Invasive Species Compendium

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Datasheet

Spodoptera littoralis
(cotton leafworm)

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Datasheet

Spodoptera littoralis (cotton leafworm)

Summary

  • Last modified
  • 11 December 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Spodoptera littoralis
  • Preferred Common Name
  • cotton leafworm
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Larva of S. littoralis, showing light and dark longitudinal bands on sides of the body and dark spots on segments of the dorsal side.
TitleLarva
CaptionLarva of S. littoralis, showing light and dark longitudinal bands on sides of the body and dark spots on segments of the dorsal side.
CopyrightBiologische Bundesanstalt, Dossenheim, Germany
Larva of S. littoralis, showing light and dark longitudinal bands on sides of the body and dark spots on segments of the dorsal side.
LarvaLarva of S. littoralis, showing light and dark longitudinal bands on sides of the body and dark spots on segments of the dorsal side.Biologische Bundesanstalt, Dossenheim, Germany
S. littoralis larva feeding on cotton leaves.
TitleLarva
CaptionS. littoralis larva feeding on cotton leaves.
Copyright©AgrEvo
S. littoralis larva feeding on cotton leaves.
LarvaS. littoralis larva feeding on cotton leaves.©AgrEvo
Adult moth of S. littoralis (museum set specimen).
TitleAdult
CaptionAdult moth of S. littoralis (museum set specimen).
Copyright©Georg Goergen/IITA Insect Museum, Cotonou, Benin
Adult moth of S. littoralis (museum set specimen).
AdultAdult moth of S. littoralis (museum set specimen).©Georg Goergen/IITA Insect Museum, Cotonou, Benin
S. littoralis egg mass on leaf of Rosa spp. from Kenya.
TitleOva
CaptionS. littoralis egg mass on leaf of Rosa spp. from Kenya.
Copyright©Maarten van Merriënboer/PPS, Aalsmeer, Netherlands
S. littoralis egg mass on leaf of Rosa spp. from Kenya.
OvaS. littoralis egg mass on leaf of Rosa spp. from Kenya.©Maarten van Merriënboer/PPS, Aalsmeer, Netherlands
Close-up of S. littoralis egg mass on leaf of Rosa spp. from Kenya.
TitleOva
CaptionClose-up of S. littoralis egg mass on leaf of Rosa spp. from Kenya.
Copyright©Maarten van Merriënboer/PPS, Aalsmeer, Netherlands
Close-up of S. littoralis egg mass on leaf of Rosa spp. from Kenya.
OvaClose-up of S. littoralis egg mass on leaf of Rosa spp. from Kenya.©Maarten van Merriënboer/PPS, Aalsmeer, Netherlands

Identity

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Preferred Scientific Name

  • Spodoptera littoralis (Boisduval)

Preferred Common Name

  • cotton leafworm

Other Scientific Names

  • Hadena littoralis Boisduval
  • Noctua gossypii
  • Prodenia littoralis (Boisduval)
  • Prodenia litura Fabricius sensu auctorum
  • Prodenia retina (Freyer)
  • Prodenia testaceoides Guenee

International Common Names

  • English: Egyptian cotton leafworm; Egyptian cotton worm; leafworm, Egyptian cotton; Mediterranean brocade moth; Mediterranean climbing cutworm; Mediterranean climbing cutworm; tobacco caterpillar; tomato caterpillar
  • Spanish: gusano negro; rosquilla negra
  • French: noctuelle méditerranéenne; ver du coton

Local Common Names

  • Denmark: afrikansk bomuldsugle
  • Finland: krysanteemiyökönnen
  • Germany: Afrikanischer Baumwollwurm
  • Sweden: afrikanskt bomullsfly

EPPO code

  • SPODLI (Spodoptera littoralis)

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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For about 60 years, the Egyptian cotton leafworm was known as Prodenia litura. However, Viette (1962) demonstrated that S. littoralis is a species separate from S. litura. Some authors mistakenly continue to regard the allopatric Old World cotton leafworm species S. littoralis and S. litura as the same species. Generally, however, it is accepted that S. littoralis is found in Africa, Madagascar, Europe and the Middle East, whereas S. litura is found in Asia, Australia and the Pacific Islands.

Description

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Eggs

Spherical, somewhat flattened, 0.6 mm in diameter, laid in clusters arranged in more or less regular rows in one to three layers, with hair scales derived from the tip of the abdomen of the female moth. Usually whitish-yellow in colour, changing to black just prior to hatching, due to the big head of the larva showing through the transparent shell (Pinhey, 1975).


Larvae

Larvae grow to 40-45 mm and are hairless, cylindrical, tapering towards the posterior and variable in colour (blackish-grey to dark green, becoming reddish-brown or whitish-yellow). The sides of the body have 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 the others, interrupting the lateral lines on the first segment. The larva of S. littoralis is figured by Bishari (1934) and Brown and Dewhurst (1975).

Pupae

When newly formed, pupae are green with a reddish colour on the abdomen, turining dark reddish-brown after a few hours. The general shape is cylindrical, 14-20 x 5 mm, tapering towards the posterior segments of the abdomen. The last segment ends in two strong straight hooks (Pinhey, 1975).

Adults

Moth with grey-brown body, 15-20 mm long; wingspan 30-38 mm; forewings grey to reddish brown with paler lines along the veins (in males, bluish areas occur on the wing base and tip); the ocellus is marked by two or three oblique whitish stripes. Hindwings are greyish white, irridescent with grey margins and usually lack darker veins (EPPO, 1997).

