Invasive Species Compendium

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Datasheet

Spodoptera litura
(taro caterpillar)

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Datasheet

Spodoptera litura (taro caterpillar)

Summary

  • Last modified
  • 11 December 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Spodoptera litura
  • Preferred Common Name
  • taro caterpillar
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • The tobacco caterpillar,S. litura, is one of the most important insect pests of agricultural crops in the Asian tropics. It is widely distributed throughout tropical and temperate Asia, Australasia and the Pacific Islands (...

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Pictures

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PictureTitleCaptionCopyright
Spodoptera litura (taro caterpillar); forewings grey to reddish-brown with a strongly variegated pattern and paler lines along the veins.
TitleAdult
CaptionSpodoptera litura (taro caterpillar); forewings grey to reddish-brown with a strongly variegated pattern and paler lines along the veins.
Copyright©E. Neering
Spodoptera litura (taro caterpillar); forewings grey to reddish-brown with a strongly variegated pattern and paler lines along the veins.
AdultSpodoptera litura (taro caterpillar); forewings grey to reddish-brown with a strongly variegated pattern and paler lines along the veins.©E. Neering
Spodoptera litura (taro caterpillar); larva hairless, variable in colour (young larvae light green, the later instars dark green to brown), sides of body with dark and light longitudinal bands.
TitleMature larva
CaptionSpodoptera litura (taro caterpillar); larva hairless, variable in colour (young larvae light green, the later instars dark green to brown), sides of body with dark and light longitudinal bands.
Copyright©E. Neering
Spodoptera litura (taro caterpillar); larva hairless, variable in colour (young larvae light green, the later instars dark green to brown), sides of body with dark and light longitudinal bands.
Mature larvaSpodoptera litura (taro caterpillar); larva hairless, variable in colour (young larvae light green, the later instars dark green to brown), sides of body with dark and light longitudinal bands.©E. Neering
Spodoptera litura (taro caterpillar); s. litura egg mass on soyabean (Note Hymenopteran egg parasites)
TitleEgg mass
CaptionSpodoptera litura (taro caterpillar); s. litura egg mass on soyabean (Note Hymenopteran egg parasites)
Copyright©Ernst Neering
Spodoptera litura (taro caterpillar); s. litura egg mass on soyabean (Note Hymenopteran egg parasites)
Egg massSpodoptera litura (taro caterpillar); s. litura egg mass on soyabean (Note Hymenopteran egg parasites)©Ernst Neering
Spodoptera litura (taro caterpillar); egg masses measure 4-7 mm in diameter and appear golden brown because they are covered with body scales of females.
TitleEgg masses
CaptionSpodoptera litura (taro caterpillar); egg masses measure 4-7 mm in diameter and appear golden brown because they are covered with body scales of females.
Copyright©ICRISAT
Spodoptera litura (taro caterpillar); egg masses measure 4-7 mm in diameter and appear golden brown because they are covered with body scales of females.
Egg massesSpodoptera litura (taro caterpillar); egg masses measure 4-7 mm in diameter and appear golden brown because they are covered with body scales of females. ©ICRISAT

Identity

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Preferred Scientific Name

  • Spodoptera litura (Fabricius)

Preferred Common Name

  • taro caterpillar

Other Scientific Names

  • Mamestra albisparsa Walker
  • Noctua elata Fabricius
  • Noctua histrionica Fabricius
  • Noctua litura Fabricius
  • Prodenia ciligera Guenée
  • Prodenia declinata Walker
  • Prodenia evanescens Butler
  • Prodenia glaucistriga Walker
  • Prodenia littoralis Fabricius
  • Prodenia litura Fabricius
  • Prodenia subterminalis Walker
  • Prodenia tasmanica Guenée
  • Prodenia testaceoides Walker
  • Spodoptera littoralis

International Common Names

  • English: armyworm; cluster caterpillar; common cutworm; cotton leafworm; cotton worm; Egyptian cotton leafworm; rice cutworm; tobacco budworm; tobacco caterpillar; tobacco cutworm; tobacco leaf caterpillar
  • Spanish: gusano del tabaco; gusano negro; lagarta; rosquilla negra
  • French: chenille defoliante; noctuelle rayee; ver du coton; ver du tabac

Local Common Names

  • Germany: Aegyptische Baumwollraupe; Asiatischer Baumwollwurm; Baumwollblattraupe; Baumwollraupe, Aegyptische; Baumwollwurm, Asiatischer
  • India: ladde purugu; telugu
  • Israel: haprodenia
  • Italy: larva bruna con macchie vellutate
  • Japan: hasumon-yoto
  • Netherlands: eiernestrups
  • Turkey: pamuk yaprak kurdu

EPPO code

  • PRODLI (Spodoptera litura)

Summary of Invasiveness

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The tobacco caterpillar,S. litura, is one of the most important insect pests of agricultural crops in the Asian tropics. It is widely distributed throughout tropical and temperate Asia, Australasia and the Pacific Islands (Feakin, 1973; Kranz et al., 1977). Records of S. litura having limited distribution in (or being eradicated from) Germany, Russian Federation, Russian Far East, the UK and Réunion may in fact refer to S. littoralis. Both S. litura and S. littoralis are totally polyphagous (Brown and Dewhurst, 1975; Holloway, 1989) and therefore have huge potential to invade new areas and/or to adapt to new climatic and/or ecological situations. The Spodoptera group consists of closely related species with similar ecology that are difficult to identify to species level.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Lepidoptera
  •                         Family: Noctuidae
  •                             Genus: Spodoptera
  •                                 Species: Spodoptera litura

Notes on Taxonomy and Nomenclature

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The two Old World cotton leafworm species, Spodoptera litura and S. littoralis, are allopatric, their ranges covering Asia and Africa, Europe and the Middle East, respectively. Many authors have regarded them as the same species.

Description

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Eggs

The eggs are spherical, somewhat flattened, and 0.6 mm in diameter. They are usually pale orange-brown or pink in colour, laid in batches and covered with hair scales from the tip of the abdomen of the female moth. Egg masses measure about 4-7 mm in diameter and appear golden brown because they are covered with body scales of females.

Larva

The larva is hairless, variable in colour (young larvae are light green, the later instars are dark green to brown on their backs, lighter underneath); sides of body with dark and light longitudinal bands; dorsal side with two dark semilunar spots laterally on each segment, except for the prothorax; spots on the first and eighth abdominal segments larger than others, interrupting the lateral lines on the first segment. Though the markings are variable, a bright-yellow stripe along the length of the dorsal surface is characteristic of S. litura larvae.

Larval instars can be distinguished on the basis of head capsule width, ranging from 2.7 to 25 mm. Body length ranges from 2.3 to 32 mm.

Pupa

The pupa is 15-20 mm long, red-brown; tip of abdomen with two small spines.

Adult

Moth, with grey-brown body, 15-20 mm long; wingspan 30-38 mm. The forewings are grey to reddish-brown with a strongly variegated pattern and paler lines along the veins (in males, bluish areas occur on the wing base and tip); the hindwings are greyish-white with grey margins, often with dark veins in S. litura (but without in S. littoralis). See also Schmutterer (1969), Cayrol (1972) and Brown and Dewhurst (1975).

