Spodoptera litura (taro caterpillar)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Air Temperature
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Wood Packaging
- Impact Summary
- Risk and Impact Factors
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Spodoptera litura (Fabricius)
Preferred Common Name
- taro caterpillar
Other Scientific Names
- Mamestra albisparsa Walker
- Noctua elata Fabricius
- Noctua histrionica Fabricius
- Noctua litura Fabricius
- Prodenia ciligera Guenée
- Prodenia declinata Walker
- Prodenia evanescens Butler
- Prodenia glaucistriga Walker
- Prodenia littoralis Fabricius
- Prodenia litura Fabricius
- Prodenia subterminalis Walker
- Prodenia tasmanica Guenée
- Prodenia testaceoides Walker
- Spodoptera littoralis
International Common Names
- English: armyworm; cluster caterpillar; common cutworm; cotton leafworm; cotton worm; Egyptian cotton leafworm; rice cutworm; tobacco budworm; tobacco caterpillar; tobacco cutworm; tobacco leaf caterpillar
- Spanish: gusano del tabaco; gusano negro; lagarta; rosquilla negra
- French: chenille defoliante; noctuelle rayee; ver du coton; ver du tabac
Local Common Names
- Germany: Aegyptische Baumwollraupe; Asiatischer Baumwollwurm; Baumwollblattraupe; Baumwollraupe, Aegyptische; Baumwollwurm, Asiatischer
- India: ladde purugu; telugu
- Israel: haprodenia
- Italy: larva bruna con macchie vellutate
- Japan: hasumon-yoto
- Netherlands: eiernestrups
- Turkey: pamuk yaprak kurdu
- PRODLI (Spodoptera litura)
Summary of InvasivenessTop of page
The tobacco caterpillar,S. litura, is one of the most important insect pests of agricultural crops in the Asian tropics. It is widely distributed throughout tropical and temperate Asia, Australasia and the Pacific Islands (Feakin, 1973; Kranz et al., 1977). Records of S. litura having limited distribution in (or being eradicated from) Germany, Russian Federation, Russian Far East, the UK and Réunion may in fact refer to S. littoralis. Both S. litura and S. littoralis are totally polyphagous (Brown and Dewhurst, 1975; Holloway, 1989) and therefore have huge potential to invade new areas and/or to adapt to new climatic and/or ecological situations. The Spodoptera group consists of closely related species with similar ecology that are difficult to identify to species level.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Lepidoptera
- Family: Noctuidae
- Genus: Spodoptera
- Species: Spodoptera litura
Notes on Taxonomy and NomenclatureTop of page
The two Old World cotton leafworm species, Spodoptera litura and S. littoralis, are allopatric, their ranges covering Asia and Africa, Europe and the Middle East, respectively. Many authors have regarded them as the same species.
DescriptionTop of page
The eggs are spherical, somewhat flattened, and 0.6 mm in diameter. They are usually pale orange-brown or pink in colour, laid in batches and covered with hair scales from the tip of the abdomen of the female moth. Egg masses measure about 4-7 mm in diameter and appear golden brown because they are covered with body scales of females.
The larva is hairless, variable in colour (young larvae are light green, the later instars are dark green to brown on their backs, lighter underneath); sides of body with dark and light longitudinal bands; dorsal side with two dark semilunar spots laterally on each segment, except for the prothorax; spots on the first and eighth abdominal segments larger than others, interrupting the lateral lines on the first segment. Though the markings are variable, a bright-yellow stripe along the length of the dorsal surface is characteristic of S. litura larvae.
Larval instars can be distinguished on the basis of head capsule width, ranging from 2.7 to 25 mm. Body length ranges from 2.3 to 32 mm.
The pupa is 15-20 mm long, red-brown; tip of abdomen with two small spines.
Moth, with grey-brown body, 15-20 mm long; wingspan 30-38 mm. The forewings are grey to reddish-brown with a strongly variegated pattern and paler lines along the veins (in males, bluish areas occur on the wing base and tip); the hindwings are greyish-white with grey margins, often with dark veins in S. litura (but without in S. littoralis). See also Schmutterer (1969), Cayrol (1972) and Brown and Dewhurst (1975).
DistributionTop of page
The tobacco caterpillar, S. litura, is one of the most important insect pests of agricultural crops in the Asian tropics. It is widely distributed throughout tropical and temperate Asia, Australasia and the Pacific Islands (Feaking, 1973; Kranz et al., 1977). Records of S. litura having limited distribution in (or being eradicated from) Germany, Russian Federation, Russian Far East, the UK and Réunion may in fact refer to S. littoralis.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 30 Jun 2021
Risk of IntroductionTop of page
S. litura and S. littoralis are listed as of quarantine significance by EPPO, CPPC, NAPPO and OIRSA.
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Protected agriculture (e.g. glasshouse production)||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Disturbed areas||Present, no further details||Natural|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Principal habitat||Natural|
|Terrestrial||Natural / Semi-natural||Riverbanks||Secondary/tolerated habitat||Natural|
|Terrestrial||Natural / Semi-natural||Wetlands||Secondary/tolerated habitat||Natural|
Hosts/Species AffectedTop of page
The host range of S. litura covers at least 120 species. Among the main crop species attacked by S. litura in the tropics are Colocasia esculenta, cotton, flax, groundnuts, jute, lucerne, maize, rice, soyabeans, tea, tobacco, vegetables (aubergines, Brassica, Capsicum, cucurbit vegetables, Phaseolus, potatoes, sweet potatoes and species of Vigna). Other hosts include ornamentals, wild plants, weeds and shade trees (for example, Leucaena leucocephala, the shade tree of cocoa plantations in Indonesia).