Distribution

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The northernly distribution limit of S. littoralis in Europe corresponds to the climatic zone in which winter frosts are infrequent. It occurs throughout Africa and extends eastwards into Turkey and north into eastern Spain, southern France and northern Italy. However, this boundary is probably the extent of migrant activity only because although the pest overwinters in southern Spain, it does not do so in northern Italy or France. In southern Greece, pupae have been observed in the soil after November and the species overwinters in this stage in Crete. Low winter temperatures are therefore an important limiting factor affecting the northerly distribution, especially in a species with no known diapause (Miller, 1976; Sidibe and Lauge, 1977).

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: 15 Dec 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresent
AngolaPresent
BeninPresent
BotswanaPresent
Burkina FasoPresent
BurundiPresent
Cabo VerdePresent
CameroonPresent
Central African RepublicPresent
ChadPresent
ComorosPresent
Congo, Democratic Republic of thePresent
Congo, Republic of thePresent
Côte d'IvoirePresent
EgyptPresent, Widespread
Equatorial GuineaPresent, Localized
EritreaPresent, Localized
EswatiniPresent
EthiopiaPresent
GambiaPresent
GhanaPresent
GuineaPresent
KenyaPresent
LibyaPresent, Widespread
MadagascarPresent
MalawiPresent
MaliPresent
MauritaniaPresent
MauritiusPresent
MoroccoPresent, Widespread
MozambiquePresent
NamibiaPresent
NigerPresent
NigeriaPresent
RéunionPresent
RwandaPresent
Saint HelenaPresent
-AscensionPresent
São Tomé and PríncipePresent
SenegalPresent
SeychellesPresent
Sierra LeonePresent
SomaliaPresent
South AfricaPresent, Widespread
SudanPresent
TanzaniaPresent
TogoPresent
TunisiaPresent, Localized
UgandaPresent
ZambiaPresent
ZimbabwePresent, Widespread

Asia

BahrainPresent
ChinaPresent
-ZhejiangPresent
IndiaPresent
IranPresent, Widespread
IraqPresent
IsraelPresent, Widespread
JordanPresent
LebanonPresent
OmanPresent
PakistanAbsent, Formerly present
Saudi ArabiaPresent
SyriaPresent
TurkeyPresent, LocalizedFirst reported: 197*
United Arab EmiratesPresent
YemenPresent, Localized

Europe

AustriaAbsent
BelgiumAbsent
CroatiaAbsent, Confirmed absent by survey
CyprusPresent, Widespread
DenmarkAbsent, Intercepted only
FinlandAbsent, Eradicated
FrancePresent, Few occurrences
GermanyAbsent, Eradicated
GreecePresent, Localized
-CretePresent
ItalyPresent, Localized1968
-SicilyPresent
MaltaPresent, Widespread
NetherlandsAbsent, EradicatedAbsent, pest eradicated, confirmed by survey. Based on long-term annual surveys, 362 survey observations in 2012.
PortugalPresent, Localized
-AzoresPresent
-MadeiraPresent
SloveniaAbsent
SpainPresent, Localized
-Balearic IslandsPresent, Localized
-Canary IslandsPresent
SwedenAbsent, Formerly present
SwitzerlandAbsent, Formerly present
United KingdomAbsent, Eradicated
-EnglandAbsent, Formerly present

Risk of Introduction

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EPPO has listed S. littoralis as an A2 quarantine pest (OEPP/EPPO, 1981). CPPC, NAPPO and OIRSA also consider it to be of quarantine significance. It is a potential pest of areas where the average annual minimal temperature is not below -10°C. S. littoralis is already fairly widespread in Mediterranean countries and does not present a phytosanitary risk there. The most significant phytosanitary risk for S. littoralis is the possible introduction into glasshouses in most parts of Europe, where it could damage many ornamental and vegetable crops. Although control with insecticides is possible, there have been many cases of resistance and the lack of available biological control methods means that introduction of S. littoralis into glasshouses could necessitate insecticide treatments that could interfere with existing biological control of other pests (EPPO, 1997).

S. littoralis first appeared in UK glasshouses in considerable numbers in 1963. It was found that the eggs were being introduced on imported cuttings, especially chrysanthemums and carnations.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial

Hosts/Species Affected

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The host range of S. littoralis covers over 40 families, containing at least 87 species of economic importance (Salama et al., 1970).