Distribution

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The tobacco caterpillar, S. litura, is one of the most important insect pests of agricultural crops in the Asian tropics. It is widely distributed throughout tropical and temperate Asia, Australasia and the Pacific Islands (Feaking, 1973; Kranz et al., 1977). Records of S. litura having limited distribution in (or being eradicated from) Germany, Russian Federation, Russian Far East, the UK and Réunion may in fact refer to S. littoralis.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 30 Jun 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

Central African RepublicPresent
GhanaPresent
RéunionPresent
Saint HelenaPresent

Asia

AfghanistanPresentInvasive
BangladeshPresent, WidespreadInvasive
British Indian Ocean Territory
-Chagos ArchipelagoPresent
BruneiPresentInvasive
CambodiaPresentInvasive
ChinaPresent, LocalizedInvasive
-AnhuiPresent
-FujianPresent
-GuangdongPresentInvasive
-GuangxiPresentInvasive
-GuizhouPresentInvasive
-HainanPresent
-HebeiPresent
-HenanPresent
-HubeiPresentInvasive
-HunanPresentInvasive
-Inner MongoliaPresent
-JiangsuPresentInvasive
-JiangxiPresent
-JilinPresentInvasive
-ShandongPresentInvasive
-ShanghaiPresent
-SichuanPresent
-YunnanPresent
-ZhejiangPresentInvasive
Cocos IslandsPresentInvasive
Hong KongPresent, Widespread
IndiaPresent, WidespreadInvasive
-Andaman and Nicobar IslandsPresentInvasive
-Andhra PradeshPresentInvasive
-AssamPresentInvasive
-BiharPresentInvasive
-DelhiPresent
-GujaratPresentInvasive
-HaryanaPresentInvasive
-Himachal PradeshPresentInvasive
-Jammu and KashmirPresentInvasive
-JharkhandPresent
-KarnatakaPresentInvasive
-KeralaPresentInvasive
-Madhya PradeshPresentInvasive
-MaharashtraPresentInvasive
-ManipurPresent
-MeghalayaPresent
-OdishaPresentInvasive
-PunjabPresentInvasive
-RajasthanPresentInvasive
-SikkimPresentInvasive
-Tamil NaduPresentInvasive
-Uttar PradeshPresentInvasive
-UttarakhandPresent
-West BengalPresentInvasive
IndonesiaPresent
-Irian JayaPresent
-JavaPresent
-Maluku IslandsPresent
-SulawesiPresent
-SumatraPresent
IranPresent
IraqPresent
JapanPresent, Widespread
-Bonin IslandsPresent
-HokkaidoPresent, Widespread
-HonshuPresent, Widespread
-KyushuPresent, Widespread
-Ryukyu IslandsPresent
-ShikokuPresent, Widespread
LaosPresent, WidespreadInvasive
MacauPresent
MalaysiaPresent, Widespread
-Peninsular MalaysiaPresent, Widespread
-SabahPresent
-SarawakPresent
MaldivesPresent
MyanmarPresentInvasive
NepalPresentInvasive
North KoreaPresent
OmanPresentInvasive
PakistanPresentInvasive
PhilippinesPresentInvasive
SingaporePresentInvasive
South KoreaPresentInvasive
Sri LankaPresentInvasive
TaiwanPresent, WidespreadInvasive
ThailandPresentInvasive
VietnamPresentInvasive

Europe

DenmarkAbsent, Intercepted onlyOne single glasshouse nursery on the island of Funen.
FrancePresent
GermanyAbsent, Eradicated
NetherlandsAbsent, Eradicated2008Absent, pest eradicated (2008), confirmed by survey. Based on long-term annual surveys, 362 survey observations in 2012.
PortugalPresentPresent based on regional distribution.
-AzoresPresent
RussiaPresent, Localized
-Central RussiaPresent, Few occurrences
-Russian Far EastPresent, Few occurrences
-Southern RussiaPresent, Localized
-Western SiberiaPresent, Few occurrences
SloveniaAbsent
United KingdomAbsent, Eradicated
-EnglandAbsent, Eradicated

North America

United StatesPresent, Localized
-FloridaAbsent, Formerly present
-HawaiiPresent

Oceania

American SamoaPresentInvasive
AustraliaPresent, LocalizedInvasive
-New South WalesPresent, LocalizedInvasive
-Northern TerritoryPresent
-QueenslandPresent, LocalizedInvasive
-VictoriaAbsent, Confirmed absent by survey
-Western AustraliaPresent, LocalizedInvasive
Christmas IslandPresentInvasive
Cook IslandsPresent
Federated States of MicronesiaPresent
-YapPresent
FijiPresentInvasive
French PolynesiaPresent, LocalizedInvasive
-Marquesas IslandsPresent
GuamPresentInvasive
KiribatiPresentInvasive
Marshall IslandsPresentInvasive
New CaledoniaPresentInvasive
New ZealandPresent, LocalizedInvasive
-Kermadec IslandsPresent
NiuePresentInvasive
Norfolk IslandPresentInvasive
Northern Mariana IslandsPresentInvasive
PalauPresent
Papua New GuineaPresentInvasive
PitcairnPresent
SamoaPresentInvasive
Solomon IslandsPresentInvasive
TokelauPresent
TongaPresentInvasive
TuvaluPresentInvasive
VanuatuPresentInvasive
Wallis and FutunaPresent, LocalizedInvasive

Risk of Introduction

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S. litura and S. littoralis are listed as of quarantine significance by EPPO, CPPC, NAPPO and OIRSA.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial ManagedCultivated / agricultural land Principal habitat Harmful (pest or invasive)
Terrestrial ManagedProtected agriculture (e.g. glasshouse production) Principal habitat Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Natural
Terrestrial Natural / Semi-naturalNatural grasslands Principal habitat Natural
Terrestrial Natural / Semi-naturalRiverbanks Secondary/tolerated habitat Natural
Terrestrial Natural / Semi-naturalWetlands Secondary/tolerated habitat Natural

Hosts/Species Affected

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The host range of S. litura covers at least 120 species. Among the main crop species attacked by S. litura in the tropics are Colocasia esculenta, cotton, flax, groundnuts, jute, lucerne, maize, rice, soyabeans, tea, tobacco, vegetables (aubergines, Brassica, Capsicum, cucurbit vegetables, Phaseolus, potatoes, sweet potatoes and species of Vigna). Other hosts include ornamentals, wild plants, weeds and shade trees (for example, Leucaena leucocephala, the shade tree of cocoa plantations in Indonesia).

