns enuigh often a highly desirable trait to esuilpe pest buildup, but it is to have been extensively studied for cotttroi 1W nati al not true resista nce, b.ca usc ilt hose va reties are plinted late, enemies( 112, 140, 141). Scoliid parasites C("a//jsoM/ci.spp. they will be itsdamaged as ain early planted susc ,'ible have beetn intrtduced itto tilePhilippites for control of variety. Ielxl ]udisiro la . The rcsearchcts w Intttrdced them le P~erhaps tileimost cosiiiopolitall of' all upland rice pests during illonthreak era claimed that the white gtti \\as are the sced pests (ripenling stagc). which pose a particular controlled (142), howevcr, it isnormal for pest epidemics to problem beta use scuds re ised IsIO for hitnit4ans. Ayl' subside' and there was no diect evidence to indictte that tfe toxic substatice introd tcld geticticllyilmay' also be toxic to parasite was responsible. A tmore promisitg titctllod might nal iniless it caln he placed old\ in thfehuff..Selectmi Idfo he to identify. iso ltc,ttass. prt~f tc,artd apply hiactelaln. hairiness or lotng ,wns %kill mIlct sthilrohleiiis dutilig fingal pathogens to tile soil. Ihe lalattese beetle in North threshing, pa rtictfarl. maitial threshinfl. [hireshers \\ill America has hcci coitrolled hv tite conmnleciall\ a\ailable complain of itcfimin atid it':teascd (fust levels. hactcrial prcpa ration itaci/hl.pu/ pav ofr milksdiscas (71). Attempts itIRRI to fitid tc.istatncc to the rice bug S,'pote, ,pr;tyefott the soil itlot\\ed ilrtier reitiain acti\c '/.plorisa have ilotbeCnl prmitisilg. lIIetIattonlids iorc fr Vea"'s. through the lenilna or palca. hillhairs sed'stVIor seeds wilh Root aphid ,sardtnctly bugs readil\slic tilib to predatots awns that deter bugs promote complaints from threshers. and parlsites once their tetndine ants are controlled. The Seed bugs li'ptocorisa and Stc'nocoris enter the seed larvac and adtilts of a coccinellid beetle '.1Scynm/s sp. prey through the opening between tile lemna and palC, but on /. /llriadlo//uai.s in slash and burn upland rice. Baits varieties with narrow openings are diifficult to mill. treated with insecticide may be used to control tending ants Breeding priorities should concentrate ott ino.tCI gruLtLIps by killing their \ourig (244). that are most difficult to c nttrol by other meains. Steil) Root-tkcditig blugs, false \wire\\oris. w\'ireor\\m.s. and horers appear to be the most widespread group that root \wcc\ils aic nomttllr of,such little ifilpoltatice that the sigiiificantly dalag s rice aind is hard to manage by other role of liatural Cticniics hias tiot beei asscssed. A possibility. Ilnca us. ho\\evrl', tiglt he tit t1g11enItaIIitic niaCIMtodcs. lint, Itese soil-itlhlfbitine mCtitodcs \W5uhtfhave to be imiass Biological control Produiced. Very fe\ stutdic., of mitatt~l eneties of rice pests in upland .\ large miltibcr of niatltrll cnemics ircknown f't envirollnletits wee fouiid in tileliterature. Therefore, we CltI'ori'S. I'C IpblCeii is thtt cltt\vortnlls ,ofonli/e the crop have not compiled a list of species. \Ve present a list of soott alter land ptel);tritiOti a1tithe bcgit iniig of the raittv natural enemies i'Otid in upland ricefields tuic latialln, Citsoln, \hfien tiatllral leniiiy populations aric lo\. Colltloil Batangas, for some conitno i'Oldiar pest species ([itble 5). WOtuld iii\ ol\e mass prodtticing andirclcaslng key spccies. Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in le egg stage is the fpirt (olthe ,Cedlitgt Ial;ot life cycle the lowlatnds or 'roi crops otithcr thlin tice. tI \%hich to f')ctis ftor tittlril elctit cottntrol h.plra;ites o1" Qu(a/iltr o/ 11l//l t'/11't('. Nat trrtfoeits lii ight Ilehc pleditors ttsiteggs ellhighly expoIsYCl. I Sgg pailsites or used agaitist ile social insects tilt and termites., Sollie predators s\tolld probahly ,t\e be ittassproduced and predator specice live ill theirf nests n nlltilnic their ip- released. Littlh i toruialioll is a' ailable ott what specie:s to pearan;IlCe aMRidbeha\vior. IJtrouMdCing a mote aggressise ant trV o ho\ to Iniss p'odLice thiCIi. species to displace pest ant codonies is a possibility (169). A etiloplhid \. I/cmi/hla:,o..l sp.. effctii\cly regu­ Patihogens, as potential natural encnies, have reccived little lates the leatf l'.cl'/oalapo//lm'2:a .spio.anoiiitr heatc and atteition (I13). Natural ncnuics asa group, however, ha\c rice antid should be collsctvcd. not been shown toIlbe iporta it in Iegulating aint and leaf beet.les whCllosehirO e am] eggs ate laid in the grutind ternite numbers. \\ould be motC difficutlt to control by niatulral enelV The nocturnal behavior aid subteitmien habitat of lole Mnll ipuhltiOn,. But species sUici ;ISthispid beetles \\'ithc,_gs crickets and field crickets protect them f'rom many ntllral and larV\le oilleascs ate vtilcratble to iparaites and enemies. There are only limited records of natural cnelics pledators. for these groups of pests: some pathogens (87) but mainly Natural enemisc, of tlirips otl rice are little known. 28 IRPS No. 123, January 1987