Both S. litura and S. littoralis are totally polyphagous (Brown and Dewhurst, 1975; Holloway, 1989).
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
Leaves are heavily attacked and bolls have large holes in them from which yellowish-green to dark-green larval excrement protrudes.
Leaves develop irregular, brownish-red patches and the stem base may be gnawed off.
The stems are often mined and young grains in the ear may be injured.
List of Symptoms/SignsTop of page
|Leaves / external feeding|
Biology and EcologyTop of page
S. litura eggs are laid in clusters of several hundreds, usually on the upper surface of the leaves. Fecundity varies from 2000 to 2600 eggs, and oviposition days vary from 6 to 8 days. The developmental thresholds and thermal requirements for different stages of S. litura are 64 day degrees above threshold 8°C, from oviposition to egg hatch, the larval period required 303 degree days and the pupal stage 155 degree days above a 10°C threshold. The response of various stages of S. litura to temperatures under constant laboratory conditions was similar to that under field conditions. The upper development threshold temperature for all stages was 37°C, and 40°C was lethal (Ranga Rao et al., 1989).
Eggs take 2-3 days to hatch, the larvae disperse quickly from the egg batch in groundnut. Newly hatched larvae can be detected by the 'scratch marks' they make on the leaf surface. The older larvae are night feeders and are usually found in the soil around the base of the plant during the daytime. They can chew large areas of leaf and at high population densities cause complete defoliation. The larvae can migrate in large groups from one field to another. In lighter soils, the larvae while hiding in the soil during daytime can also cause damage to groundnut pods.
The larvae go through six instars and the final instars weigh up to 800 mg. Individual larvae can consume around 4 g fresh weight of groundnut foliage. However, 80% of the total consumption is in the final instar.
Pupation takes place in the soil close to the plants. The pupal period lasts about 7-10 days.
After adult emergence, peak oviposition occurs on the second night. Females mate three or four times during their lifetime, while males mate up to 10 times.
In Andhra Pradesh, India, S. litura completes 12 generations a year, each lasting slightly more than a month in winter and less than a month in the hot season.
ClimateTop of page
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
|Cf - Warm temperate climate, wet all year||Tolerated||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Tolerated||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Tolerated||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Absolute minimum temperature (ºC)||6|
|Mean annual temperature (ºC)||10||37|
|Mean maximum temperature of hottest month (ºC)||37|
|Mean minimum temperature of coldest month (ºC)||8|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Bacillus thuringiensis kurstaki||Pathogen|
|Chelonus formosanus||Parasite||Eggs; Arthropods|Larvae|
|Copidosoma floridanum||Parasite||Eggs; Arthropods|Larvae|
|Cytoplasmic polyhedrosis virus (CPV)||Pathogen||Arthropods|Larvae|
|Eurytoma syleptae||Hyperparasite||Arthropods|Larvae; Arthropods|Pupae|
|Microchelonus heliopae||Parasite||Eggs; Arthropods|Larvae|
Notes on Natural EnemiesTop of page
After Rao et al., 1993.
S. litura is known to be atttacked by many natural enemies at various life stages. Altogether, about 131 species of natural enemies have been reported from different parts of the world.
Four species of trichogrammatids, one scelionid and one braconid which had been reported as egg parasitoids of S. litura, an unidentified Chelonus species and species of Telenomus, have also been reported as both egg and larval parasitoids. A total of 10 egg parasitoids have been reported from different parts of the host distribution. Among the trichogrammatids, T. chilonis from India (Joshi et al., 1979; Patel et al., 1971) and T. dendrolimi from China (Chiu and Chou, 1976) are the most common. These species are often reported from the eggs of several other hosts.
Generally, the larval stage of S. litura is more prone to parasitism. Larval parasitoids of S. litura attack young to mature larvae and a few also attack eggs and larvae, and larvae and prepupae. Fifty-eight parasitoid species have been reported to attack the larval stage of this species. Of these, 47% were braconids, 19% ichneumonids, 16% tachinids, 10% eulophids, 3% chalcids, and 2% scelionids, encrytids and muscids. In general, 84% were Hymenoptera, and 16% Diptera.