Both S. litura and S. littoralis are totally polyphagous (Brown and Dewhurst, 1975; Holloway, 1989).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Abelmoschus esculentus (okra)MalvaceaeMain
Acacia mangium (brown salwood)FabaceaeMain
Agastache foeniculumLamiaceaeUnknown
Allium cepa (onion)LiliaceaeMain
Allium sativum (garlic)LiliaceaeUnknown
Amaranthus (amaranth)AmaranthaceaeMain
    Amaranthus blitum (livid amaranth)AmaranthaceaeOther
    Angelica keiskeiApiaceaeUnknown
    Annona squamosa (sugar apple)AnnonaceaeOther
      ApiosUnknown
      ArachisFabaceaeUnknown
      Arachis hypogaea (groundnut)FabaceaeMain
      Asparagus officinalis (asparagus)LiliaceaeUnknown
      Averrhoa carambola (carambola)OxalidaceaeUnknown
      Bacopa monnieri (water hyssop)ScrophulariaceaeUnknown
      BegoniaBegoniaceaeMain
        Beta vulgaris var. saccharifera (sugarbeet)ChenopodiaceaeMain
          Boehmeria nivea (ramie)UrticaceaeMain
          BrassicaBrassicaceaeMain
            Brassica juncea (mustard)BrassicaceaeOther
            Brassica oleracea (cabbages, cauliflowers)BrassicaceaeUnknown
            Brassica oleracea var. botrytis (cauliflower)BrassicaceaeMain
            Brassica oleracea var. capitata (cabbage)BrassicaceaeMain
            Brassica rapa (field mustard)BrassicaceaeUnknown
            Brassica rapa cultivar group CaixinBrassicaceaeMain
              Brassica rapa subsp. chinensis (Chinese cabbage)BrassicaceaeUnknown
              Brassica rapa subsp. pekinensisBrassicaceaeUnknown
              Callistephus chinensis (China aster)AsteraceaeUnknown
                Camellia sinensis (tea)TheaceaeMain
                  Capsicum annuum (bell pepper)SolanaceaeOther
                  Capsicum frutescens (chilli)SolanaceaeMain
                    Carica papaya (pawpaw)CaricaceaeUnknown
                    Chenopodiastrum murale (nettleleaf goosefoot)ChenopodiaceaeOther
                    Chenopodium album (fat hen)ChenopodiaceaeOther
                    Chrysanthemum (daisy)AsteraceaeOther
                      Cicer arietinum (chickpea)FabaceaeMain
                      CitrusRutaceaeMain
                        Coffea (coffee)RubiaceaeMain
                          Coleus blumei (common coleus)LamiaceaeUnknown
                          Colocasia esculenta (taro)AraceaeMain
                          Convolvulus arvensis (bindweed)ConvolvulaceaeOther
                          Corchorus (jutes)TiliaceaeMain
                            Corchorus olitorius (jute)TiliaceaeMain
                            Coriandrum sativum (coriander)ApiaceaeMain
                            Crotalaria juncea (sunn hemp)FabaceaeMain
                              CynaraAsteraceaeMain
                                Cynara cardunculus var. scolymus (globe artichoke)AsteraceaeMain
                                  Dahlia coccineaAsteraceaeOther
                                  Daucus carota (carrot)ApiaceaeOther
                                  DendrobiumOrchidaceaeUnknown
                                  Dicliptera chinensisUnknown
                                  Dimocarpus longan (longan tree)SapindaceaeUnknown
                                  Diospyros kaki (persimmon)EbenaceaeUnknown
                                  Eucalyptus globulus (Tasmanian blue gum)MyrtaceaeUnknown
                                  Fabaceae (leguminous plants)FabaceaeMain
                                    Foeniculum vulgare (fennel)ApiaceaeMain
                                      Fragaria ananassa (strawberry)RosaceaeMain
                                        Gaillardia pulchella (Indian blanket)AsteraceaeUnknown
                                        Gerbera (Barbeton daisy)AsteraceaeOther
                                        Ginkgo biloba (kew tree)GinkgoaceaeUnknown
                                        Gladiolus hybrids (sword lily)IridaceaeMain
                                          Glycine max (soyabean)FabaceaeMain
                                            Gossypium (cotton)MalvaceaeMain
                                              Gossypium hirsutum (Bourbon cotton)MalvaceaeMain
                                              Helianthus annuus (sunflower)AsteraceaeMain
                                              Heliotropium indicum (Indian heliotrope)BoraginaceaeUnknown
                                              Hevea brasiliensis (rubber)EuphorbiaceaeMain
                                                Hibiscus cannabinus (kenaf)MalvaceaeUnknown
                                                Hibiscus rosa-sinensis (China-rose)MalvaceaeOther
                                                Ipomoea aquatica (swamp morning-glory)ConvolvulaceaeMain
                                                  Ipomoea batatas (sweet potato)ConvolvulaceaeMain
                                                    Jatropha curcas (jatropha)EuphorbiaceaeMain
                                                    Lathyrus odoratus (sweet pea)FabaceaeMain
                                                      Lilium (lily)LiliaceaeMain
                                                        Linum usitatissimum (flax)Main
                                                        Lycium chinense (chinese wolfberry)SolanaceaeUnknown
                                                        Malus (ornamental species apple)RosaceaeUnknown
                                                        Malus domestica (apple)RosaceaeMain
                                                          Mangifera indica (mango)AnacardiaceaeUnknown
                                                          Manihot esculenta (cassava)EuphorbiaceaeMain
                                                          Medicago sativa (lucerne)FabaceaeMain
                                                            Melissa officinalis (Lemon balm)LamiaceaeUnknown
                                                            Mentha (mints)LamiaceaeUnknown
                                                            Mentha arvensis (Corn mint)LamiaceaeUnknown
                                                              Morus alba (mora)MoraceaeMain
                                                                Musa (banana)MusaceaeMain
                                                                  Musa paradisiacaUnknown
                                                                  Musa x paradisiaca (plantain)MusaceaeUnknown
                                                                  Nelumbo nucifera (sacred lotus)NelumbonaceaeUnknown
                                                                  Nicotiana tabacum (tobacco)SolanaceaeMain
                                                                  Ocimum basilicum (basil)LamiaceaeOther
                                                                  Origanum majorana (sweet marjoram)LamiaceaeUnknown
                                                                  Oryza sativa (rice)PoaceaeMain
                                                                  Papaver (poppies)PapaveraceaeMain
                                                                    Paulownia tomentosa (paulownia)ScrophulariaceaeMain
                                                                      Pelargonium hortorumGeraniaceaeUnknown
                                                                      Perilla frutescensLamiaceaeUnknown
                                                                      Phaseolus (beans)FabaceaeMain
                                                                        Piper aduncum (spiked pepper)PiperaceaeUnknown
                                                                        Piper nigrum (black pepper)PiperaceaeMain
                                                                          Piper umbellatumPiperaceaeUnknown
                                                                          Pisum sativum (pea)FabaceaeOther
                                                                          Platostoma chinensisLamiaceaeUnknown
                                                                          Plectranthus amboinicus (Indian borage)LamiaceaeUnknown
                                                                          Poaceae (grasses)PoaceaeMain
                                                                            Polyphagous (polyphagous)Main
                                                                              Prunus mume (Japanese apricot tree)RosaceaeOther
                                                                                Psophocarpus tetragonolobus (winged bean)FabaceaeMain
                                                                                  Raphanus sativus (radish)BrassicaceaeMain
                                                                                  Reynoutria multifloraUnknown
                                                                                  Ricinus communis (castor bean)EuphorbiaceaeMain
                                                                                  Rosa (roses)RosaceaeMain
                                                                                    Rosmarinus officinalis (rosemary)LamiaceaeUnknown
                                                                                    Ruellia tuberosaUnknown
                                                                                    Ruta graveolens (common rue)RutaceaeUnknown
                                                                                    Salvia officinalis (common sage)LamiaceaeUnknown
                                                                                    Sesbania grandiflora (sesbania)FabaceaeMain
                                                                                      Sesbania sesban (sesban)FabaceaeMain
                                                                                      Solanum aculeatissimumSolanaceaeUnknown
                                                                                      Solanum lycopersicum (tomato)SolanaceaeMain
                                                                                        Solanum melongena (aubergine)SolanaceaeMain
                                                                                        Solanum tuberosum (potato)SolanaceaeMain
                                                                                        Sorghum bicolor (sorghum)PoaceaeMain
                                                                                        SpathoglottisUnknown
                                                                                        Spinacia oleracea (spinach)ChenopodiaceaeOther
                                                                                        Syzygium aromaticum (clove)MyrtaceaeMain
                                                                                          Tagetes (marigold)AsteraceaeOther
                                                                                            Tectona grandis (teak)LamiaceaeMain
                                                                                            Theobroma cacao (cocoa)MalvaceaeMain
                                                                                              Trianthema portulacastrum (horse purslane)AizoaceaeMain
                                                                                              Trifolium alexandrinum (Berseem clover)FabaceaeOther
                                                                                              Trigonella foenum-graecum (fenugreek)FabaceaeMain
                                                                                                Vigna mungo (black gram)FabaceaeMain
                                                                                                Vigna radiata (mung bean)FabaceaeMain
                                                                                                Vigna unguiculata (cowpea)FabaceaeMain
                                                                                                Vitis vinifera (grapevine)VitaceaeMain
                                                                                                Zea mays (maize)PoaceaeMain
                                                                                                Zinnia elegans (zinnia)AsteraceaeMain

                                                                                                  Growth Stages

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                                                                                                  Flowering stage, Fruiting stage, Vegetative growing stage

                                                                                                  Symptoms

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                                                                                                  On most crops, damage arises from extensive feeding by larvae, leading to complete stripping of the plants.

                                                                                                  On Cotton

                                                                                                  Leaves are heavily attacked and bolls have large holes in them from which yellowish-green to dark-green larval excrement protrudes.

                                                                                                  On Tobacco

                                                                                                  Leaves develop irregular, brownish-red patches and the stem base may be gnawed off.

                                                                                                  On Maize

                                                                                                  The stems are often mined and young grains in the ear may be injured.

                                                                                                  List of Symptoms/Signs

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                                                                                                  SignLife StagesType
                                                                                                  Leaves / external feeding

                                                                                                  Biology and Ecology

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                                                                                                  S. litura eggs are laid in clusters of several hundreds, usually on the upper surface of the leaves. Fecundity varies from 2000 to 2600 eggs, and oviposition days vary from 6 to 8 days. The developmental thresholds and thermal requirements for different stages of S. litura are 64 day degrees above threshold 8°C, from oviposition to egg hatch, the larval period required 303 degree days and the pupal stage 155 degree days above a 10°C threshold. The response of various stages of S. litura to temperatures under constant laboratory conditions was similar to that under field conditions. The upper development threshold temperature for all stages was 37°C, and 40°C was lethal (Ranga Rao et al., 1989).