Table S. Parasites recorded from upland rice insect pests in Tanauan, Batangas, Philippines, 1977-84. Order Family Species Host stage

Leaffolders Cnaphalocrocis, Marasinia spy. I lymttenoptera a Braconidae Cardiochilesphilipplinensis Ashmead Larva Cotesia angustillasis(Gahan) Larva Cotesia or. cypris (Nixon) Larva Cotesia or. taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Egg to larva Chelonits Spp. Egg to larva Alacroce'trts philippinensis (Ashtmead) Larva Orgilts spp. Larva Iclineumonidac Isclutojppa hireator(Fabricius) Larva Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva a Tenelchha stangli (Ashmead) Larva ir'ichomma cnaphalocrosis Uchida Larva Bethylidac Goniozms or. trianguli ,r Kieffer Larva Elasmidac ''lasmtusalhopictus Crawford Larva Elasmtus spp. Larva Encyrtidac Copidosomopsisnacoh'iae(F:idy) Egg to larva Chalcididac Brach ,nteria e.vcarinata Gahan Pupa Biachyi'erialasus (Walker) Pupa Diptera Tachinidac Zt'gohothria ciliata (Wulp) Larva Argyrophyl.x nigrotihialis(Blaranov) Larva Yellow stern borer Scirpopiaga incertulas l-ymenoptera Braconidac Braco,, chinensisSzeplegeti Larva CIthelotis spp. Larva Chelonus niunakatae Munakata Egg to larva Cotesi, flaripes Cameron a Larva Steno/racon nicerill'i(Bingham) Larva Tropobracon schoenobii (Viereck) Larva Ichneumonidac Amatroinorpiaaccepta metathoracica(Ashmiead) Larva Frihorus sinicus (I lolmgren) Larva Ischnojoppaluteator (Fabricius) Larva Isotima nr. dammeratiti Rohwer Larva a Tenelucha philippinensis (Ashmead) Larva Tenehicha stangli (Ashmcad) Larva Trichomma enaphalocrosis Uchida Lara Xanthopimnpla stemmator (Thunberg) Larva to pupa Charops hrachYpterumn Gupta and Malteswary Larva Chalcididac Brachtm eria spp. Pupa Eulophidac Tetrasticttus schoenohii Ferriere Egg Trichogranmmatidae Trichorigranm chihunis Ishi Egg Tt ichograintnta aponiicuitt (Ashniead) Egg a Scelionidac Telenomits rowani (Gahan) Fgg Pteromalidae Trichoinalopsis apattelctena (Crawford) Egg, Diptera pupa Tachinidae J'eirhaea spp. Larva Z4'gobothria ciliata (Wulp) Larva Gol fringed borer ('hilo auricilius Ilymenoptera lecneumonirlac Trichomm enop/talocrosisUchida Larva a Aanthol'imlh steiniator (Thtinberg) Loarva to pupa Pink ste't I:oret Sesamia Ieioenoptera inferens Braconidae St.,,oIn'acoil nic'rilhi (Bingh am) l.arva Armvtnyorms Mythimna separata. Spodoptera mauritia acronyetoides Diptera Tachinidae Zvgohothriaatropi'ora( Robinea i -Desvoid y)" Larva Ilyinenoptera Eulophidae I:t-phctrtiscitapadoe (Ashimead) Larva (halcididae Brachpin ria spip. Larva Braconidac ('oesia spp. larva Rice I'elop ida math ias aidgreen /iortedci ciliar NIelan itis eda ismtne IHyntenoptera ('halcididae Brat l'hincria sp. iii. matuqitatt Cameron I)iptera Larva Tachinidae ,,Irgyrolhirlaxi "trotlhia/is(marumtv)a1 Larva fuown se tilo per Mois frugalis Diptera Tachnidae Irgt'rophrlax 1igrothiahiv 3a rat ov) Larva Ilymenoptera lelineutionidae ,'anthopimpa pmnctata (I'abricius) Larva to pupa aMost dominant. IRPS No. 123, January 1987 29