In India, 32 different species of parasitoids have been reported as larval parasitoids of S. litura. Among these, Apanteles and species of Bracon were the most commonly reported. Rai (1974) surveyed vegetable crops in the state of Karnataka and found that 10% of larval mortality was caused by Chelonus formosanus. Jayanth and Nagarkatti (1984) reported the emergence of up to 12 tachninid parasitoids (Peribaea orbata) from a single S. litura larva in Karnataka state, India.
Rao and Satyanarayana (1984), during a pest survey of natural enemies of S. litura in Andhra Pradesh, India, reported Zele chlorophthalma as a larval parasitoid.
Sathe (1987) in a survey for natural enemies of S. litura in Maharashtra region of India reported Campoletes chlorideae and Apanteles colemani. During the same survey two new Braconid species (Enicospilus sp. and Echthromorpha sp.) were found responsible for the 5% parasitization of S. litura, while A. colemani and A. prodeniae parasitized up to 20% larvae.
Relatively few pupal parasitoids have been reported from S. litura. Eight parasitoid species have been reported from the pupal stage of S. litura, one of which is a larval-pupal parasitoid (Ichneumon sp.) and one a prepupal parasitoid (species of Chelonus).
Altogether 36 predatory insects from 14 families and 12 species of spiders, representing six families were reported to feed on S. litura eggs, larvae and pupae in different parts of the world. Of the total predators reported to feed on S. litura, 50% of the insect predatory fauna and 83% of the spiders were from India.
Nosema carpocapsae was found to infect S. litura larvae in New Zealand (Malone and Wigley, 1980), India (Narayanan and Jayaraj, 1979), Japan (Watanabe, 1976) and China (Tsai et al., 1978; Li and Wenn, 1987).
So far four fungi have been reported to infect S. litura and cause physiological disorders in larval growth and development: Aspergillus flavus, Beauveria bassiana, Nomuraea rileyi and Metarhizium anisopliae. Zaz and Kishwaha (1983) reported B. bassiana infecting S. litura in cauliflower crops in Rajasthan. Siddaramaiah et al. (1986) reported an incidence of larval infection with M.anisopliae in groundnut in Karnataka. The infection first appeared in the second fortnight of June, was highest in mid-August, and decreased by November.
Viral diseases of this species have been reported from China, Japan, India and New Zealand. Among the viruses, nuclear polyhedrosis viruses are the most common and potent. Narayanan (1985) from Karnataka, reported the occurrence of a granulosis virus in dead S. litura larvae. Eggs and all six larval instars were highly susceptible to the virus, the mortality was 100% in eggs and first to fifth-instar larvae and 50% in the last larval instar. The disease killed older larvae more rapidly than younger ones.
Four nematode species have been reported parasitizing S. litura in India and one of them has also been reported in Japan. Bhatnagar et al. (1985) found S. litura larvae parasitized by the mermithid nematodes Ovomermis albicans, Hexamermis sp. and Pentatomermis sp. They observed more nematode activity on alfisols than on vertisols. They also discussed the population dynamics and distribution of nematodes and the arthropod hosts. Kondo and Ishibashi (1984) explained the infectivity and propagation of entomogenous nematodes Steinernema sp. on S. litura from Japan.
Means of Movement and DispersalTop of page
The moths have a flight range of 1.5 km during a period of 4 h overnight, facilitating dispersion and oviposition on different hosts (Salama and Shoukry, 1972). They can accordingly fly quite long distances. The caterpillars can migrate over short distances.
In international trade, eggs or larvae may be present on planting material, cut flowers or vegetables; for example, the introduction of S. litura into the UK was on aquatic plants imported from Singapore (Aitkenhead et al., 1974). The pupae can be present in soil. The pupae are long-lived and could be transported over a considerable distance if not crushed, but to establish a viable population several specimens of both sexes need to be transported.
Pathway CausesTop of page
Pathway VectorsTop of page
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bulbs/Tubers/Corms/Rhizomes||eggs; larvae; pupae||Yes||Yes||Pest or symptoms usually invisible|
|Flowers/Inflorescences/Cones/Calyx||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Fruits (inc. pods)||eggs; larvae; pupae||Yes||Yes||Pest or symptoms usually invisible|
|Leaves||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Stems (above ground)/Shoots/Trunks/Branches||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|True seeds (inc. grain)||eggs; larvae; pupae||Yes||Yes||Pest or symptoms usually invisible|
|Plant parts not known to carry the pest in trade/transport|
|Growing medium accompanying plants|
Wood PackagingTop of page
|Wood Packaging liable to carry the pest in trade/transport||Timber type||Used as packing|
|Loose wood packing material||No|
|Processed or treated wood||No|
|Solid wood packing material with bark||No|
|Solid wood packing material without bark||No|
Impact SummaryTop of page
|Environment (generally)||Positive and negative|
ImpactTop of page
S. litura larvae are polyphagous defoliators, seasonally common in annual and perennial agricultural systems in tropical and temperate Asia. This noctuid is often found as part of a complex of lepidopteran and non-lepidopteran foliar feeders but may also damage tubers and roots. Hosts include field crops grown for food and fibre, plantation and forestry crops, as well as certain weed species.