In many of the published reports of host plants, it is difficult to distinguish between S. littoralis and S. litura but the tabular data refers entirely to records from the distribution area of the former. Both species 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 nilotica (gum arabic tree)FabaceaeOther
    Actinidia arguta (tara vine)ActinidiaceaeOther
      Alcea rosea (Hollyhock)MalvaceaeOther
        Allium ampeloprasum (wild leek)LiliaceaeUnknown
        Allium cepa (onion)LiliaceaeMain
        Allium fistulosum (Welsh onion)LiliaceaeOther
          Amaranthus (amaranth)AmaranthaceaeMain
          Anemone (windflower)RanunculaceaeOther
            Antirrhinum majus (snapdragon)ScrophulariaceaeWild host
              Apium graveolens (celery)ApiaceaeOther
                Arachis hypogaea (groundnut)FabaceaeMain
                Asparagus officinalis (asparagus)LiliaceaeOther
                  Beta vulgaris (beetroot)ChenopodiaceaeMain
                    Beta vulgaris var. ciclaChenopodiaceaeUnknown
                    Beta vulgaris var. saccharifera (sugarbeet)ChenopodiaceaeMain
                      Brassica oleracea (cabbages, cauliflowers)BrassicaceaeMain
                      Brassica oleracea var. capitata (cabbage)BrassicaceaeOther
                        Brassica rapa subsp. chinensis (Chinese cabbage)BrassicaceaeMain
                          Brassica rapa subsp. pekinensisBrassicaceaeOther
                            Brassicaceae (cruciferous crops)BrassicaceaeMain
                              CaladiumAraceaeOther
                                Callistephus chinensis (China aster)AsteraceaeOther
                                  Camellia sinensis (tea)TheaceaeMain
                                    CannaCannaceaeOther
                                      Capsicum (peppers)SolanaceaeOther
                                        Capsicum annuum (bell pepper)SolanaceaeMain
                                        Casuarina equisetifolia (casuarina)CasuarinaceaeWild host
                                          Chloris gayana (Rhodes grass)PoaceaeOther
                                            Chrysanthemum indicum (chrysanthemum)AsteraceaeMain
                                              Citrullus lanatus (watermelon)CucurbitaceaeMain
                                                CitrusRutaceaeMain
                                                  Citrus aurantium (sour orange)RutaceaeMain
                                                    Coffea arabica (arabica coffee)RubiaceaeMain
                                                      Convolvulus (morning glory)ConvolvulaceaeOther
                                                        Corchorus capsularis (white jute)TiliaceaeOther
                                                          Corchorus olitorius (jute)TiliaceaeMain
                                                            CryptomeriaTaxodiaceaeOther
                                                              Cucurbita (pumpkin)CucurbitaceaeMain
                                                                Cucurbita pepo (marrow)CucurbitaceaeMain
                                                                  Cynara cardunculus var. scolymus (globe artichoke)AsteraceaeMain
                                                                  Dalbergia sissooFabaceaeOther
                                                                    Datura (thorn-apple)SolanaceaeOther
                                                                      Daucus carota (carrot)ApiaceaeMain
                                                                        Dianthus barbatus (sweet williams)CaryophyllaceaeOther
                                                                          Dianthus caryophyllus (carnation)CaryophyllaceaeMain
                                                                            Eucalyptus globulus (Tasmanian blue gum)MyrtaceaeWild host
                                                                              EuphorbiaceaeEuphorbiaceaeOther
                                                                                Fabaceae (leguminous plants)FabaceaeMain
                                                                                  Ficus carica (common fig)MoraceaeMain
                                                                                  Ficus sycomorus (sycamore fig)MoraceaeUnknown
                                                                                  Fragaria vesca (wild strawberry)RosaceaeWild host
                                                                                    Gerbera (Barbeton daisy)AsteraceaeMain
                                                                                      Gladiolus hybrids (sword lily)IridaceaeOther
                                                                                        GlycineFabaceaeUnknown
                                                                                        Glycine max (soyabean)FabaceaeMain
                                                                                          Gossypium (cotton)MalvaceaeMain
                                                                                          Gossypium barbadense (Gallini cotton)MalvaceaeMain
                                                                                            Gossypium hirsutum (Bourbon cotton)MalvaceaeUnknown
                                                                                            Guizotia abyssinica (niger)AsteraceaeWild host
                                                                                              Helianthus annuus (sunflower)AsteraceaeMain
                                                                                                Helianthus tuberosus (Jerusalem artichoke)AsteraceaeMain
                                                                                                  Hibiscus cannabinus (kenaf)MalvaceaeOther
                                                                                                    Hibiscus mutabilis (cottonrose)MalvaceaeOther
                                                                                                      Indigofera tinctoria (true indigo)FabaceaeOther
                                                                                                        Ipomoea batatas (sweet potato)ConvolvulaceaeMain
                                                                                                          Jatropha curcas (jatropha)EuphorbiaceaeWild host
                                                                                                            Lactuca sativa (lettuce)AsteraceaeMain
                                                                                                              LantanaVerbenaceaeOther
                                                                                                                Luffa aegyptiaca (loofah)CucurbitaceaeOther
                                                                                                                  LycopersiconSolanaceaeOther
                                                                                                                    Malus sylvestris (crab-apple tree)RosaceaeWild host
                                                                                                                      Medicago sativa (lucerne)FabaceaeMain
                                                                                                                      Melilotus spp.FabaceaeOther
                                                                                                                        Mentha spicata (Spear mint)LamiaceaeOther
                                                                                                                          Monstera deliciosa (ceriman)AraceaeWild host
                                                                                                                            Morus (mulberrytree)MoraceaeMain
                                                                                                                              Morus alba (mora)MoraceaeUnknown
                                                                                                                              Musa (banana)MusaceaeOther
                                                                                                                                Musa x paradisiaca (plantain)MusaceaeMain
                                                                                                                                  Nicandra physalodes (apple of Peru)SolanaceaeWild host
                                                                                                                                    Nicotiana tabacum (tobacco)SolanaceaeMain
                                                                                                                                      Opuntia (Pricklypear)CactaceaeOther
                                                                                                                                        Oryza sativa (rice)PoaceaeMain
                                                                                                                                          Persea americana (avocado)LauraceaeMain
                                                                                                                                          Petroselinum crispum (parsley)ApiaceaeUnknown
                                                                                                                                          Phaseolus (beans)FabaceaeMain
                                                                                                                                            Phaseolus vulgaris (common bean)FabaceaeMain
                                                                                                                                            Phoenix dactylifera (date-palm)ArecaceaeOther
                                                                                                                                              Piper (pepper)PiperaceaeOther
                                                                                                                                                Pistia stratiotes (water lettuce)AraceaeOther
                                                                                                                                                  Pisum sativum (pea)FabaceaeMain
                                                                                                                                                    Poaceae (grasses)PoaceaeMain
                                                                                                                                                      Polyphagous (polyphagous)Main
                                                                                                                                                        Populus alba (silver-leaf poplar)SalicaceaeOther
                                                                                                                                                          Portulaca oleracea (purslane)PortulacaceaeOther
                                                                                                                                                            Prunus domestica (plum)RosaceaeMain
                                                                                                                                                              Prunus salicina (Japanese plum)RosaceaeOther
                                                                                                                                                                Psidium guajava (guava)MyrtaceaeMain
                                                                                                                                                                  Punica granatum (pomegranate)PunicaceaeMain
                                                                                                                                                                    Quercus petraea (durmast oak)FagaceaeWild host
                                                                                                                                                                      Raphanus sativus (radish)BrassicaceaeMain
                                                                                                                                                                        Ricinus communis (castor bean)EuphorbiaceaeMain
                                                                                                                                                                        Rosa (roses)RosaceaeMain
                                                                                                                                                                          Saccharum officinarum (sugarcane)PoaceaeMain
                                                                                                                                                                            Salvia officinalis (common sage)LamiaceaeOther
                                                                                                                                                                              Senecio (Groundsel)AsteraceaeWild host
                                                                                                                                                                                Sesamum indicum (sesame)PedaliaceaeOther
                                                                                                                                                                                  Sesbania sesban (sesban)FabaceaeOther
                                                                                                                                                                                    Solanum lycopersicum (tomato)SolanaceaeMain
                                                                                                                                                                                    Solanum melongena (aubergine)SolanaceaeMain
                                                                                                                                                                                      Solanum tuberosum (potato)SolanaceaeMain
                                                                                                                                                                                        Sorghum bicolor (sorghum)PoaceaeMain
                                                                                                                                                                                        Spinacia oleracea (spinach)ChenopodiaceaeMain
                                                                                                                                                                                          Tectona grandis (teak)LamiaceaeWild host
                                                                                                                                                                                            Theobroma cacao (cocoa)MalvaceaeMain
                                                                                                                                                                                              Trifolium (clovers)FabaceaeOther
                                                                                                                                                                                                Trifolium alexandrinum (Berseem clover)FabaceaeOther
                                                                                                                                                                                                  Trifolium repens (white clover)FabaceaeOther
                                                                                                                                                                                                    Trifolium spp.FabaceaeOther
                                                                                                                                                                                                      Trigonella foenum-graecum (fenugreek)FabaceaeOther
                                                                                                                                                                                                        Triticum aestivum (wheat)PoaceaeMain
                                                                                                                                                                                                          Verbena (vervain)VerbenaceaeOther
                                                                                                                                                                                                            Vicia faba (faba bean)FabaceaeMain
                                                                                                                                                                                                              Vigna angularis (adzuki bean)FabaceaeOther
                                                                                                                                                                                                                Vigna mungo (black gram)FabaceaeMain
                                                                                                                                                                                                                  Vigna radiata (mung bean)FabaceaeMain
                                                                                                                                                                                                                    Vigna unguiculata (cowpea)FabaceaeMain
                                                                                                                                                                                                                    Viola odorata (English violet)ViolaceaeOther
                                                                                                                                                                                                                      Vitis (grape)VitaceaeUnknown
                                                                                                                                                                                                                      Vitis vinifera (grapevine)VitaceaeMain
                                                                                                                                                                                                                        Zea mays (maize)PoaceaeMain
                                                                                                                                                                                                                        Zinnia elegans (zinnia)AsteraceaeOther