                                                                                                  Eggs take 2-3 days to hatch, the larvae disperse quickly from the egg batch in groundnut. Newly hatched larvae can be detected by the 'scratch marks' they make on the leaf surface. The older larvae are night feeders and are usually found in the soil around the base of the plant during the daytime. They can chew large areas of leaf and at high population densities cause complete defoliation. The larvae can migrate in large groups from one field to another. In lighter soils, the larvae while hiding in the soil during daytime can also cause damage to groundnut pods.

                                                                                                  The larvae go through six instars and the final instars weigh up to 800 mg. Individual larvae can consume around 4 g fresh weight of groundnut foliage. However, 80% of the total consumption is in the final instar.

                                                                                                  Pupation takes place in the soil close to the plants. The pupal period lasts about 7-10 days.

                                                                                                  After adult emergence, peak oviposition occurs on the second night. Females mate three or four times during their lifetime, while males mate up to 10 times.

                                                                                                  In Andhra Pradesh, India, S. litura completes 12 generations a year, each lasting slightly more than a month in winter and less than a month in the hot season.

                                                                                                  Climate

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                                                                                                  ClimateStatusDescriptionRemark
                                                                                                  C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
                                                                                                  Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
                                                                                                  Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
                                                                                                  Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

                                                                                                  Air Temperature

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                                                                                                  Parameter Lower limit Upper limit
                                                                                                  Absolute minimum temperature (ºC) 6
                                                                                                  Mean annual temperature (ºC) 10 37
                                                                                                  Mean maximum temperature of hottest month (ºC) 37
                                                                                                  Mean minimum temperature of coldest month (ºC) 8

                                                                                                  Natural enemies

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                                                                                                  Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                                                                  Apanteles colemani Parasite Arthropods|Larvae
                                                                                                  Apanteles prodeniae Parasite Arthropods|Larvae
                                                                                                  Aspergillus ochraceus Pathogen
                                                                                                  Bacillus thuringiensis Pathogen Arthropods|Larvae
                                                                                                  Bacillus thuringiensis kurstaki Pathogen
                                                                                                  Beauveria bassiana Pathogen Arthropods|Larvae
                                                                                                  Brachymeria lasus Parasite
                                                                                                  Bracon brevicornis Parasite Arthropods|Larvae
                                                                                                  Campoletis chlorideae Parasite Arthropods|Larvae
                                                                                                  Charops bicolor Parasite Arthropods|Larvae
                                                                                                  Charops obtusus Parasite Arthropods|Larvae
                                                                                                  Chelonus formosanus Parasite Eggs; Arthropods|Larvae
                                                                                                  Copidosoma floridanum Parasite Eggs; Arthropods|Larvae
                                                                                                  Cotesia marginiventris Parasite Arthropods|Larvae
                                                                                                  Cotesia ruficrus Parasite Arthropods|Larvae
                                                                                                  Cotesia sesamiae Parasite Arthropods|Larvae
                                                                                                  Cytoplasmic polyhedrosis virus (CPV) Pathogen Arthropods|Larvae
                                                                                                  Elasmus nephantidis Parasite
                                                                                                  Eocanthecona furcellata Predator
                                                                                                  Eriborus argenteopilosus Parasite Arthropods|Larvae
                                                                                                  Euplectrus Parasite Arthropods|Larvae
                                                                                                  Euplectrus gopimohani Parasite Arthropods|Larvae
                                                                                                  Eurytoma syleptae Hyperparasite Arthropods|Larvae; Arthropods|Pupae
                                                                                                  Glyptapanteles ashmeadi Parasite Arthropods|Larvae
                                                                                                  Granulosis virus Pathogen Arthropods|Larvae
                                                                                                  Metarhizium anisopliae Pathogen Arthropods|Larvae
                                                                                                  Metopius rufus Parasite Arthropods|Larvae
                                                                                                  Microchelonus heliopae Parasite Eggs; Arthropods|Larvae
                                                                                                  Microgaster tuberculifera Parasite Arthropods|Larvae
                                                                                                  Microplitis Parasite Arthropods|Larvae
                                                                                                  Microplitis pallidipes Parasite Arthropods|Larvae
                                                                                                  Microplitis prodiniae Parasite Arthropods|Larvae
                                                                                                  Netelia ferruginea Parasite Arthropods|Larvae
                                                                                                  Nomuraea rileyi Pathogen Arthropods|Larvae
                                                                                                  Nosema carpocapsae Pathogen Arthropods|Larvae
                                                                                                  Nucleopolyhedrosis virus Pathogen Arthropods|Larvae
                                                                                                  Oxyopes sertatus Predator
                                                                                                  Peribaea orbata Parasite Arthropods|Larvae
                                                                                                  Rhynocoris kumari Predator
                                                                                                  Rhynocoris marginatus Predator
                                                                                                  Telenomus hawai Parasite Eggs
                                                                                                  Telenomus remus Parasite Eggs
                                                                                                  Telenomus spodopterae Parasite Eggs
                                                                                                  Temelucha biguttula Parasite Arthropods|Larvae
                                                                                                  Tetrastichus israeli Parasite
                                                                                                  Trichogramma chilonis Parasite Eggs
                                                                                                  Trichogramma dendrolimi Parasite Eggs
                                                                                                  Trichospilus pupivora Parasite
                                                                                                  Winthemia Parasite Arthropods|Pupae
                                                                                                  Zele chlorophthalma Parasite Arthropods|Larvae

                                                                                                  Notes on Natural Enemies

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                                                                                                  After Rao et al., 1993.

                                                                                                  S. litura is known to be atttacked by many natural enemies at various life stages. Altogether, about 131 species of natural enemies have been reported from different parts of the world.

                                                                                                  Egg Parasitoids

                                                                                                  Four species of trichogrammatids, one scelionid and one braconid which had been reported as egg parasitoids of S. litura, an unidentified Chelonus species and species of Telenomus, have also been reported as both egg and larval parasitoids. A total of 10 egg parasitoids have been reported from different parts of the host distribution. Among the trichogrammatids, T. chilonis from India (Joshi et al., 1979; Patel et al., 1971) and T. dendrolimi from China (Chiu and Chou, 1976) are the most common. These species are often reported from the eggs of several other hosts.

                                                                                                  Larval Parasites

                                                                                                  Generally, the larval stage of S. litura is more prone to parasitism. Larval parasitoids of S. litura attack young to mature larvae and a few also attack eggs and larvae, and larvae and prepupae. Fifty-eight parasitoid species have been reported to attack the larval stage of this species. Of these, 47% were braconids, 19% ichneumonids, 16% tachinids, 10% eulophids, 3% chalcids, and 2% scelionids, encrytids and muscids. In general, 84% were Hymenoptera, and 16% Diptera.

                                                                                                  In India, 32 different species of parasitoids have been reported as larval parasitoids of S. litura. Among these, Apanteles and species of Bracon were the most commonly reported. Rai (1974) surveyed vegetable crops in the state of Karnataka and found that 10% of larval mortality was caused by Chelonus formosanus. Jayanth and Nagarkatti (1984) reported the emergence of up to 12 tachninid parasitoids (Peribaea orbata) from a single S. litura larva in Karnataka state, India.

                                                                                                  Rao and Satyanarayana (1984), during a pest survey of natural enemies of S. litura in Andhra Pradesh, India, reported Zele chlorophthalma as a larval parasitoid.

                                                                                                  Sathe (1987) in a survey for natural enemies of S. litura in Maharashtra region of India reported Campoletes chlorideae and Apanteles colemani. During the same survey two new Braconid species (Enicospilus sp. and Echthromorpha sp.) were found responsible for the 5% parasitization of S. litura, while A. colemani and A. prodeniae parasitized up to 20% larvae.

                                                                                                  Pupal Parasitoids

                                                                                                  Relatively few pupal parasitoids have been reported from S. litura. Eight parasitoid species have been reported from the pupal stage of S. litura, one of which is a larval-pupal parasitoid (Ichneumon sp.) and one a prepupal parasitoid (species of Chelonus).

