Helicoverpa armigera (cotton bollworm)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Helicoverpa armigera (Hübner, 1809)
Preferred Common Name
- cotton bollworm
Other Scientific Names
- Chloridea armigera Hübner
- Chloridea obsoleta
- Helicoverpa obsoleta Auct.
- Heliothis armigera Hübner
- Heliothis fusca Cockerell
- Heliothis obsoleta Auct.
- Heliothis rama Bhattacherjee & Gupta
- Noctua armigera Hübner
International Common Names
- English: African cotton bollworm; corn earworm; gram pod borer; grub, tomato; old world bollworm; tobacco budworm
- Spanish: gusano bellotero del algodon; gusano de la cápsula; gusano del elote del maiz; gusano verde de la cápsula; noctua del tomate; oruga de las cápsulas del algodón; oruga del choclo
- French: chenille des epis du mais; noctuelle des tomates; ver de la capsule
Local Common Names
- Denmark: amerikansk bomuldsugle
- Germany: Altweltlicher Baumwollkapselwurm
- Italy: elotide del cotone; elotide del granturco; elotide del pomodoro; elotide del tabacco; nottua del granturco; nottua gialla del granturco
- Netherlands: mimosa-rups
- HELIAR (Helicoverpa armigera)
- HELIRA (Heliothis rama)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Lepidoptera
- Family: Noctuidae
- Genus: Helicoverpa
- Species: Helicoverpa armigera
Notes on Taxonomy and NomenclatureTop of page The taxonomic situation is complicated and presents several problems. Hardwick (1965) reviewed the New World corn earworm species complex and the Old World African bollworm, most of which had previously been referred to as a single species (Heliothis armigera or H. obsoleta), and pointed out that there was a complex of species and subspecies involved. He proposed that the New World H. zea (first used in 1955) was distinct from the Old World H. armigera on the basis of male and female genitalia, and he described the new genus Helicoverpa to include these important pest species. Some 80 or more species were formerly placed in Heliothis (sensu lato), and Hardwick referred 17 species (including 11 new species) to Helicoverpa on the basis of differences in both male and female genitalia. Within this new genus the zea group contains eight species, and the armigera group two species with three subspecies. See also Hardwick (1970).
Because the old name of Heliothis for the pest species (four major pest species and three minor) is so well established in the literature, and as dissection of genitalia is required for identification, there has been resistance to the name change (e.g. Heath and Emmet, 1983), but Hardwick's work is generally accepted and so the name change must also be accepted (see Matthews, 1991).
DescriptionTop of page Egg
Yellowish-white and glistening at first, changing to dark-brown before hatching; pomegranate-shaped, 0.4-0.6 mm in diameter; the apical area surrounding the micropyle is smooth, the rest of the surface sculptured in the form of approximately 24 longitudinal ribs, alternate ones being slightly shorter, with numerous finer transverse ridges between them; laid on plants which are flowering, or are about to produce flowers.
The first and second larval instars are generally yellowish-white to reddish-brown in colour, without prominent markings; head, prothoracic shield, supra-anal shield and prothoracic legs are very dark-brown to black, as are also the spiracles and tuberculate bases to the setae, which give the larva a spotted appearance; prolegs are present on the third to sixth, and tenth, abdominal segments. A characteristic pattern develops in subsequent instars. Fully grown larvae are ca 30-40 mm long; the head is brown and mottled; the prothoracic and supra-anal plates and legs are pale-brown, only claws and spiracles remaining black; the skin surface consists of close-set, minute tubercles. Crochets on the prolegs are arranged in an arc. The final body segment is elongated. Colour pattern: a narrow, dark, median dorsal band; on each side, first a broad pale band, then a broad dark band; on the lateral line, a broad, very light band on which the row of spiracles shows up clearly. The underside is uniformly rather pale. On the basic dorsal pattern, numerous very narrow, somewhat wavy or wrinkled longitudinal stripes are superimposed. Colour is extremely variable and the pattern described may be formed from shades of green, straw-yellow, and pinkish- to reddish-brown or even black.
Mahogany-brown, 14-18 mm long, with smooth surface, rounded both anteriorly and posteriorly, with two tapering parallel spines at posterior tip.