                                                                                                                                                                                                                          Growth Stages

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                                                                                                                                                                                                                          Flowering stage, Fruiting stage, Seedling 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, the larvae feed on the leaves creating large holes of irregular shape and usually all that remains are the bigger veins. The larvae may also bore into the bud or young boll and consume the whole contents, causing them to be shed or dry up (Bishari, 1934). Bolls have large holes in them from which yellowish- to dark-green larval excrement protrudes. On tobacco, leaves develop irregular, brownish-red patches and the stem base may be gnawed off. Maize stems are often mined by S. littoralis and young grains in the ear may also be damaged.

                                                                                                                                                                                                                          List of Symptoms/Signs

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                                                                                                                                                                                                                          SignLife StagesType
                                                                                                                                                                                                                          Fruit / frass visible
                                                                                                                                                                                                                          Fruit / internal feeding
                                                                                                                                                                                                                          Fruit / obvious exit hole
                                                                                                                                                                                                                          Fruit / premature drop
                                                                                                                                                                                                                          Leaves / external feeding
                                                                                                                                                                                                                          Leaves / shredding

                                                                                                                                                                                                                          Biology and Ecology

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                                                                                                                                                                                                                          Female moths lay most of their egg masses (20-1000 eggs) on the lower surface of younger leaves or upper parts of the plant (Khalifa et al., 1982). On cotton, the first three larval instars feed mainly on the lower surface of the leaves, whereas later instars feed on both surfaces. The larvae feed mainly in the dark, although this behaviour pattern may be less noticeable in early instars (Hassan et al., 1960). Pinhey (1975) recorded that >50% of the nocturnal larval population consisted of early instar larvae. In summer the majority of fifth- and sixth-instar larvae leave the plants during mid-morning until sunset, returning to climb the plant at night (Baker and Miller, 1974). Third- and fourth-instars rest on the plant and remain stationary unless overcrowded.