                                                                                                  Predators

                                                                                                  Altogether 36 predatory insects from 14 families and 12 species of spiders, representing six families were reported to feed on S. litura eggs, larvae and pupae in different parts of the world. Of the total predators reported to feed on S. litura, 50% of the insect predatory fauna and 83% of the spiders were from India.

                                                                                                  Diseases

                                                                                                  Protozoa
                                                                                                  Nosema carpocapsae was found to infect S. litura larvae in New Zealand (Malone and Wigley, 1980), India (Narayanan and Jayaraj, 1979), Japan (Watanabe, 1976) and China (Tsai et al., 1978; Li and Wenn, 1987).

                                                                                                  Fungi
                                                                                                  So far four fungi have been reported to infect S. litura and cause physiological disorders in larval growth and development: Aspergillus flavus, Beauveria bassiana, Nomuraea rileyi and Metarhizium anisopliae. Zaz and Kishwaha (1983) reported B. bassiana infecting S. litura in cauliflower crops in Rajasthan. Siddaramaiah et al. (1986) reported an incidence of larval infection with M.anisopliae in groundnut in Karnataka. The infection first appeared in the second fortnight of June, was highest in mid-August, and decreased by November.

                                                                                                  Viruses
                                                                                                  Viral diseases of this species have been reported from China, Japan, India and New Zealand. Among the viruses, nuclear polyhedrosis viruses are the most common and potent. Narayanan (1985) from Karnataka, reported the occurrence of a granulosis virus in dead S. litura larvae. Eggs and all six larval instars were highly susceptible to the virus, the mortality was 100% in eggs and first to fifth-instar larvae and 50% in the last larval instar. The disease killed older larvae more rapidly than younger ones.

                                                                                                  Nematodes

                                                                                                  Four nematode species have been reported parasitizing S. litura in India and one of them has also been reported in Japan. Bhatnagar et al. (1985) found S. litura larvae parasitized by the mermithid nematodes Ovomermis albicans, Hexamermis sp. and Pentatomermis sp. They observed more nematode activity on alfisols than on vertisols. They also discussed the population dynamics and distribution of nematodes and the arthropod hosts. Kondo and Ishibashi (1984) explained the infectivity and propagation of entomogenous nematodes Steinernema sp. on S. litura from Japan.

                                                                                                  Means of Movement and Dispersal

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                                                                                                  Natural dispersal

                                                                                                  The moths have a flight range of 1.5 km during a period of 4 h overnight, facilitating dispersion and oviposition on different hosts (Salama and Shoukry, 1972). They can accordingly fly quite long distances. The caterpillars can migrate over short distances.

                                                                                                  Accidental introduction

                                                                                                  In international trade, eggs or larvae may be present on planting material, cut flowers or vegetables; for example, the introduction of S. litura into the UK was on aquatic plants imported from Singapore (Aitkenhead et al., 1974). The pupae can be present in soil. The pupae are long-lived and could be transported over a considerable distance if not crushed, but to establish a viable population several specimens of both sexes need to be transported.

                                                                                                  Pathway Causes

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                                                                                                  CauseNotesLong DistanceLocalReferences
                                                                                                  Aid Yes Yes
                                                                                                  Botanical gardens and zoos Yes
                                                                                                  Breeding and propagation Yes
                                                                                                  Crop production Yes Yes
                                                                                                  Cut flower trade Yes

                                                                                                  Pathway Vectors

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                                                                                                  VectorNotesLong DistanceLocalReferences
                                                                                                  Aircrafteggs, larvae, pupae Yes
                                                                                                  Land vehicleseggs, larvae, pupae, moths Yes Yes
                                                                                                  WindAdult moth Yes
                                                                                                  Plants or parts of plants Yes
                                                                                                  Containers and packaging - wood Yes
                                                                                                  Containers and packaging - non-wood Yes

                                                                                                  Plant Trade

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                                                                                                  Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
                                                                                                  Bulbs/Tubers/Corms/Rhizomes eggs; larvae; pupae Yes Yes Pest or symptoms usually invisible
                                                                                                  Flowers/Inflorescences/Cones/Calyx eggs; larvae; pupae Yes Pest or symptoms usually visible to the naked eye
                                                                                                  Fruits (inc. pods) eggs; larvae; pupae Yes Yes Pest or symptoms usually invisible
                                                                                                  Leaves eggs; larvae; pupae Yes Pest or symptoms usually visible to the naked eye
                                                                                                  Stems (above ground)/Shoots/Trunks/Branches eggs; larvae; pupae Yes Pest or symptoms usually visible to the naked eye
                                                                                                  True seeds (inc. grain) eggs; larvae; pupae Yes Yes Pest or symptoms usually invisible
                                                                                                  Plant parts not known to carry the pest in trade/transport
                                                                                                  Bark
                                                                                                  Growing medium accompanying plants
                                                                                                  Roots
                                                                                                  Wood

                                                                                                  Wood Packaging

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                                                                                                  Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
                                                                                                  Loose wood packing material No
                                                                                                  Non-wood No
                                                                                                  Processed or treated wood No
                                                                                                  Solid wood packing material with bark No
                                                                                                  Solid wood packing material without bark No

                                                                                                  Impact Summary

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                                                                                                  CategoryImpact
                                                                                                  Economic/livelihood Negative
                                                                                                  Environment (generally) Positive and negative

                                                                                                  Impact

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                                                                                                  S. litura larvae are polyphagous defoliators, seasonally common in annual and perennial agricultural systems in tropical and temperate Asia. This noctuid is often found as part of a complex of lepidopteran and non-lepidopteran foliar feeders but may also damage tubers and roots. Hosts include field crops grown for food and fibre, plantation and forestry crops, as well as certain weed species.

                                                                                                  Most work on the economic impact of S. litura has been conducted in India where it is a serious pest of a range of field crops. It has caused 12-23% damage to tomatoes in the monsoon season, and 9-24% damage in the winter (Patnaik, 1998). In a 40- to 45-day-old potato crop, damage ranged from 20 to 100% in different parts of the field depending on moisture availability. Larval populations peaked at four fully-grown larvae per square metre when the crop was 60-70 days old. Larvae also attacked exposed tubers when young succulent leaves were unavailable. Up to 2% of tubers were damaged in August-September and February (Trivedi, 1988). S. litura is also a pest of sugarbeet, with infestations commencing in March and peaking in late March and April (Chatterjee and Nayak, 1987). Severe infestations led to skeletonisation of leaves as well as feeding holes in roots that rendered the crop 'virtually unfit for marketing'. Late harvested crops were most severely affected and, in extreme cases, 100% of the roots were damaged, leading to 'considerable' yield reduction. Work on this species in a complex of other sugarbeet defoliators (Spodoptera exigua and Spilosoma obliqua) led to the development of an interactive exponential model based on length and severity of defoliation. It explained 88-90% of the variability in root and sugar yields and suggested the need for pest control when defoliation exceeded 25% during April. Control was not required if the pest appeared after the first week of May (Singh and Sethi, 1993).

                                                                                                  S. litura is one of six defoliating pests of fodder cowpea which, in a field experiment, were responsible for consuming up to 85.5% of leaf area (Ram et al., 1989). Aroid tuber crops (including taro (Colocasia esculenta)) suffered yield losses of up to 29% as a result of infestation by S. litura, Aphis gossypii and spider mites (Pillai et al., 1993). S. litura is also a member of a complex that causes extensive defoliation of soyabean (Bhattacharjee and Ghude, 1985). Defoliation as severe as 48.7% during the pre-bloom stage of growth caused no 'marked' difference from a control treatment in which defoliation was prevented by repeated insecticide application. Number and weight of pods and grains per plant were, however, reduced when defoliation occurred at, or after, blooming.

                                                                                                  In groundnut, S. litura is one of several pests that can be important during the pegging, podding and pod maturation stages of growth (Singh and Sachan, 1992). Several studies have aimed at quantifying the damage attributable to S. litura. Field experiments by Panchabhavi and Raj (1987) extended over 2 years and used artificial infestation of groundnut plots of 15 m² with differing densities of S. litura. Infestation levels of just three egg masses (of 250 eggs each) caused significant loss of groundnut pods and haulms. Infestation with 12 egg masses per plot led to a haulm yield reduction of up to 43.7% and a pod yield reduction as high as 27% compared with an insecticide-protected control treatment. In other field experiments over 3 consecutive years, leaf damage attributed to S. litura tended to decline with delayed sowing time irrespective of groundnut cultivar (Patil et al., 1996). Leaf damage fell from 51.8% for mid-June sown crops to 19.2% for late-July sown crops. Mean pod yields were 2.68 and 0.99 t/ha, respectively.