Most work on the economic impact of S. litura has been conducted in India where it is a serious pest of a range of field crops. It has caused 12-23% damage to tomatoes in the monsoon season, and 9-24% damage in the winter (Patnaik, 1998). In a 40- to 45-day-old potato crop, damage ranged from 20 to 100% in different parts of the field depending on moisture availability. Larval populations peaked at four fully-grown larvae per square metre when the crop was 60-70 days old. Larvae also attacked exposed tubers when young succulent leaves were unavailable. Up to 2% of tubers were damaged in August-September and February (Trivedi, 1988). S. litura is also a pest of sugarbeet, with infestations commencing in March and peaking in late March and April (Chatterjee and Nayak, 1987). Severe infestations led to skeletonisation of leaves as well as feeding holes in roots that rendered the crop 'virtually unfit for marketing'. Late harvested crops were most severely affected and, in extreme cases, 100% of the roots were damaged, leading to 'considerable' yield reduction. Work on this species in a complex of other sugarbeet defoliators (Spodoptera exigua and Spilosoma obliqua) led to the development of an interactive exponential model based on length and severity of defoliation. It explained 88-90% of the variability in root and sugar yields and suggested the need for pest control when defoliation exceeded 25% during April. Control was not required if the pest appeared after the first week of May (Singh and Sethi, 1993).
S. litura is one of six defoliating pests of fodder cowpea which, in a field experiment, were responsible for consuming up to 85.5% of leaf area (Ram et al., 1989). Aroid tuber crops (including taro (Colocasia esculenta)) suffered yield losses of up to 29% as a result of infestation by S. litura, Aphis gossypii and spider mites (Pillai et al., 1993). S. litura is also a member of a complex that causes extensive defoliation of soyabean (Bhattacharjee and Ghude, 1985). Defoliation as severe as 48.7% during the pre-bloom stage of growth caused no 'marked' difference from a control treatment in which defoliation was prevented by repeated insecticide application. Number and weight of pods and grains per plant were, however, reduced when defoliation occurred at, or after, blooming.
In groundnut, S. litura is one of several pests that can be important during the pegging, podding and pod maturation stages of growth (Singh and Sachan, 1992). Several studies have aimed at quantifying the damage attributable to S. litura. Field experiments by Panchabhavi and Raj (1987) extended over 2 years and used artificial infestation of groundnut plots of 15 m² with differing densities of S. litura. Infestation levels of just three egg masses (of 250 eggs each) caused significant loss of groundnut pods and haulms. Infestation with 12 egg masses per plot led to a haulm yield reduction of up to 43.7% and a pod yield reduction as high as 27% compared with an insecticide-protected control treatment. In other field experiments over 3 consecutive years, leaf damage attributed to S. litura tended to decline with delayed sowing time irrespective of groundnut cultivar (Patil et al., 1996). Leaf damage fell from 51.8% for mid-June sown crops to 19.2% for late-July sown crops. Mean pod yields were 2.68 and 0.99 t/ha, respectively.
Another field study determined the effect of artificially infesting individual groundnut plants with third-instar S. litura larvae 15, 30 or 45 days after emergence (d.a.e.) (Dhir et al., 1992). The most severe damage occurred when plants were infested with three larvae 15 d.a.e. These lost 98.3% of leaf area, and pod yield was reduced by 50%. Even single larvae caused the leaf area to be more than halved and pod yield to fall by 27.3%. Plants infested 30 d.a.e. suffered similar levels of damage but those infested 45 d.a.e. were less severely affected.
Selveraj et al. (2014) conducted a study of the ecological factors on the incidence and development of S. litura on cotton. They found a positive correlation with relative humidity, sunshine hours and dewfall, but a negative correlation with wind velocity. Determination of the effects of different weather factors on the population and incidence of S. litura in cotton is essential for effective pest management.
S. litura causes damage to many species of forest and plantation trees and shrubs (Roychoudhury et al., 1995). It is responsible for brown flag syndrome in banana (Ranjith et al., 1997), and 5-10% fruit damage in grapes (Balikai et al., 1999). In Paulownia nurseries and plantations a complex of at least 24 defoliating pest species causes damage. Within this complex, S. litura was considered the most important noctuid species, with an incidence of 72% in weekly surveys (Kumar and Ahmad, 1998). Peak activity occurred in July and September, with an average of 6.5 and 5.2 larvae per plant in these months, respectively. During this period, many plants were completely defoliated by S. litura. In teak, it is one of about 139 defoliators that attack all stages from seedlings to mature trees (Roychoudhury et al., 1995). S. litura is abundant on teak in June and July and damage incidence in seedlings has been reported to be as high as 56%. Late-instar larvae were found to feed preferentially on mature teak leaves, whilst early instars fed on leaves of intermediate age. High concentrations of polyphenols in young leaves (Roychoudhury et al., 1995) may reduce their attractiveness to S. litura larvae but differing levels of susceptibility among nine teak clones were attributed to the nitrogen:potassium ratio of the foliage (Roychoudhury et al., 1998).