                                                                                                                                                                                                                          On pupation the fully grown larva pushes the loose surface of the soil downwards until it reaches more solid ground 3-5 cm deep. It then creates a clay 'cell' or cocoon in which it usually pupates within 5-6 hours (Pinhey, 1975).

                                                                                                                                                                                                                          Emergence of adult moths occurs at night and they have a life span of 5-10 days (Shalama and Shoukry, 1972). The reproductive capacity, egg facility and life span of moths are affected by the difference in ages between males and females. The highest ratio of egg fertility was obtained by mating between 4-day-old males with fresh females (Nasr and Nassif, 1978). There is also a correlation between the host plant and the longevity and fecundity of S. littoralis (Dimetry and Nadia, 1972). The majority of adults mate on the first night of emergence, copulation lasting for 20 minutes to 2 hours. Approximately 50% of mated females lay their eggs on the same night of mating, before sunrise (Hassan et al., 1960). Adults fly at night, mostly between 20.00 and midnight (Nasr et al., 1981). Flight activity is governed by atmospheric conditions, increases in relative humidity and decreases in air temperature inducing flight (Hassan et al., 1960). The flight range during a 4-hour-period can be up to 1.5 km (Salama and Shoukry, 1972).

                                                                                                                                                                                                                          The moths have chemoreceptors on the ventral surface of the tarsi and the distal portion of the proboscis. These are highly sensitive and respond to a certain number of sugars mainly present in nectar. Pheromones (comprising of tetradecadien-1-ol acetates) have been isolated and successfully used in traps (Kehat and Gordon, 1975; Campion, 1977).

                                                                                                                                                                                                                          The minimum constant temperature for normal development in all stages is 13-14°C. Resistance to cold generally increases through the larval stages and is greatest in the pupal stage (Miller, 1977). At 18°C, egg, larval and pupal stages last 9, 34 and 27 days, respectively. At 36°C, egg, larval and pupal stages last 2, 10 and 8 days, respectively. Data on survival and development at different temperatures are provided by Sidibe and Lauge (1977), Baker and Miller (1974) and Ocete Rubio (1984). Information on development on different host crops is given by Dimetry and Nadia (1972), Hirakly and Bishara (1974), Abdal-Fattah (1977), Zoebelein (1977) and Badr et al. (1983). Studies in Egypt indicate that there are seven overlapping generations of S. littoralis when feeding on cotton, and that there are three peak infestation periods (El-Shafei et al., 1981; Khalifa et al., 1982).