                                                                                                  Another field study determined the effect of artificially infesting individual groundnut plants with third-instar S. litura larvae 15, 30 or 45 days after emergence (d.a.e.) (Dhir et al., 1992). The most severe damage occurred when plants were infested with three larvae 15 d.a.e. These lost 98.3% of leaf area, and pod yield was reduced by 50%. Even single larvae caused the leaf area to be more than halved and pod yield to fall by 27.3%. Plants infested 30 d.a.e. suffered similar levels of damage but those infested 45 d.a.e. were less severely affected.

                                                                                                  Selveraj et al. (2014) conducted a study of the ecological factors on the incidence and development of S. litura on cotton. They found a positive correlation with relative humidity, sunshine hours and dewfall, but a negative correlation with wind velocity. Determination of the effects of different weather factors on the population and incidence of S. litura in cotton is essential for effective pest management.

                                                                                                  S. litura causes damage to many species of forest and plantation trees and shrubs (Roychoudhury et al., 1995). It is responsible for brown flag syndrome in banana (Ranjith et al., 1997), and 5-10% fruit damage in grapes (Balikai et al., 1999). In Paulownia nurseries and plantations a complex of at least 24 defoliating pest species causes damage. Within this complex, S. litura was considered the most important noctuid species, with an incidence of 72% in weekly surveys (Kumar and Ahmad, 1998). Peak activity occurred in July and September, with an average of 6.5 and 5.2 larvae per plant in these months, respectively. During this period, many plants were completely defoliated by S. litura. In teak, it is one of about 139 defoliators that attack all stages from seedlings to mature trees (Roychoudhury et al., 1995). S. litura is abundant on teak in June and July and damage incidence in seedlings has been reported to be as high as 56%. Late-instar larvae were found to feed preferentially on mature teak leaves, whilst early instars fed on leaves of intermediate age. High concentrations of polyphenols in young leaves (Roychoudhury et al., 1995) may reduce their attractiveness to S. litura larvae but differing levels of susceptibility among nine teak clones were attributed to the nitrogen:potassium ratio of the foliage (Roychoudhury et al., 1998).

                                                                                                  Two separate Indian studies concerned the potential of S. litura as a biological control agent of weeds. Sites of larval feeding on leaves and petioles of the aquatic weed Eichhornia crassipes were said to be vulnerable to infection by fungi, and overall damage levels by S. litura were estimated to be 20-25% (Jamil et al., 1984). It was recognized, however, that the potential use of this polyphagous agent needs caution because preferred hosts include important crops such as castor and cotton. In a second study, S. litura attacked the weeds, Marsilea quadrifolia, Ammannia baccifera and Eclipta alba, and although up to 10 larvae were recorded per plant on nearby rice, no damage was reported to this crop (Sain et al., 1983).

                                                                                                  Studies elsewhere in southern Asia illustrate the economic impact of S. litura. In Pakistan, it is one of several lepidopteran pests attacking a wide range of crops including cotton and rice (Ahmad and Kamaluddin, 1987), as well as cabbage, tobacco, groundnut, soyabean, lucerne, gram, cowpea, tomato, cauliflower, carrot, onion, brinjal, turnip, radish and spinach (Maree et al., 1999). In the latter study, S. litura was present in a December-planted cabbage crop during January-March with densities peaking at 4.55/plant on 2 February. Damage was highest in early February and there was a positive correlation between plant damage and pest density. Fieldwork in Bangladesh led to a 10% visual damage spraying threshold being proposed for S. litura and Plutella xylostella (Ali and Bakshi, 1994).

                                                                                                  In eastern Asia, damage by S. litura occurs as far north as Japan, extending south to Indonesia. Japanese soyabean field plots artificially infested with one or two S. litura egg masses per plant suffered estimated leaf area reductions of 14.3 and 23.2% and yield losses of 13.9 and 24.7%, respectively, compared with control plots (Higuchi et al., 1994). Similar work indicated that yield losses caused by early infestation (at the flowering and pod-development stages) were attributable chiefly to reduced production of pods per plant, whilst infestations on more mature plants affected yield by reducing the weight of individual beans (Higuchi, 1991). In studies with red (adzuki) bean, the effect of defoliation by S. litura on yield was investigated by artificial defoliation. Combined with observations that the average leaf area consumed by S. litura was 203.9 cm², of which 82% was consumed by final-instar larvae, an action threshold of two final-instar larvae/plant (causing 5% yield loss) was estimated (Katayama and Sano, 1989). Equivalent thresholds were developed for protected crops in Korea (Nakasuji and Matsuzaki, 1977). In this instance, a 10% yield loss threshold was applied and the threshold densities of S. litura were 4.6 and 15.4 neonates or 0.8 and 2.6 egg masses/m², for aubergine and sweet peppers, respectively. S. litura is also found in Welsh onion crops in Korea, but is less damaging than other lepidopterans such as Liriomyza chinensis (Ahn et al., 1991).

                                                                                                  A lepidopteran pest complex including S. litura, Pieris rapa and Plutella xylostella damages cabbage in China (Zhou et al., 1996). Experiments to determine larval feeding capacity showed S. litura to be intermediate amongst these species (Zhu et al., 1994). In the Zhejiang region of China, S. litura is usually first recorded in June on legumes, aubergines, tomatoes, sweet peppers and tree fruits, and subsequently damages cabbage seedlings in July (Chen et al., 1999). Damage becomes more severe in August and September when cauliflower, sweet potato and legumes may also be attacked. The pest appears unable to overwinter in this area and is virtually absent after the end of October. In Taiwan, S. litura attacks gladiolus foliage (Wang, 1982; Liu, 1998) as well as soyabean and adzuki bean (Lee, 1989). Damage ranged between 3 and 13% over three consecutive years for the latter crops.

                                                                                                  In South-East Asia, S. litura has been recorded damaging tobacco in Malaysia (Hamid et al., 1992), mungbean (Vigna radiata) in Thailand (Sepswasdi et al., 1991), tree legumes in Indonesia (Matsumoto, 2000) and Paraserianthes falcataria seedlings in the Philippines (Braza, 1990).

                                                                                                  Roa et al. (2012) studied the impact of climate change on the development of S. litura in the near future and reported a change in the nutrient value of groundnut foliage and as a result a higher consumption, lower digestive efficiency, slower growth and longer time to pupation (1 day more than ambient).

                                                                                                  Risk and Impact Factors

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                                                                                                  Invasiveness
                                                                                                  • Invasive in its native range
                                                                                                  • Proved invasive outside its native range
                                                                                                  • Has a broad native range
                                                                                                  • Abundant in its native range
                                                                                                  • Highly adaptable to different environments
                                                                                                  • Is a habitat generalist
                                                                                                  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
                                                                                                  • Tolerant of shade
                                                                                                  • Capable of securing and ingesting a wide range of food
                                                                                                  • Highly mobile locally
                                                                                                  • Benefits from human association (i.e. it is a human commensal)
                                                                                                  • Fast growing
                                                                                                  • Has high reproductive potential
                                                                                                  • Gregarious
                                                                                                  Impact outcomes
                                                                                                  • Host damage
                                                                                                  • Infrastructure damage
                                                                                                  • Negatively impacts agriculture
                                                                                                  • Negatively impacts livelihoods
                                                                                                  Impact mechanisms
                                                                                                  • Herbivory/grazing/browsing
                                                                                                  Likelihood of entry/control
                                                                                                  • Highly likely to be transported internationally accidentally
                                                                                                  • Highly likely to be transported internationally deliberately
                                                                                                  • Difficult to identify/detect as a commodity contaminant
                                                                                                  • Difficult to identify/detect in the field
                                                                                                  • Difficult/costly to control

                                                                                                  Detection and Inspection

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                                                                                                  The presence of newly hatched larvae can be detected by the 'scratch' marks they make on the leaf surface. The older larvae are night-feeders and are usually found in the soil around the base of plants during the day. They chew large areas of the leaf, and can, at high population densities, strip a crop of its leaves. In such cases, larvae migrate in large groups from one field to another in search of food.