Normally, predatory thrips prey on herbivorous thrips. strepsiptera on both. Spiders dominate tilenymphal/adult Natural enemies of stem bugs --Scolinoplara and Tibra'a predators. Spittle bugs move to rice from pasture grasses - are, in order of importance, egg par~isites and predators, and biocontrol would be successful only ifcarried out before fungal pathogens, and nymphal, adult parasites. Egg tiley immigrate into ricefields. parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released. %% ould follow those outlined for stein bugs. Armyworms are normally lwld in :hcck by the activities Coml)arison of" environnewtns. Natural enemies are of egg and larval parasites. When these natural enemies fail, perhaps faced with even greaterconstraints than insect pests usually because of drought. armyworns become epidemic. to survive ind reproduce in upland environments. They It may be worthwhile, therefore. to release parasites during have adapted an,i survived by having wide host ranges, the the rainy months following a drought. Virus diseases of ability to aestivate or hibernate over unfavorable seasons, larvae would beanotheravnue to exploreforarnyworms. and the ability to disperse, or by other mechanisms. Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers, usually by rekasing adapted species of natural enemies collected in various exotic species( 166). This approach has not met with success, environments in the Philippines, we can understand how and new avenues should be explored. The role of predators different rice environments are. arid pathogens is little inderstood and deserves greater Anagqrms olptahiis,a mymarid planthopper egg parasite, attention. Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Table high but should be supplemented with effective egg 6). It is selective for planthoppers and even parasitizes eggs predators. Orthopterans are potential egg predators. of the n-iize planthopper Peregrinitsmaidis itt upland areas. Metioche vittaticollis (Sta ) Ana.vipha /ni.fiwnnis (Saus- A../lavohis also parasitizes maize planthopper eggs but is sure), and Conocephahs hongip'nni. (de i-laan) feed on most prevalent on BPH in the irrigated wetlands. (ona­ stem borer eggs. .etioche and Ana.xil)ha specialize in eggs tocerus spp., like A. optalhilis on BPH and WBPI I,attacks with no hair such as Chilo spp. The omnivorous ('ono- the eggs of GLH in ell rie environments. It is also found cephahsspecializes in eggs cov red with hairs: it is a proven parasitizing the eggs ef the white rice licahopper (o/imna egg predator of Scirlouphaga spp. but has the discouraging spetra. Oirosita naias, a trichogrammatid egg parasite, habit of eating rice guin (65). attacks BIPH and WBPH in the wetlands and has no other Grasshoppers have egg (scelionid) and nymphalad ult host. A related species, 0. aesopi, specializes in G1-H in (nemestrinid, tachinid, and sarcophagid) v irasites. Among wetland environments. Dryinid, strepsipt-'an, and pipun­ pathogens, protozoans have been recorded most frequently. culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas, environments. Among the dryinids, Pseuhdgonatoips*flavi- Leaffolders have rich complexes of natural enemies, some feimur prefers the lowlands and BPH. while P. nuch has no adapted to upland rice (22. 27), ranging from egg predators environmental prefirences but is adapted only to WBPH. (gryllids, coccinellids) to larval parasites (braconids. ichncu- The main d ryinid species on GLH, ;aptogonatop)usspp., is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira! and fungal diseases). Efforts to augment leafhoppers within Amnrasca. Cicadulina,and Ba/chaltua. these natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing. releasing)and viral or fungal pathogens (culturing, lealhoppers. The Elenchus species in the Philippines attack spraying). all species of planthoppers within the genera Sogawilla, Parasitization of the large, conspicuous larvae of Sogatodels, Nilapars'ata,Harualia. and Opiconsiva. El/en­ skippers, green horned caterpillars, and other polyphagous cus.rva.suntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low. ELg but prefers BPH in rainfed arid WBPIH in wetland environ­ parasites and predators are perhaps the key to their control. ments. Ilalictoph(utsmtnroeiand H..Vpectrus attack G LH Gall midges are parasitized by pterormalids and platy- and COfimra.vcctra, respectively. gasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers. Each seasons. Shoot aphids are preyed upon by coccinellids, species has a unique environmental preference. Tomos­ syrphids, and chrysopids. varireIa or':aetora prefers Ne7 otetltix nigropictrs, and Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp., which predators as well as nymphial/ad nlt parasitesand predators* are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments. T suhvirescensattacks Nephotettix virescens pathogens. The principal egg parasites are myrnarids and and prefers the lowlands. Pilpuiulus nttillattls is an trichogramniatids. Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix species. bugs that suck out the yolk. Nymphal; ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers, pipunculids on lealhopper's, and the Philippines attack species of C(naplwocroci.s and 30 IRPS No. 123, January 1987