                                                                                                                                                                                                                          Natural enemies

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                                                                                                                                                                                                                          Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                                                                                                                                                                                          Andrallus spinidens Predator
                                                                                                                                                                                                                          Apanteles colemani Parasite Larvae
                                                                                                                                                                                                                          Apanteles prodeniae Parasite Larvae
                                                                                                                                                                                                                          Apanteles vitripennis Parasite Larvae
                                                                                                                                                                                                                          Aspergillus flavus Antagonist
                                                                                                                                                                                                                          Bacillus cereus Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis aizawai Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis alesti Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis colmeri Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis entomocidus Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis galleriae Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis kenyae Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis kurstaki Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis sotto Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis subsp. dendrolimus Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis subtoxicus Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis thuringiensis Pathogen Larvae
                                                                                                                                                                                                                          Bacillus thuringiensis tolworthi Pathogen Larvae
                                                                                                                                                                                                                          Barylypa humeralis Parasite
                                                                                                                                                                                                                          Beauveria bassiana Pathogen
                                                                                                                                                                                                                          Bessa remota Parasite Larvae
                                                                                                                                                                                                                          Blaptostethus piceus Predator
                                                                                                                                                                                                                          Blepharella lateralis Parasite Larvae
                                                                                                                                                                                                                          Borrelinavirus litura Pathogen
                                                                                                                                                                                                                          Brachymeria excarinata Parasite
                                                                                                                                                                                                                          Brachymeria lasus Parasite
                                                                                                                                                                                                                          Bracon brevicornis Parasite Larvae
                                                                                                                                                                                                                          Bracon hebetor Parasite Larvae Israel
                                                                                                                                                                                                                          Brinckochrysa scelestes Predator Karnataka
                                                                                                                                                                                                                          Calosoma blaptoides Predator Larvae Guam
                                                                                                                                                                                                                          Calosoma chlorostictum Predator Larvae
                                                                                                                                                                                                                          Campoletis chlorideae Parasite Larvae Maharashtra
                                                                                                                                                                                                                          Cantheconidia furcellata Predator
                                                                                                                                                                                                                          Charops obtusus Parasite
                                                                                                                                                                                                                          Cheiracanthium mildei Predator
                                                                                                                                                                                                                          Chelonus aegyptia Parasite Larvae
                                                                                                                                                                                                                          Chelonus curvimaculatus Parasite Larvae
                                                                                                                                                                                                                          Chelonus formosanus Parasite Larvae
                                                                                                                                                                                                                          Chelonus inanitus Parasite Eggs/Larvae Egypt; Israel cabbages; polyphagous
                                                                                                                                                                                                                          Chelonus insularis Parasite Larvae Egypt; India clovers; polyphagous
                                                                                                                                                                                                                          Chelonus oculator Parasite Özkan and Özmen (2001)
                                                                                                                                                                                                                          Chrysoperla carnea Predator
                                                                                                                                                                                                                          Coccinella undecimpunctata Predator
                                                                                                                                                                                                                          Compsilura concinnata Parasite Larvae
                                                                                                                                                                                                                          Copidosoma maculata Parasite
                                                                                                                                                                                                                          Cotesia marginiventris Parasite Larvae Egypt clovers
                                                                                                                                                                                                                          Cotesia ruficrus Parasite Larvae
                                                                                                                                                                                                                          Cotesia telengai Parasite Larvae
                                                                                                                                                                                                                          Cybocephalus micans Predator
                                                                                                                                                                                                                          cytoplasmic polyhedrosis viruses Pathogen Larvae
                                                                                                                                                                                                                          Damaster blaptoides Predator Larvae
                                                                                                                                                                                                                          Drino imberbis Parasite Larvae
                                                                                                                                                                                                                          Eocanthecona furcellata Predator
                                                                                                                                                                                                                          Eriborus argenteopilosus Parasite
                                                                                                                                                                                                                          Euborellia annulipes Predator
                                                                                                                                                                                                                          Euplectrus laphygmae Parasite
                                                                                                                                                                                                                          Euplectrus platyhypenae Parasite Pakistan
                                                                                                                                                                                                                          Exorista larvarum Parasite Larvae
                                                                                                                                                                                                                          Glyptapanteles africanus Parasite Larvae Gujarat
                                                                                                                                                                                                                          Glyptapanteles ashmeadi Parasite Larvae
                                                                                                                                                                                                                          Gnathonarium exsciccatum Predator
                                                                                                                                                                                                                          Granulosis virus Pathogen
                                                                                                                                                                                                                          Harpactor costalis Predator
                                                                                                                                                                                                                          Heterorhabditis heliothidis Parasite
                                                                                                                                                                                                                          Heterorhabditis indicus Parasite
                                                                                                                                                                                                                          Homolobus truncatoides Parasite Larvae
                                                                                                                                                                                                                          Hyposoter didymator Parasite Larvae Cape Verde Cajanus cajan; tomatoes
                                                                                                                                                                                                                          Hyposoter exiguae Parasite Egypt clovers
                                                                                                                                                                                                                          Labidura riparia Predator
                                                                                                                                                                                                                          Lasiochalcidia erythropus Parasite
                                                                                                                                                                                                                          Lecanicillium lecanii Pathogen
                                                                                                                                                                                                                          Lespesia archippivora Parasite Larvae Guam
                                                                                                                                                                                                                          Mallada boninensis Predator
                                                                                                                                                                                                                          Metarhizium anisopliae Pathogen Karnataka
                                                                                                                                                                                                                          Metaseiulus occidentalis Predator
                                                                                                                                                                                                                          Meteorus gyrator Parasite Larvae Egypt cabbages; Corchorus olitorius; Vigna unguiculata
                                                                                                                                                                                                                          Meteorus pulchricornis Parasite Larvae
                                                                                                                                                                                                                          Meteorus rubens Parasite Larvae Israel
                                                                                                                                                                                                                          Microchelonus blackburni Parasite Larvae
                                                                                                                                                                                                                          Microchelonus heliopae Parasite Larvae Gujarat
                                                                                                                                                                                                                          Microplitis demolitor Parasite Larvae Egypt clovers
                                                                                                                                                                                                                          Microplitis manilae Parasite Larvae
                                                                                                                                                                                                                          Microplitis rufiventris Parasite Larvae Egypt; Israel
                                                                                                                                                                                                                          Nemorilla maculosa Parasite Larvae
                                                                                                                                                                                                                          Nesidiocoris tenuis Predator
                                                                                                                                                                                                                          Nomuraea rileyi Pathogen Larvae
                                                                                                                                                                                                                          Nosema infuscatellus Pathogen
                                                                                                                                                                                                                          Nosema liturae Pathogen Guangzhou
                                                                                                                                                                                                                          Nosema mesnili Pathogen
                                                                                                                                                                                                                          Nucleopolyhedrosis virus Pathogen
                                                                                                                                                                                                                          Oncocephalus annulipes Predator
                                                                                                                                                                                                                          Orius albidipennis Predator
                                                                                                                                                                                                                          Orius laevigatus Predator
                                                                                                                                                                                                                          Ovomermis albicans Parasite
                                                                                                                                                                                                                          Paederus alfierii Predator
                                                                                                                                                                                                                          Palexorista laxa Parasite Larvae
                                                                                                                                                                                                                          Peribaea orbata Parasite Larvae Egypt; New Caledonia cabbages; clovers; Corchorus olitorius; Malva pariflora; polyphagous
                                                                                                                                                                                                                          Peribaea palaestina Parasite Larvae
                                                                                                                                                                                                                          Philonthus stragulatus Predator
                                                                                                                                                                                                                          Pimpla hypochondriaca Parasite
                                                                                                                                                                                                                          Podisus maculiventris Predator
                                                                                                                                                                                                                          Podisus sagitta Predator
                                                                                                                                                                                                                          Polistes chinensis chinensis Predator
                                                                                                                                                                                                                          Polistes jadwigae Predator
                                                                                                                                                                                                                          Pseudomonas aeruginosa Pathogen
                                                                                                                                                                                                                          Rhynocoris marginatus Predator
                                                                                                                                                                                                                          Sarcophaga misera Parasite
                                                                                                                                                                                                                          Scymnus interruptus Predator
                                                                                                                                                                                                                          Scymnus syriacus Predator
                                                                                                                                                                                                                          Serratia marcescens Pathogen
                                                                                                                                                                                                                          Spodophagus lepidopterae Parasite
                                                                                                                                                                                                                          Steinernema carpocapsae Parasite
                                                                                                                                                                                                                          Steinernema feltiae Parasite
                                                                                                                                                                                                                          Steinernema glaseri Parasite
                                                                                                                                                                                                                          Telenomus nawaii Parasite Eggs Guam; Samoa bananas; polyphagous
                                                                                                                                                                                                                          Telenomus remus Parasite Eggs Australia; Cape Verde; Guam; Gujarat; India; Israel; Karnataka; New Zealand; Pakistan Cajanus cajan; polyphagous; tobacco; tomatoes
                                                                                                                                                                                                                          Telenomus spodopterae Parasite Eggs Egypt clovers
                                                                                                                                                                                                                          Tetrastichus howardi Parasite Karnataka
                                                                                                                                                                                                                          Tetrastichus israeli Parasite
                                                                                                                                                                                                                          Trichogramma buesi Parasite Eggs
                                                                                                                                                                                                                          Trichogramma cacoeciae Parasite Eggs
                                                                                                                                                                                                                          Trichogramma chilonis Parasite Eggs Karnataka
                                                                                                                                                                                                                          Trichogramma evanescens Parasite Eggs
                                                                                                                                                                                                                          Trichogramma perkinsi Parasite Eggs
                                                                                                                                                                                                                          Trichogramma rhenana Parasite Eggs
                                                                                                                                                                                                                          Trichogrammatoidea armigera Parasite Eggs
                                                                                                                                                                                                                          Trichogrammatoidea australicum Parasite Eggs
                                                                                                                                                                                                                          Trichogrammatoidea lutea Parasite Eggs
                                                                                                                                                                                                                          Trichospilus pupivora Parasite
                                                                                                                                                                                                                          Ummeliata insecticeps Predator
                                                                                                                                                                                                                          Vairimorpha necatrix Pathogen
                                                                                                                                                                                                                          Xenorhabdus nematophilus Pathogen
                                                                                                                                                                                                                          Zele chlorophthalma Parasite Larvae Egypt cabbages; Corchorus olitorius
                                                                                                                                                                                                                          Zele nigricornis Parasite Larvae