                                                                                                  Similarities to Other Species/Conditions

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                                                                                                  S. litura can be easily confused with S. littoralis as in both cases adults and larvae are similar, and they can be distinguished only through examination of genitalia. On dissection of the genitalia, ductus and ostium bursae are the same length in female littoralis, different lengths in litura. The shape of the juxta in males is very characteristic, and the ornamentation of the aedeagus vesica is also diagnostic. The presence of newly hatched larvae can be detected by the 'scratch marks' they make on the leaf surface.

                                                                                                  For more information on the morphological discrimination between the adult, pupal and larval stages of the two species, see Mochida (1973).

                                                                                                  An EPPO standard provides guidance for the identification of S. littoralis, S. litura, S. frugiperda and S. eridania (OEPP/EPPO, 2015).

                                                                                                  Prevention and Control

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                                                                                                  Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

                                                                                                  Introduction

                                                                                                  After Rao et al. (1993).

                                                                                                  The green revolution in Asia brought with it an increased awareness of the potential of insecticides for increasing the sustainability of rice production. Unfortunately, the involvement of farms in insecticide-related technologies did not proceed as fast as the rate of subsidy spread and the overspill of insecticide usage into the fields of legume growers and horticulturalists. Legume pests are increasing in economic importance throughout Asia due to the destruction of natural control systems, and the build-up of insecticide resistance following the 'spraymania' of many farmers. If this is to be counteracted, natural control needs to be given increased emphasis as a component of the IPM approach. S. litura populations in groundnut fields are increasing in number and intensity, especially in fields where insecticides have been applied (Stechmann and Semisi, 1984; Rao and Shanower, 1988).

                                                                                                  Chemical Control

                                                                                                  In the past, the control of arthropods depended mostly on inexpensive and efficient insecticides. But in recent years populations of many pests including S. litura have developed resistance to many commercially available pesticides (Ramakrishnan et al., 1984; Naeem Abbas et al., 2014). Studies at ICRISAT between 1991 and 1996 revealed the occurrence of resistance to cypermethrin, fenvalerate and quinalphos, by 197-, 121-, 29- and 362-fold, respectively (JA Wightman, ICRISAT, Andhra Pradesh, India, personal communication, 1996).

                                                                                                  The control of arthropod pests is therefore becoming increasingly difficult and it is vital that all biological alternatives to insecticides need to be given greater priority, both in research and application. 

                                                                                                  New insecticides have been tested to deal with resistant strains of this moth and some promising results are coming forward (Venkateswarlu et al., 2005). Neem oil microemulsion proved significantly superior than macroemulsion (Swaran Dhingra et al., 2006).

                                                                                                  New molecules such as chlorantraniliprole, spinosad and emamectin benzoate have shown promising results against S. litura (Gadhiya et al., 2014) but chlorantraniliprole gave the highest cost: benefit ratio among pesticides tested by Patil et al. (2014) on soyabeans.

                                                                                                  Chatterjee and Mondal (2012) tested a number of new chemicals and their application methods on different vegetable crops in India and South-East Asia against lepidopterous pests and found flubendiamide, spinosad and chlorfenapyr to be the most effective.

                                                                                                  Following studies on the sublethal effects of mathofenozide, Shahout et al. (2011) concluded that the effects of methoxyfenozide with its sterilizing properties, if used strategically on S. litura, might induce changes in the population dynamics of this pest in vegetable crops and could be considered a potent insecticidal compound for controlling this pest.

                                                                                                  Suganthy and Sakthivel (2013) studied different bio-pesticides against S. litura infesting fields of Gloriosa superba and showed that flavonoids could be used as an alternative to chemical pesticides in the gloriosa ecosystem and as a component in organic pest management.

                                                                                                  Plant oils and insecticides mixtures (synthetic pyrethroids) gave a higher mortality rate on 8-day-old larvae of S. litura than the synthethic pyrethroids alone (Anju and Srivastava, 2012).

                                                                                                  Biological Control

                                                                                                  In the past the mass releases of egg and larval parasitoids for the control of S. litura in different crops in different geographical regions had achieved only partial success (Patel et al., 1979; Michael et al., 1984). Observations in ICRISAT groundnut fields revealed more leaves with defoliator damage in insecticide applied fields than unsprayed areas (Wightman et al., 1990). Similar observations were also made during farmers' field surveys in the post-rainy season in coastal Andhra Pradesh, India (Rao and Shanower, 1988). Stechmann and Semisi (1984) also shared the same opinion after surveying taro fields in Western Samoa. In view of the development of insecticidal resistance and the destruction of the natural enemies, and the polyphagous nature of this species, there is a need to give more consideration to the role of natural enemies as a component of integrated approaches to managing S. litura.

                                                                                                  Egg parasitoids
                                                                                                  Mass releases of an indigenous egg-larval parasite Chelonus heliopae in 1971-73 in Anand, Gujarat, India, against S. litura in cauliflower proved ineffective in controlling the pest. During 1974, weekly release of Telenomus remus, an egg parasitoid, in a tobacco nursery did not result in any parasitism. However, five weekly releases of 50,000 parasites per 0.2 ha and two releases of 15,000 parasitoids per 0.2 ha in cauliflower resulted in 60% parasitism (Patel et al., 1979).

                                                                                                  T. remus was introduced to Western Samoa and was recorded by Braune (1982) as a common egg-larval parasitoid of S. litura, with parasitism averaging 54%. Complete parasitization was observed only in small egg masses (up to 150 eggs) and the percentage of parasitization decreased with an increase in size of egg mass. T. remus could oviposit only in host eggs on the surface of the host egg mass. Thus the effectiveness of T. remus was limited to the large compact egg masses of S. litura.

                                                                                                  Larval parasitoids
                                                                                                  Six parasitoid species, Apanteles ruficrus, Cotesia marginiventris, Apanteles kazak, Campoletes chloridae, Hyposoter didymator and T. remus were introduced to Western Australia from overseas in 1978-83 and released against S. litura and 11 other economically important pests. The highest level of parasitism by A. ruficrus (80% and above) was noticed in Mythimna sp. (Michael et al., 1984).

                                                                                                  In Western Samoa, Stechmann and Semisi (1984) collected information on S. litura damage levels in relation to natural populations of Apanteles spp. in taro fields. They found that this pest was more severe on taro where insecticides and herbicides were widely used, which perhaps created imbalance between the pest and its natural enemies. Barrion and Litsinger (1987) reported the presence of Peribaea orbata as a gregarious larval parasitoid on S. litura.

                                                                                                  Wang et al. (2014) studied the relationship between the larval parasitoid Meteorus pulchricornis and the bacterium Empedobacter brevis. The study suggested that the bacteria has a negative effect on M. pulchricornis, but the impact could be alleviated by using low bacteria concentration and extending the time between the application and wasp release in biological control practices.

                                                                                                  Predators
                                                                                                  The biology of Canthoconidia furcellata was studied in the laboratory with a view to using this predator in an integrated pest management programme for tobacco pests. Chu and Chu (1975) studied the effects of temperature on the growth of C. furcellata and found that 71,216 and 134 degree days were required for egg, nymph and adult stages, respectively. It was concluded that there are five to six generations per year of this predator in northern Taiwan.

                                                                                                  Nakasuji et al. (1976) observed a predatory wasp, preferentially selecting fifth- and sixth-instar larvae over early instars. The wasps were more active and attacked more larvae in fields with high larval density than those with low larval density. However, the percentage of predation was lower in the field with highest density of S. litura larvae.

                                                                                                  Deng and Jim (1985) reported Conocephalus sp. as a new predator on egg masses of S. litura in Guanxi, China. This katydid was successfully reared on an artificial diet. Field releases of nymphs and adults of Conocephalus sp. were attempted for control of Scirpophaga incertulus.

                                                                                                  Pathogens
                                                                                                  Ansari et al. (1987) reported Serratia marcescens from Karnataka, India, attacking larvae of the noctuids Helicoverpa armigera and S. litura. In laboratory tests, S. litura was found to be more susceptible to the bacterium than H. armigera. The bacterium was equally pathogenic when ingested through artificial diet or the natural food plant, but pathogenicity by contact application to the body of larvae was poor.