Table 6. Dominant natural enemies of rice pests in upland, rainfed lowland, and irrigated lowland rice environments of the Philippines.

Natural enemy Environment Host stage Species Upand Rainfed lowland Irrigated lowland

Brown planthop.- * A'ilapar'ata lugens Parasite Egg Mymaridae Anagrus optabilis (Perkins) x x X Anagrusflaveolus Waterhouse x Trichogrammatidac Oligosita naiasGirault x Nymph/Adult Dryinidac Pseudogotatopus laiifemttr Esaki et Hashimoto x x Elenchidae Elenchus yasurnatsui Kifune and Hirashima x X X Whitebacked planthoppers Sogatella furcifera Egg and Sogatodes pusanus Mymaridae Anagrus optabilis (Perkins) x x x Trichogrammatidac Ol~gositanaias Girault x x Nymph/Adult Dryinidae Pseudogonatopusnudus Perkins X x X Elenchidae Elenchus yasutatsui Kifune et Hirashima x x Green leafhopper Nephotettix virescens Parasite Egg Mymaridae Gonatocerus spp. X x x Trichogrammatidae Oligosita aesopi Girault x x Nymph/Adult Pipunculidac Tomosvaryella subviresccns Loew X X Tomosvaryella oryzaetora (Koizumi) X X Pipunclusmutillatus(de Meijere) X Dryinidae Ilaplogonatopusspp. X Rice leaffolders Cnaphalocrocis,Marasmia Egg Copidosomopsis nacoleiae (Eady) x X x Larva BraLonidae Cardiochiles philipphnensisAshnead X lchneumonidac Trichiomnia cnaphalocrosisUchida X Temelucha stangli (Ashmead) X Pupa Chalcididae Brachymeria excarinataGahan x x x Brachymerialasus (Walker) x x Brachymeria X nr. tachardiaegroup x x x Yellow stem borer Scirpophaga incertulas Parasite Egg Scelionidae Telenomus rowani (Gahan) X X x Eulophidac Tettasticus schoenobhi Ferriere x Larva Braconidae Stenobraconnicevillei (Bingham) x Iclhncurnonidac Temelucha phiiippinensis(Ashinead) x Temelucha stangli (Ashmead) X Ainauromorpha X accepta metathoracica(Ashmead) x continued on next page IRPS No. 123, January 1987 31

Table 6 cntinued.