                                                                                                                                                                                                                          Notes on Natural Enemies

                                                                                                                                                                                                                          Top of page
                                                                                                                                                                                                                          Natural enemies that have been observed in the field (Hegazi et al., 1977). Delvare and Rasplus (1994) described the pteromalid parasitoid, Spodophagus lepidopterae, from S. littoralis from Madagascar. Most work on the natural enemies of S. littoralis has been done in Egypt and Spain.

                                                                                                                                                                                                                          Generalist predators of S. littoralis include ladybirds, which feed on egg masses and young larvae, Paedeus fuscipes (staphylinid rover beetle), Orius albidipennis, Labidura riparia, Creontiades pallidus, Calosoma chlorostictum and Polistes gallicus.

                                                                                                                                                                                                                          Impact

                                                                                                                                                                                                                          Top of page
                                                                                                                                                                                                                          S. littoralis is one of the most destructive agricultural lepidopterous pests within its subtropical and tropical range. It can attack numerous economically important crops throughout the year (EPPO, 1997). On cotton, the pest may cause considerable damage by feeding on the leaves, fruiting points, flower buds and occasionally on bolls. When groundnuts are infested, larvae first select young folded leaves for feeding, but in severe attacks, leaves of any age are stripped off. Sometimes, even the ripening kernels in the pods in the soil may be attacked. Pods of cowpeas and the seeds they contain are also often badly damaged. In tomatoes, larvae bore into the fruit, which is thus rendered unsuitable for consumption. Numerous other crops are attacked, mainly on their leaves.

                                                                                                                                                                                                                          In Europe, damage caused by S. littoralis was minimal until about 1937. In 1949, there was a catastrophic population explosion in southern Spain, which affected lucerne, potatoes and other vegetable crops. At present, this noctuid is of great economic importance in Cyprus, Israel, Malta, Morocco and Spain (except the north). In Italy, it is especially important on protected crops of ornamentals and vegetables (Inserra and Calabretta, 1985; Nucifora, 1985). In Greece, S. littoralis causes slight damage in Crete on lucerne
                                                                                                                                                                                                                          and clover only.

                                                                                                                                                                                                                          In North Africa, tomato, Capsicum, cotton, maize and other vegetables are affected. In Egypt, it is one of the most serious cotton pests.

                                                                                                                                                                                                                          Detection and Inspection

                                                                                                                                                                                                                          Top of page
                                                                                                                                                                                                                          Pheromeone traps can be used to monitor the incidence of S. littoralis (Rizk et al., 1990). Kehat and Dunkelblum (1993) found that the minor sex pheromone component, (9Z,12Z)-9,12-tetradecadienyl acetate in addition to the major component (9Z,11Z)-9,11-tetradecadienyl acetate was required for in order to attract males.