Two separate Indian studies concerned the potential of S. litura as a biological control agent of weeds. Sites of larval feeding on leaves and petioles of the aquatic weed Eichhornia crassipes were said to be vulnerable to infection by fungi, and overall damage levels by S. litura were estimated to be 20-25% (Jamil et al., 1984). It was recognized, however, that the potential use of this polyphagous agent needs caution because preferred hosts include important crops such as castor and cotton. In a second study, S. litura attacked the weeds, Marsilea quadrifolia, Ammannia baccifera and Eclipta alba, and although up to 10 larvae were recorded per plant on nearby rice, no damage was reported to this crop (Sain et al., 1983).
Studies elsewhere in southern Asia illustrate the economic impact of S. litura. In Pakistan, it is one of several lepidopteran pests attacking a wide range of crops including cotton and rice (Ahmad and Kamaluddin, 1987), as well as cabbage, tobacco, groundnut, soyabean, lucerne, gram, cowpea, tomato, cauliflower, carrot, onion, brinjal, turnip, radish and spinach (Maree et al., 1999). In the latter study, S. litura was present in a December-planted cabbage crop during January-March with densities peaking at 4.55/plant on 2 February. Damage was highest in early February and there was a positive correlation between plant damage and pest density. Fieldwork in Bangladesh led to a 10% visual damage spraying threshold being proposed for S. litura and Plutella xylostella (Ali and Bakshi, 1994).
In eastern Asia, damage by S. litura occurs as far north as Japan, extending south to Indonesia. Japanese soyabean field plots artificially infested with one or two S. litura egg masses per plant suffered estimated leaf area reductions of 14.3 and 23.2% and yield losses of 13.9 and 24.7%, respectively, compared with control plots (Higuchi et al., 1994). Similar work indicated that yield losses caused by early infestation (at the flowering and pod-development stages) were attributable chiefly to reduced production of pods per plant, whilst infestations on more mature plants affected yield by reducing the weight of individual beans (Higuchi, 1991). In studies with red (adzuki) bean, the effect of defoliation by S. litura on yield was investigated by artificial defoliation. Combined with observations that the average leaf area consumed by S. litura was 203.9 cm², of which 82% was consumed by final-instar larvae, an action threshold of two final-instar larvae/plant (causing 5% yield loss) was estimated (Katayama and Sano, 1989). Equivalent thresholds were developed for protected crops in Korea (Nakasuji and Matsuzaki, 1977). In this instance, a 10% yield loss threshold was applied and the threshold densities of S. litura were 4.6 and 15.4 neonates or 0.8 and 2.6 egg masses/m², for aubergine and sweet peppers, respectively. S. litura is also found in Welsh onion crops in Korea, but is less damaging than other lepidopterans such as Liriomyza chinensis (Ahn et al., 1991).
A lepidopteran pest complex including S. litura, Pieris rapa and Plutella xylostella damages cabbage in China (Zhou et al., 1996). Experiments to determine larval feeding capacity showed S. litura to be intermediate amongst these species (Zhu et al., 1994). In the Zhejiang region of China, S. litura is usually first recorded in June on legumes, aubergines, tomatoes, sweet peppers and tree fruits, and subsequently damages cabbage seedlings in July (Chen et al., 1999). Damage becomes more severe in August and September when cauliflower, sweet potato and legumes may also be attacked. The pest appears unable to overwinter in this area and is virtually absent after the end of October. In Taiwan, S. litura attacks gladiolus foliage (Wang, 1982; Liu, 1998) as well as soyabean and adzuki bean (Lee, 1989). Damage ranged between 3 and 13% over three consecutive years for the latter crops.
In South-East Asia, S. litura has been recorded damaging tobacco in Malaysia (Hamid et al., 1992), mungbean (Vigna radiata) in Thailand (Sepswasdi et al., 1991), tree legumes in Indonesia (Matsumoto, 2000) and Paraserianthes falcataria seedlings in the Philippines (Braza, 1990).
Roa et al. (2012) studied the impact of climate change on the development of S. litura in the near future and reported a change in the nutrient value of groundnut foliage and as a result a higher consumption, lower digestive efficiency, slower growth and longer time to pupation (1 day more than ambient).
Risk and Impact FactorsTop of page
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Tolerant of shade
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Benefits from human association (i.e. it is a human commensal)
- Fast growing
- Has high reproductive potential
- Host damage
- Infrastructure damage
- Negatively impacts agriculture
- Negatively impacts livelihoods
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
Detection and InspectionTop of page
Similarities to Other Species/ConditionsTop of page
S. litura can be easily confused with S. littoralis as in both cases adults and larvae are similar, and they can be distinguished only through examination of genitalia. On dissection of the genitalia, ductus and ostium bursae are the same length in female littoralis, different lengths in litura. The shape of the juxta in males is very characteristic, and the ornamentation of the aedeagus vesica is also diagnostic. The presence of newly hatched larvae can be detected by the 'scratch marks' they make on the leaf surface.