Natural enemy Environment Host stage Species Upland Rainfed lowland Irrigated lowland Larva-pupa Gold fringed borer Chilo attrici/ius lchneumonidae Xanthopimpla stemmator (Thunberg) x a Pupa x Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax Pink stem borerb Sesamnia inferens Parasite Larva Braconidae Stenobracon nicevillei (Bingham) x b b Larva Armyworms Mvthinna, Spodoptera Tachinidac Zygobothria arroptvora (Robineau-Desvoidy) x x Braconidac Cotesia spp. X F-.gg Skipper Pelopidas mathias and green horned caterpillar Melanitisleda ismene Trichogramrnatidac Trichogrannaspp. x Scelionidae Undetermined Scelionid x Larva Tachinidae Argy'roph /lax nitrotibialis (Baranov) X x Pteromalidae Trichomnalopsis apanteloctena (Crawford) X Predator Coenagrionidae Agriocnenis femina femina Brauer x x Ischnura senegalensis 'tambur Miridae CrtorhinsliidipennisReuter x x Nabidac Stcnonabis tagalicus (Stal) x Veliidae Alicrovelia douglasi atrolineata (Bergroth) x Coccinellidae Micraspis crocea (Mulsant) X x x Vespidae Eumenes campaniformis (Fabricius) x Formicidac Diaecamna spp. x Odontoponera transversa (F. Smith) x Carabidae Ch/aenius spp. x Ophionea W.- aJasciataSchmidt-Goebel x Ophioneaishii ishii Habu x X Gryllidav Anaxipta longipennis (Saussur-) x x Metioche ittaticollis (Sial) x x x Tettigoniidae Conocephalus longipennis (de Haan) x x x a No rearing made on rainfed lowland. bNo rearing made on rainfed lowland or Irrigated lowland.

Mr'asmia.Among the Laval parasites, the braconid Cardio- lowlands and parasitizes oriental maize borer OsIrinia chilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae. The leaffolder pupal parasites ment. It also parasitizes Hydelepta indicata, a common not only occur equally in all environments but also have leaffolder of legumes. The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia, Maruca, phalocrosis and Temehcha stangli occur more in the Hydelepta, and Honiona- in maize and legumes. lowlands. T. stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB) and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environments larvae. T. cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25). 32 IRPS No. 123, January 1987