                                                                                                  Zaz and Kushwaha (1983) found Bacillus thuringiensis to be an effective microbial insecticide against S. litura larvae in cauliflower fields in Rajasthan, India.

                                                                                                  The efficiency of B. thuringiensis was enhanced significantely though protoplast fusion with a strain of Bacillus subtilis (Kannan Revathi et al., 2014).

                                                                                                  Asayama and Ohoishi (1980) from Japan and Phadke and Rao (1978) from India, investigated the pathogenicity of a green muscardine fungus Nomuraea rileyi. Laboratory studies in India indicated that this fungus was harmless to eggs of an egg parasitoid, Telenomus preditor, on Achaea janata and recommended the combined use of the fungus and the egg parasitoid in biocontrol programmes against A. janata. This may also apply to S. litura management.

                                                                                                  Laboratory studies were undertaken to evaluate the bioefficacy of Beauveria bassiana against third-instar larvae of S. litura. B. bassiana was identified, isolated and maintained from field-collected cadavers of lepidopteran larvae. Minimum mortality was observed in the control, i.e. 23.3%, and the percentage mortality increased as the number of spores increased (Gupta and Bhupendra Kumar, 2014).

                                                                                                  Research was also carried out on entophytic fungi (Khuskia oryzae and Cladosporium uredinicola), which showed adverse effects on survival and fitness of the insects (Abhinay et al., 2014).

                                                                                                  Krishnaiah et al. (1985) conducted field trials with a nuclear polyhedrosis virus against S. litura damage in black gram (Vigna mungo) fields in Andhra Pradesh, India. Two sprays of virus suspension gave effective control similar to chemical insecticides tested.

                                                                                                  Chari et al. (1985) evaluated the effectiveness of integrated management of natural enemies and viral diseases to control S. litura on tobacco seedlings in Gujarat, India. They concluded that a combination of biological control agents, insect growth regulators, antifeedants and a trap crop on all sides of the nursery was an ecologically sound procedure for the control of S. litura.

                                                                                                  Different doses of SpltMNPV on final instars of S. litura showed dose-related mortality, but sublethal doses on subsequent generations needs to be considered in the design of baculovirus-based pest management (Mohammad Monobrullah and Umi Shankar, 2008).

                                                                                                  Integrated Pest Management

                                                                                                  In recent years, due to crop failures experienced despite the use of several combinations of chemicals, an integrated approach based on cultural and biocontrol with efficient monitoring using pheromones has been developed (JA Wightman, ICRISAT, Andhra Pradesh, India, personal communication, 1996). The IPM technology that has been developed and implemented in irrigated groundnut where S. litura is endemic has the following components:

                                                                                                  - clean cultivation to expose Spodoptera pupae to natural enemies and weather-related factors

                                                                                                  - sunflower, taro (Zhou, 2009) and castor plants (that attract Spodoptera) to be sown as trap crops both around and within fields

                                                                                                  - pheromone traps to predict Spodoptera egg laying

                                                                                                  - mechanical collection of egg masses and larvae from trap plants on alternate days following the 'warning' from the pheromone traps

                                                                                                  - application of fungicide (chlorothalonil) at the appearance of the first leaf spot lesions, and again after 10 days

                                                                                                  - an application of neem kernel extract during the early stages of crop growth if necessary

                                                                                                  Pongamia glabra oil treatment on tomato plants gave significant reductions on the populations of S. litura while no adverse effects againsts it natural enemies (Marimuthu, 2008)

                                                                                                  - application of nuclear polyhedrosis virus at 500 larval equivalents per hectare in the evening if needed.

                                                                                                  Sahayaraj (2011) gives a summary of different types of plant extracts used by farmer on groundnuts and discusses their effiency.

                                                                                                  Monitoring

                                                                                                  Developments in pheromone technology have made it possible to monitor S. litura in the field, to improve on timing of plant protection measures within groundnut IPM programmes.

                                                                                                  The identification of a male sex pheromone of S. litura, (ZE) 9,11-tetradecadienyl acetate and (ZE) 9,12-tetradecadienyl acetate by Youshima et al. (1974) has enabled effective monitoring of this species for several years. The basic work regarding trap design, height, longevity of the septa, and the potential role of this technology in groundnut has been thoroughly studied at ICRISAT Center, Hyderabad, India over the past decade. These studies have clearly indicated the migratory behaviour of the species in different areas. At present, pheromone technology has given high priority in monitoring for timing of plant protection measures within groundnut IPM programmes. The studies on trap density in groundnut situations indicated no significant differences in moth catches when there were four or more traps per hectare. No decline was noticed in moth catch with increase in trap density. This indirectly suggests a limitation in utilizing the technology in mass trapping operations (Ranga Rao et al., 1989).

                                                                                                  However, there have been some promising results in monitoring the population of moths on Chinese cabbage (Yang Song et al., 2009); spraying times and costs of chemical pesticides against S. litura were significantly reduced by the adoption of sex pheromone trapping.

                                                                                                  Population projections based on life tables and stage-specific consumption rates can reveal the stage structure and damage potential of the pest population of the moths (Tuan et al., 2014). This method could prove to be more reliable as the data obtained by pheromone traps. It is evident that these life tables have to be developed for each area where the moth occurs and one should take into account climate change and yearly temperature and rainfall patterns. It was already established that minimum temperature is the predominant factor that influences pheromone traps whereas wind velocity is predominant in light traps. The overall influence of all the weather factors was high in case of pheromone traps compared to light traps (Prasad et al., 2009).

                                                                                                  Host-Plant Resistance

                                                                                                  The development of resistance to S. litura in suitable groundnut varieties has been regarded as a high priority for Asian groundnut farmers for a number of years. The results of experiments carried out in 1986 and 1987 (data in Wightman and Ranga Rao, 1993) indicated the possibility that ICGV 86031 had some resistance to S. litura combined with high yield in the post-rainy season. This hope was substantiated in further tests on the ICRISAT research farm and in farmers' fields in coastal Andhra Pradesh (southern India). In the limited trials that have been carried out, farmers had sufficient confidence to grow this variety without protecting it with insecticides. They were rewarded with higher yields and lower variable costs than neighbouring farmers who grew locally acceptable varities but applied insecticides to kill defoliators. PI 269116, PI 269118 and PI 262042 had resistance to S. litura, but none were outstanding (Campbell and Wynne, 1980).

                                                                                                  Bioassays carried out with larvae as preliminaries to detect the mechanism of resistance (independent tests by Ranga Rao (ICRISAT) and Padgham (NRI)) revealed no antibiosis effect on second- to sixth-instar larvae when fed mature leaves of ICGV 86031. The main mechanism of resistance is currently thought to be tolerance, manifested as the enhanced ability of vegetative tissue to regrow following defoliation.

                                                                                                  However, first-instar larvae suffered 56% mortality when fed on ICGV 86031 compared with 12% mortality when fed on susceptible ICG 221. Padgham also found that newly hatched larvae had a marked propensity to vacate the leaves of this variety in the first 2 hours of free life. This suggests that the resistance factor which influences the neonates is associated with the leaf surface, because their feeding activity is restricted to scraping the leaf surface. The antixenosis demonstrated by ICGV 86031 is likely to increase the first-instar mortality that is characteristic of r-strategist noctuids (Kyi et al., 1991) and will therefore contribute to the determination of the level of damage caused by the older larvae among which mortality is comparatively low.

                                                                                                  Amin et al. (2011) investigated the morphological and biochemical characteristics of three varieties of cotton and observed their effect on feeding and growth of S. litura. At least one variety was not suited for cotton growers. In a study of the interaction between the virus and the parasitoid, Guo Huifang et al. (2013) showed that the use of an appropriate concentration has the potential to improve the efficiency of the biological control.

                                                                                                  Gaps in Knowledge/Research Needs

                                                                                                  Top of page

                                                                                                  S. litura belongs to a species complex 'Spodoptera' consisting of very similar species with similar ecology in other continents. Correct identification is essential for establishing the correct distribution pattern of this pest.

                                                                                                  References

                                                                                                  Top of page

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