For more information on the morphological discrimination between the adult, pupal and larval stages of the two species, see Mochida (1973).
An EPPO standard provides guidance for the identification of S. littoralis, S. litura, S. frugiperda and S. eridania (OEPP/EPPO, 2015).
Prevention and ControlTop 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.
After Rao et al. (1993).
The green revolution in Asia brought with it an increased awareness of the potential of insecticides for increasing the sustainability of rice production. Unfortunately, the involvement of farms in insecticide-related technologies did not proceed as fast as the rate of subsidy spread and the overspill of insecticide usage into the fields of legume growers and horticulturalists. Legume pests are increasing in economic importance throughout Asia due to the destruction of natural control systems, and the build-up of insecticide resistance following the 'spraymania' of many farmers. If this is to be counteracted, natural control needs to be given increased emphasis as a component of the IPM approach. S. litura populations in groundnut fields are increasing in number and intensity, especially in fields where insecticides have been applied (Stechmann and Semisi, 1984; Rao and Shanower, 1988).
In the past, the control of arthropods depended mostly on inexpensive and efficient insecticides. But in recent years populations of many pests including S. litura have developed resistance to many commercially available pesticides (Ramakrishnan et al., 1984; Naeem Abbas et al., 2014). Studies at ICRISAT between 1991 and 1996 revealed the occurrence of resistance to cypermethrin, fenvalerate and quinalphos, by 197-, 121-, 29- and 362-fold, respectively (JA Wightman, ICRISAT, Andhra Pradesh, India, personal communication, 1996).
The control of arthropod pests is therefore becoming increasingly difficult and it is vital that all biological alternatives to insecticides need to be given greater priority, both in research and application.
New insecticides have been tested to deal with resistant strains of this moth and some promising results are coming forward (Venkateswarlu et al., 2005). Neem oil microemulsion proved significantly superior than macroemulsion (Swaran Dhingra et al., 2006).
New molecules such as chlorantraniliprole, spinosad and emamectin benzoate have shown promising results against S. litura (Gadhiya et al., 2014) but chlorantraniliprole gave the highest cost: benefit ratio among pesticides tested by Patil et al. (2014) on soyabeans.
Chatterjee and Mondal (2012) tested a number of new chemicals and their application methods on different vegetable crops in India and South-East Asia against lepidopterous pests and found flubendiamide, spinosad and chlorfenapyr to be the most effective.
Following studies on the sublethal effects of mathofenozide, Shahout et al. (2011) concluded that the effects of methoxyfenozide with its sterilizing properties, if used strategically on S. litura, might induce changes in the population dynamics of this pest in vegetable crops and could be considered a potent insecticidal compound for controlling this pest.
Suganthy and Sakthivel (2013) studied different bio-pesticides against S. litura infesting fields of Gloriosa superba and showed that flavonoids could be used as an alternative to chemical pesticides in the gloriosa ecosystem and as a component in organic pest management.
Plant oils and insecticides mixtures (synthetic pyrethroids) gave a higher mortality rate on 8-day-old larvae of S. litura than the synthethic pyrethroids alone (Anju and Srivastava, 2012).
In the past the mass releases of egg and larval parasitoids for the control of S. litura in different crops in different geographical regions had achieved only partial success (Patel et al., 1979; Michael et al., 1984). Observations in ICRISAT groundnut fields revealed more leaves with defoliator damage in insecticide applied fields than unsprayed areas (Wightman et al., 1990). Similar observations were also made during farmers' field surveys in the post-rainy season in coastal Andhra Pradesh, India (Rao and Shanower, 1988). Stechmann and Semisi (1984) also shared the same opinion after surveying taro fields in Western Samoa. In view of the development of insecticidal resistance and the destruction of the natural enemies, and the polyphagous nature of this species, there is a need to give more consideration to the role of natural enemies as a component of integrated approaches to managing S. litura.
Mass releases of an indigenous egg-larval parasite Chelonus heliopae in 1971-73 in Anand, Gujarat, India, against S. litura in cauliflower proved ineffective in controlling the pest. During 1974, weekly release of Telenomus remus, an egg parasitoid, in a tobacco nursery did not result in any parasitism. However, five weekly releases of 50,000 parasites per 0.2 ha and two releases of 15,000 parasitoids per 0.2 ha in cauliflower resulted in 60% parasitism (Patel et al., 1979).
T. remus was introduced to Western Samoa and was recorded by Braune (1982) as a common egg-larval parasitoid of S. litura, with parasitism averaging 54%. Complete parasitization was observed only in small egg masses (up to 150 eggs) and the percentage of parasitization decreased with an increase in size of egg mass. T. remus could oviposit only in host eggs on the surface of the host egg mass. Thus the effectiveness of T. remus was limited to the large compact egg masses of S. litura.