Tetrastictu schoenobiispecializes only in YSB and occurs However, of the 10 most prevalent ricefield spiders. 9 were mainly in irrigated lowlands -- further proof that YSB is abundant in upland rice, showing wide adaptation. All three most adapted to the wetter environments, spider guilds -- orb-web, space-web., and huting spiders ­ Larval parasites of YSB are highly environment-specific. were prevalent in upland ricefields. There were, however, The braconid Stenohracon n'icevillei occurs only in upland differences in environmental prelerences for some spider rice but also attacks the pink stem borer (PSB). Tiei'nchwa species. phii'iplP'wnsis, an ichneumonid. occurs in upland rice and Of the orb-wcavers, T'etragniathamanlihulatawas parti­ also parasitizes the larvae of (7hi/u stqpre.sais.,inatro- cila rlv abundant, 7. japonica was low, and Lt'ucatge morpha accepta melaihoracica, like 7. stang/i, is mnost dehcorata was absent in upland rice. adapted to the irrigated lowlands. As in other environments, Aiypeiwnaifirtosana was the The dark-headed stem borer(l)HSB) (ih po!.chit3sus most dominant space-web spider, however, its relative was not collected in all rice environments, and the gold numbers were lowerthan for lowland sites. Among hunting fringed borer Chilo auricilius nd PSB were collected only spiders, O.\x|'opes javanus and L'cosa leatcost fgla were in upland rice. Xaantiholiplastenrnalor,an ichneumonid more abundant and Lvcosa lpseudoanmnlata was less larval parasite, appears to be widely adapted, whereas abundant in upland compared with lowland rice envi­ Tri'homaslopisapanitI)cteiia,a pteromalid pupal parasite, roniments. occurs mostly in wetlands and also parasitizes larvae of Extensive rearing of rice insect pests in upland, rainfed Chilo suppr'essais and Pelhuida.s mathias, lowland, and irrigated sites over an 8-yr period in the Armyworms Myit hinin and Spodujipera have two main Philippines provided insight into the effectiveness of larval parasites - a tachinid Zrgohothriaatropivora in parasites as natural enemies (Table 7, 8). Most of toe data rainfed environments and a braconid Cotesia spp. in are from large sample sizes taken over at least one crop, irrigated lowlands, which overcomes the pitfalls expressed by Van Driesche The butterfly pests Pelopidas and .4elanitishave tricho- (238). grammatid egg parasites found only in irrigated lowland VSB egg parasitization was surprisingly similar across all environments, environments. Larval parasitization was highly dynamic Two larval parasites show environmental preferences -- among sites and even years within the same site but showed Argrroph '/a.v nigrotihialis, a tachinid in rainfed habitats, somewhat lowerlevels in uplandsthan in irrigated wetlands. and Trichonialopsisapantelkcnela in irrigated ricefields. A. This would have been predicted for egg parasites as well, limgrotibialisalsoattacks the sweet potato hornwormAgrius since YSB is adapted to the lowlands and so must be its convolvtdi, and T. apanteloctena parasitizes DH S1B. parasites. This isevidence that Telenomus rowani is as good Predators as a group show more distinct environmental at dispersing as its main hosts. T.rowani clings onto the anal preferences. The aquatic damselflies Coenagrionidae are tufts of female YSB and parasitizes eggs as they are laid. most abundant in the lowlands. Cvriorhinus is also most This phoresy may explain its wide adaptation. The more adapted to the lowlands. The nabid Stenonabis tagalicus specialized lowland YSB larval parasites, therefore, appear occurs only in the uplands and isalso prevalent on legumes. more effective than their upland counterparts with more The coccinellid lady beetles, wasps, and ants are more alternate hosts. prevalent in upland areas. AlicrasIpis cit-ea larvae prey ona Low collection levels of PSB larvae, especially in the variety of aphids attacking legumes and maize. Etunienes wetlands, make comparisons between environments campan./oni.,a vespid mud wasp, makes nests in trees and difficult. Low levels of larval parasifization were recorded in is therefore most adapted to the more botanically diverse both rainfed environments. Except in the Cagayan rainfed uplands. Soil-dwelling ants cannot tolerate flooding, wetland site in 1980, low levels ef larval parasitization were Upland environments are habitats to arboreal carabid evident at irrigated and rainfed sitesalike. No differences by beetles. Three species of Chlaeniusbeetles prey on leaffolder environment were evident. Higher levels of parasitization larvae and are perhaps more important than parasites (27). occurred in Sarawak during a 1967 outbreak, also from The gryllids and tettigoniids are widely adapted egg tachinids. Rothschild (197) indicated that higher parasi­ predators, not only on rice but also on maize and legumes. tization occurred in upland rice areas. They feed on eggs of most insects that are laid on leaves. Skipper and green horned caterpillar larval parasitization The spidercommunity of upland rice environments isrich rates were generally low and showed no environmental in species. At one site (Batangas, Philippines) 31 species effect. htighest activity occurred in Cagayan in 1980 (48% have been recorded (23). Of the 176 spider species recorded parasitization). Although larval parasitization levels of RLF in Philippine ricefields, about one-half (82) occur in upland were variable year to year, activity appeared greater in the rice. The spiderspecies of upland rice environments overlap irrigated lowlands than in the uplands. Again, the highest more with those of rainfed lowland than with those of" levels were recorded in Cagayan in 1980. Cagayan has irrigated lowland rice (24). The same study showed that of extensive grasslands and fallow around the rice areas and a the three environments, irrigated rice has (ie greatest spider relatively long rainfall period. Rainfall in 1930 was good for species diversity, foiiowed by rainfed lowland and upland. crop growth and apparently for increase of larval parasites. IRPS No. 123, January 1987 33

Table 7. Parasitization rates of rice planthoppers and green leafhopper in upland, rainfed lowland, and irrigated lowland environments. Philip­ pines, 1976-84.0

Egg parasitization (%) Nymphal/adult parasitization (%) Environment and site Year BPH W13P GLH BPII WBPH GLIH Upland Batangas 1976 2(47) 9(1800) 8 (101) 1979 8(39) 14 (212) 6 (127) 1980 15 17 12 1982 4(51) 12(304) 9(116) Rainfed lowland Iloilo 1976 0 (69) - 0 (471) 0 (200) - 4 (600) 1977 9 (214) 7 (462) 10 (725) 1978 11 - 11 10 (1259) 11 (270) 10 (362) 1981 8(235) 3 (313) 12(870) Cagayan 1980 44 46 52 6 (900) 5 (900) 5 (900) 1981 48 55 56 3(800) 1 (800) 3(800) 1982 40 44 49 1984 6(14) 3 (81) 28(211) Pangasinan 1976 7 (197) 10 (274) 9 (764) 1978 13 (436) 13(379) 14 (457) 17(198) 3 (116) 5 (647) 1979 11 (486) 16(871) 7 (1204) 1982 18 (275) 19(88) 8(305) Irrigatedlowland Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 56 26 16 1983 7(278) 16 (1650) 27(1921) aSources of data: All sites: A. T. Barrion, unpublished, 1977-82; Batangas: R. I. Apostol, unpublished; 1lo1lo and Pangasinan: 136; Cagayan: B. Canapi, unpublished; Li'una: 175; I'. C. Pantua, unpublished; 118; 234; 43; Carino and Shepard, unpublished. BPH = brown planthopper Nila. parvara htgens, W131Pi = whitebacked planthopper Sogatelia furcifera, GLH = green leafhopper Nephotettrix virescens. Figiires in parentheses are sample sizes.