                                                                                                                                                                                                                          Similarities to Other Species/Conditions

                                                                                                                                                                                                                          Top of page

                                                                                                                                                                                                                          S. littoralis is often confused with S. litura, and the variability and similarity of the two species makes correct identification difficult and examination of adult genitalia is often the only certain method. For more information on morphological discrimination between the adult, pupal and larval stages of the two species, refer to Schmutterer (1969), Cayrol (1972), Mochida (1973) and Brown and Dewhurst (1975).

                                                                                                                                                                                                                          Although markings on larvae are variable, a bright-yellow stripe along the length of the dorsal surface is characteristic of S. litura.

                                                                                                                                                                                                                          On dissection of the genitalia, the ductus and ostium bursae are the same length in female S. littoralis, whereas they are different lengths in S. litura. The shape of the juxta in males in both species is very characteristic, and the ornamentation of the aedeagus vesica is also diagnostic.

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

                                                                                                                                                                                                                          Prevention and Control

                                                                                                                                                                                                                          Top of page

                                                                                                                                                                                                                          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.

                                                                                                                                                                                                                          Biological Control

                                                                                                                                                                                                                          Numerous studies have been carried out on possible biological control of S. littoralis. Parasitoids (braconids, encyrtids, tachinids and ichneumonids) and predators have been extensively documented. A nuclear polyhedrosis virus has been evaluated against S. litura (Elnagar and El-Sheikh, 1990; Jones et al., 1994), whereas fungi and microsporidia have also been recorded as pathogens. Parasitic nematodes such as Neoaplectana carpocapsae have also been evaluated. However, direct use of these biocontrol agents has not been commercailized. Treatment with Bacillus thuringiensis has been used (Navon et al., 1983), but only some strains are effective as S. littoralis is resistant to many strains (Salama et al., 1989).

                                                                                                                                                                                                                          Chemical Control

                                                                                                                                                                                                                          The chemical control of S. littoralis has been extensively reported, especially in relation to cotton in Egypt. Numerous organophosphorus, synthetic pyrethroids and other insecticides have been used, with appearance of resistance and cross resistance in many cases (Issa et al., 1984a; 1984b; Abo-El-Ghar et al., 1986). However, compulsory limitation of the application of synthetic pyrethroids to one per year on cotton in Egypt has stopped the appearance of new resistance (Sawicki, 1986).

                                                                                                                                                                                                                          Chemicals used against species of Spodoptera also include insect growth regulators. There is interest, especially in India, in various antifeedant compounds or extracts, and in natural products, such as azadirachtin and neem extracts.

                                                                                                                                                                                                                          IPM

                                                                                                                                                                                                                          Integrated pest management techniques, favouring beneficial arthropods, are applied against S. littoralis on cotton in Egypt. These involve hand collection of egg masses, use of microbial pesticides and insect growth regulators and slow-release pheromone formulations for mating disruption. If these measures are taken, relatively few applications of conventional insecticides are necessary (Campion and Nesbitt, 1982; Hosny et al., 1983; Campion and Hosny, 1987). Damage thresholds have been established by Hosny et al. (1986). Pheromones have also been used for mass trapping using a lure and kill strategy (McVeigh and Bettany, 1987) and for monitoring populations. Souka (1980) experimented with irradiation for sterile-insect release, but this technique has not been widely applied in the field.

                                                                                                                                                                                                                          Phytosanitary Measures

                                                                                                                                                                                                                          For planting material, EPPO recommends (OEPP/EPPO, 1990) absence of the pests from the place of production during the last 3 months, or treatment of the consignment. For cut flowers, pre-export inspection is considered sufficient.

                                                                                                                                                                                                                          Cold storage of chrysanthemum and carnation cuttings for at least 10 days at a temperature not exceeding 1.7°C will kill all stages of S. littoralis, but may damage the plants. Storage at slightly higher temperatures or shorter durations does not eradicate S. littoralis, but differences in response to cold have been observed both between strains and within developmental stages of the pest (Powell and Gostick, 1971; Miller, 1976). Irradiation has been investigated as a treatment for cut flowers (Navon et al., 1988). For cut chrysanthemum flowers, Wang and Lin (1984) suggest enclosing buds in perforated polythene bags to exclude the pest and dipping the cut stems in insecticide solutions.
                                                                                                                                                                                                                           

                                                                                                                                                                                                                          References

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                                                                                                                                                                                                                          Nasr ESA, El-Rafie K, Hosny MM, Badawi A, 1973. Effect of temperature and relative humidity on the life-cycle of the cotton leafworm Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae). Bulletin de la Societe Entomologique D'Egypte, 57:139-144

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                                                                                                                                                                                                                          Sadek, M. M., 2011. Complementary behaviors of maternal and offspring Spodoptera littoralis: oviposition site selection and larval movement together maximize performance. Journal of Insect Behavior, 24(1), 67-82. doi: 10.1007/s10905-010-9238-4

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                                                                                                                                                                                                                          Schmutterer, H., 1971. Contribution to the knowledge of the crop pest fauna in Ethiopia. Zeitschrift fur Angewandte Entomologie, 67(4), 371-389.

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                                                                                                                                                                                                                          GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
                                                                                                                                                                                                                          Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

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