Six parasitoid species, Apanteles ruficrus, Cotesia marginiventris, Apanteles kazak, Campoletes chloridae, Hyposoter didymator and T. remus were introduced to Western Australia from overseas in 1978-83 and released against S. litura and 11 other economically important pests. The highest level of parasitism by A. ruficrus (80% and above) was noticed in Mythimna sp. (Michael et al., 1984).
In Western Samoa, Stechmann and Semisi (1984) collected information on S. litura damage levels in relation to natural populations of Apanteles spp. in taro fields. They found that this pest was more severe on taro where insecticides and herbicides were widely used, which perhaps created imbalance between the pest and its natural enemies. Barrion and Litsinger (1987) reported the presence of Peribaea orbata as a gregarious larval parasitoid on S. litura.
Wang et al. (2014) studied the relationship between the larval parasitoid Meteorus pulchricornis and the bacterium Empedobacter brevis. The study suggested that the bacteria has a negative effect on M. pulchricornis, but the impact could be alleviated by using low bacteria concentration and extending the time between the application and wasp release in biological control practices.
The biology of Canthoconidia furcellata was studied in the laboratory with a view to using this predator in an integrated pest management programme for tobacco pests. Chu and Chu (1975) studied the effects of temperature on the growth of C. furcellata and found that 71,216 and 134 degree days were required for egg, nymph and adult stages, respectively. It was concluded that there are five to six generations per year of this predator in northern Taiwan.
Nakasuji et al. (1976) observed a predatory wasp, preferentially selecting fifth- and sixth-instar larvae over early instars. The wasps were more active and attacked more larvae in fields with high larval density than those with low larval density. However, the percentage of predation was lower in the field with highest density of S. litura larvae.
Deng and Jim (1985) reported Conocephalus sp. as a new predator on egg masses of S. litura in Guanxi, China. This katydid was successfully reared on an artificial diet. Field releases of nymphs and adults of Conocephalus sp. were attempted for control of Scirpophaga incertulus.
Ansari et al. (1987) reported Serratia marcescens from Karnataka, India, attacking larvae of the noctuids Helicoverpa armigera and S. litura. In laboratory tests, S. litura was found to be more susceptible to the bacterium than H. armigera. The bacterium was equally pathogenic when ingested through artificial diet or the natural food plant, but pathogenicity by contact application to the body of larvae was poor.
Zaz and Kushwaha (1983) found Bacillus thuringiensis to be an effective microbial insecticide against S. litura larvae in cauliflower fields in Rajasthan, India.
The efficiency of B. thuringiensis was enhanced significantely though protoplast fusion with a strain of Bacillus subtilis (Kannan Revathi et al., 2014).
Asayama and Ohoishi (1980) from Japan and Phadke and Rao (1978) from India, investigated the pathogenicity of a green muscardine fungus Nomuraea rileyi. Laboratory studies in India indicated that this fungus was harmless to eggs of an egg parasitoid, Telenomus preditor, on Achaea janata and recommended the combined use of the fungus and the egg parasitoid in biocontrol programmes against A. janata. This may also apply to S. litura management.
Laboratory studies were undertaken to evaluate the bioefficacy of Beauveria bassiana against third-instar larvae of S. litura. B. bassiana was identified, isolated and maintained from field-collected cadavers of lepidopteran larvae. Minimum mortality was observed in the control, i.e. 23.3%, and the percentage mortality increased as the number of spores increased (Gupta and Bhupendra Kumar, 2014).
Research was also carried out on entophytic fungi (Khuskia oryzae and Cladosporium uredinicola), which showed adverse effects on survival and fitness of the insects (Abhinay et al., 2014).
Krishnaiah et al. (1985) conducted field trials with a nuclear polyhedrosis virus against S. litura damage in black gram (Vigna mungo) fields in Andhra Pradesh, India. Two sprays of virus suspension gave effective control similar to chemical insecticides tested.
Chari et al. (1985) evaluated the effectiveness of integrated management of natural enemies and viral diseases to control S. litura on tobacco seedlings in Gujarat, India. They concluded that a combination of biological control agents, insect growth regulators, antifeedants and a trap crop on all sides of the nursery was an ecologically sound procedure for the control of S. litura.
Different doses of SpltMNPV on final instars of S. litura showed dose-related mortality, but sublethal doses on subsequent generations needs to be considered in the design of baculovirus-based pest management (Mohammad Monobrullah and Umi Shankar, 2008).