Tauher et al (230) reported that internal, external, and These beneficial organisms must be conserved by applying genetic factors influence the seasonal activity of parasitoids. insecticides judiciously, particularly sprays. Seed or soil Hopper egg parasites, but not nymphal/adult parasites, placement of chemicals minimizes exposure to natural were also parlicularly abundant at the Cagayan site: enemies. The strategy to derive the greatest benefit from possibly the egg parasites have more alternative hopper prey natural enemies isto allow the greatest number of beneficial in the weedy areas than nymphal.:adult paisites. species to thrive. Then, perhaps at least one species will be The seasonal dynamics of hopper egg parasites can be effective against each pest at any tine, overcoming the seen for upland rice in Batangas(Fig. I. 2). and for rainfed variable abundance of each species within and between lowland lice in Pangasinan( 135)and Cagayan( 105). Levels years. of BPH,WBPI-L. and GLH egg parasitization were corn- Programs to introduce exotic species or mass produce parable in upland and irrigated lowland environments. indigenous ones are ambitious and expensive because they Except for some high rates of paraMsitization early in the require trained people on a sustained, not ad hoc, basis. season, the parasitization rate tended to be steady in all envilonments. However, hopper nymphal/adult parasi- Chemical control tizution levels were more similar in the rainfed environments Insecticides offer rapid and efficient control of upland insect and lower than in irrigated rice. pests (190) but-should be used only after other control Overall, parasites do not appear highly effective by measures have been considered. Insecticides are rarely used themselves against upland or even lowland pests. Among on upland rice because of cost (32). Upland rice yields are natural enemies, parasites have attracted the most attention normally too low to justify the expenditure. Also, most because they can be more readily assesscd, upland rice is grown as a cash crop. Spraying upland rice is Predators and pathogens lend themselves better to more difficult than lowland rice because water is less management practices. Predators, the most important accessible. Government subsidy programs to provide pesti­ group of natural enemies in riccfields, are difficult to cides to tribal peoples for upland rice production have quantify. On the other hand, pathogens appear less im- created problems of toxicity to humans because isolated portant but can be readily cultured and disseminated, people have had no experience to be able to handle The natural enemy commulnity of upland rice is rich in pesticides safely. species and differs significantly from that of lowland rice. However, ins-eticide use has beenjustified when a crop is 0Z

Table 8. Parasitization rates of lepidopterous pests in upland, rainfed lowland, and irrigated lowland environments. Philippines, 1976-84.a

Parasitization (5) Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooper Egg Larva larva larva larva larva larva Upland Batangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101) 1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30) 1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26) Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2? 13) 15 (180) 1978 5 (76) 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191) 1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104) 1981 56 (1322) 18(304) 0(5) 12(51) 18(138) Pangasinan 1976 60 ( )b 7 7 62(82) 0(111) 5 (39) 1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7) 0(11) 4 (28) 15 (286) 1980 62 (' 48(177) 61(140) 1981 0(5) 12 (2.) 13(45) 14 (92) Irrigatedlowland Laguna 1974 62 8 1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60) 1979 83(2017) 18(110) 8(62) 1980 11 (89) 22(400) 78 (1625) 20(75) 0 (16) 4 (72) 9 (80) 1983 60 7 (270) 30 aSources of data: All sites: A. T. Barrion, unpublished, 1977-82; Batangas: R. F. Apostol, unpublished; Iloilo and Pangasnan: 136; Cagayan: B. Canapi, unpublished; Laguna; 175; P. Pantua, unpublished; 118; 234; 143; C. Carino and Shepard, unpublished. YSB = yellow stem borer Scirpophagaincertulas,GFB = gold fringed borer Chilo Sesamia inferens, earcutting auricilius PSB = pink stem borer caterpillar Alythimna separata, swarming caterpillar Spcdoptera mauritia acronyctoides, skipper Pelopidas mathias, ismene, RLF = rice leaffoldersCnaphalocrocis reen horned caterpillar Melanitls leda medinalis and Marasmiaspp., brown semilooper Mocisfrugalis Figures in parentheses are sample sizes. Egg masses. IRPS No. 123, January 1987 35

Table 9. Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often are yield of Dagge upland rice. Batangas, Philippines, 1976.a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o), but usually from different pests. The principal (kg ai/ha) seed treatment (kg ai/ha) (no.Ira-row) (t/ha) groups are soil pests feeding on sown seed and roots, at 14 DE followed by seedling pests. These groups have largely gone 50 None - 35 a 2.0 a unnoticed, leading researchers to conclude that stem borers 50 Carbofuran F 1.0 38 ab 2.3 a and seed bugs are the most important pests (17). Yield loss 100 None 1.0- 4551 beCabofuranFc 2.3 a trials that pinpoint major pests need to includc seed 100 Dieldrin WP 1.0 52 cd 2.2 a treatments. Granules applied in the furrow may not give 150 None - 59 d 2.2 a adequate protection from many seed pests. aAv of 8 fields. DE = days after crop emergence. In a column, means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of , 5%level by D)MRT. F = flowable, Wi' = wettable powder, both pests and their natural enemie:s. Irrigation has homo­ genized lowland rice environments. Flooded soils tend to threatened by epidemics of locusts or armyworms. New have the same properties and to eliminate soil pests. The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost. benefited specialist pests. Seed treatment is normally inexpensive and can be Upland rice has no specialist species. The economicallyjustified in -nanycases(66). Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved. Rice in the uplands exists only because (247) is most effective in controlling seedling maggots. of man, and even though lowland rice predates upland rice, Foliar sprays have to be repeated to obtain control and specialist species evolved. Wheat, maize, and sorghum do often are washed off by frequent early season rains, not have monophagous pests, leading us to conclude that Granular insecticides are effective, but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae. (247). When directed against first-instar larvae, banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cycles, grubs at low dosages (0.25 kg aii ha) (134). Seed treatments undergoing dormancy, or being dispersive. Each pest has a are not promising for white grub control (248). unique life history strategy suited to highly unpredictable Seed treatment is also effective a.inst ants and may be environments. In agivenyear, someinsect specics an better cheaper than increasing the seeding rate. In Batangas, exploit upland rice as a food source, but no pest is a senious treating 50 kg seed/ ha was as effective as increasing the one every year on upland rice. seeding rate to 100 kg/ha and using untreated seeds (Table The low yielding potential of upland rice means less 9). In Brazil, seed treatments control termites and other soil quantity and lower quality of food for pests to exploit, but insects including the lessercornstalk borer(66). Domiciano also less incentive for farmers to undertake control. (58), however. found seed treatments inconsistent in their The most effective control would be early and synchro­ effect against high populations of lesser corn stalk borer. nous planting of early-maturing upland rice varieties to Baiting can be an inexpensive way of controlling ants, prevent insect pest buildup. Breeding for insect resistance seedling maggot flies, mole crickets, and field crickets. Baits should initially focus on stem borers because they are very made of locally available material and impregnated with difficult to control by other means. Biocontrol efforts insecticide can be sparingly distributed, taking ad'antage of should first stress conservation of natural enemies, pre­ the pests' ability to disperse and encounter bait sites. dators, and pathogens rather than parasites. Efficient The new low-volume, hand-held, controlled-droplet chemical control includes seed treatment or baiting for seed, sprayers may offer an advantage over high-volume knap- seedling, and some root pests, and using controlled droplet sack sprayes for controlling foliar pests. Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice, and bugs when economic thresholds are reached. better droplet penetration should result. Controlled droplet applicators would be ideal for seedbugs and defoliators.

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice, mainly beca use of the addition of soil pests. Generally, population levels of lowland rice pests are lower in upland than lowland rice, leading many people to conclude that insect pests are not important on upland rice. 36 IRPS No. 123, January 1987

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