Integrated Pest Management
In recent years, due to crop failures experienced despite the use of several combinations of chemicals, an integrated approach based on cultural and biocontrol with efficient monitoring using pheromones has been developed (JA Wightman, ICRISAT, Andhra Pradesh, India, personal communication, 1996). The IPM technology that has been developed and implemented in irrigated groundnut where S. litura is endemic has the following components:
- clean cultivation to expose Spodoptera pupae to natural enemies and weather-related factors
- sunflower, taro (Zhou, 2009) and castor plants (that attract Spodoptera) to be sown as trap crops both around and within fields
- pheromone traps to predict Spodoptera egg laying
- mechanical collection of egg masses and larvae from trap plants on alternate days following the 'warning' from the pheromone traps
- application of fungicide (chlorothalonil) at the appearance of the first leaf spot lesions, and again after 10 days
- an application of neem kernel extract during the early stages of crop growth if necessary
- Pongamia glabra oil treatment on tomato plants gave significant reductions on the populations of S. litura while no adverse effects againsts it natural enemies (Marimuthu, 2008)
- application of nuclear polyhedrosis virus at 500 larval equivalents per hectare in the evening if needed.
Sahayaraj (2011) gives a summary of different types of plant extracts used by farmer on groundnuts and discusses their effiency.
Developments in pheromone technology have made it possible to monitor S. litura in the field, to improve on timing of plant protection measures within groundnut IPM programmes.
The identification of a male sex pheromone of S. litura, (ZE) 9,11-tetradecadienyl acetate and (ZE) 9,12-tetradecadienyl acetate by Youshima et al. (1974) has enabled effective monitoring of this species for several years. The basic work regarding trap design, height, longevity of the septa, and the potential role of this technology in groundnut has been thoroughly studied at ICRISAT Center, Hyderabad, India over the past decade. These studies have clearly indicated the migratory behaviour of the species in different areas. At present, pheromone technology has given high priority in monitoring for timing of plant protection measures within groundnut IPM programmes. The studies on trap density in groundnut situations indicated no significant differences in moth catches when there were four or more traps per hectare. No decline was noticed in moth catch with increase in trap density. This indirectly suggests a limitation in utilizing the technology in mass trapping operations (Ranga Rao et al., 1989).
However, there have been some promising results in monitoring the population of moths on Chinese cabbage (Yang Song et al., 2009); spraying times and costs of chemical pesticides against S. litura were significantly reduced by the adoption of sex pheromone trapping.
Population projections based on life tables and stage-specific consumption rates can reveal the stage structure and damage potential of the pest population of the moths (Tuan et al., 2014). This method could prove to be more reliable as the data obtained by pheromone traps. It is evident that these life tables have to be developed for each area where the moth occurs and one should take into account climate change and yearly temperature and rainfall patterns. It was already established that minimum temperature is the predominant factor that influences pheromone traps whereas wind velocity is predominant in light traps. The overall influence of all the weather factors was high in case of pheromone traps compared to light traps (Prasad et al., 2009).
The development of resistance to S. litura in suitable groundnut varieties has been regarded as a high priority for Asian groundnut farmers for a number of years. The results of experiments carried out in 1986 and 1987 (data in Wightman and Ranga Rao, 1993) indicated the possibility that ICGV 86031 had some resistance to S. litura combined with high yield in the post-rainy season. This hope was substantiated in further tests on the ICRISAT research farm and in farmers' fields in coastal Andhra Pradesh (southern India). In the limited trials that have been carried out, farmers had sufficient confidence to grow this variety without protecting it with insecticides. They were rewarded with higher yields and lower variable costs than neighbouring farmers who grew locally acceptable varities but applied insecticides to kill defoliators. PI 269116, PI 269118 and PI 262042 had resistance to S. litura, but none were outstanding (Campbell and Wynne, 1980).
Bioassays carried out with larvae as preliminaries to detect the mechanism of resistance (independent tests by Ranga Rao (ICRISAT) and Padgham (NRI)) revealed no antibiosis effect on second- to sixth-instar larvae when fed mature leaves of ICGV 86031. The main mechanism of resistance is currently thought to be tolerance, manifested as the enhanced ability of vegetative tissue to regrow following defoliation.
However, first-instar larvae suffered 56% mortality when fed on ICGV 86031 compared with 12% mortality when fed on susceptible ICG 221. Padgham also found that newly hatched larvae had a marked propensity to vacate the leaves of this variety in the first 2 hours of free life. This suggests that the resistance factor which influences the neonates is associated with the leaf surface, because their feeding activity is restricted to scraping the leaf surface. The antixenosis demonstrated by ICGV 86031 is likely to increase the first-instar mortality that is characteristic of r-strategist noctuids (Kyi et al., 1991) and will therefore contribute to the determination of the level of damage caused by the older larvae among which mortality is comparatively low.
Amin et al. (2011) investigated the morphological and biochemical characteristics of three varieties of cotton and observed their effect on feeding and growth of S. litura. At least one variety was not suited for cotton growers. In a study of the interaction between the virus and the parasitoid, Guo Huifang et al. (2013) showed that the use of an appropriate concentration has the potential to improve the efficiency of the biological control.
Gaps in Knowledge/Research NeedsTop of page
S. litura belongs to a species complex 'Spodoptera' consisting of very similar species with similar ecology in other continents. Correct identification is essential for establishing the correct distribution pattern of this pest.
ReferencesTop of page
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