Stout-bodied moth of typical noctuid appearance, with 3.5-4 cm wing span; broad across the thorax and then tapering, 14-18 mm long; colour variable, but male usually greenish-grey and female orange-brown. Forewings have a line of seven to eight blackish spots on the margin and a broad, irregular, transverse brown band. Hindwings are pale-straw colour with a broad dark-brown border that contains a paler patch; they have yellowish margins and strongly marked veins and a dark, comma-shaped marking in the middle. Antennae are covered with fine hairs.
For more information, see Dominguez Garcia-Tejero (1957), Hardwick (1965), Cayrol (1972), Delatte (1973), King (1994).
DistributionTop of page The first issue of the CIE distribution map (CIE, 1952) included the American continent, but the species found there is now known to be Helicoverpa zea (Boddie), the New World bollworm (EPPO/CABI, 1992).
See also CABI/EPPO (1998, No. 77).
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Afghanistan||Present||IIE, 1993; EPPO, 2014|
|Armenia||Present||IIE, 1993; EPPO, 2014|
|Azerbaijan||Present||IIE, 1993; EPPO, 2014|
|Bangladesh||Widespread||IIE, 1993; EPPO, 2014|
|Bhutan||Present||IIE, 1993; EPPO, 2014|
|Bismarck Archipelago||Present||IIE, 1993|
|Brunei Darussalam||Present||Waterhouse, 1993|
|Cambodia||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|China||Restricted distribution||IIE, 1993; EPPO, 2014|
|-Anhui||Present||IIE, 1993; EPPO, 2014|
|-Fujian||Present||IIE, 1993; EPPO, 2014|
|-Guangdong||Present||IIE, 1993; EPPO, 2014|
|-Guangxi||Present||IIE, 1993; EPPO, 2014|
|-Guizhou||Present||IIE, 1993; EPPO, 2014|
|-Hainan||Present||IIE, 1993; EPPO, 2014|
|-Hebei||Widespread||IIE, 1993; EPPO, 2014|
|-Heilongjiang||Present||IIE, 1993; EPPO, 2014|
|-Henan||Widespread||IIE, 1993; EPPO, 2014|
|-Hong Kong||Widespread||EPPO, 2014|
|-Hubei||Present||IIE, 1993; EPPO, 2014|
|-Hunan||Present||IIE, 1993; EPPO, 2014|
|-Jiangsu||Widespread||IIE, 1993; EPPO, 2014|
|-Jiangxi||Present||IIE, 1993; EPPO, 2014|
|-Liaoning||Present||IIE, 1993; EPPO, 2014|
|-Nei Menggu||Present||IIE, 1993; EPPO, 2014|
|-Qinghai||Present||Zhang et al., 2001|
|-Shandong||Widespread||IIE, 1993; EPPO, 2014|
|-Sichuan||Present||IIE, 1993; EPPO, 2014|
|-Tibet||Present||IIE, 1993; EPPO, 2014|
|-Xinjiang||Widespread||IIE, 1993; EPPO, 2014|
|-Yunnan||Present||IIE, 1993; EPPO, 2014|
|-Zhejiang||Present||IIE, 1993; EPPO, 2014|
|Cocos Islands||Present||EPPO, 2014|
|Georgia (Republic of)||Present||IIE, 1993; EPPO, 2014|
|India||Widespread||IIE, 1993; EPPO, 2014|
|-Andaman and Nicobar Islands||Present||EPPO, 2014|
|-Andhra Pradesh||Present||IIE, 1993; EPPO, 2014|
|-Assam||Present||IIE, 1993; EPPO, 2014|
|-Bihar||Present||IIE, 1993; EPPO, 2014|
|-Chhattisgarh||Present||Netam et al., 2007|
|-Delhi||Present||IIE, 1993; EPPO, 2014|
|-Gujarat||Present||IIE, 1993; EPPO, 2014|
|-Haryana||Present||IIE, 1993; EPPO, 2014|
|-Himachal Pradesh||Present||IIE, 1993; EPPO, 2014|
|-Indian Punjab||Present||IIE, 1993; EPPO, 2014|
|-Jammu and Kashmir||Present||IIE, 1993; EPPO, 2014|
|-Jharkhand||Present||Rabindra and Devendera, 2007|
|-Karnataka||Present||IIE, 1993; EPPO, 2014; Manjula et al., 2015|
|-Kerala||Present||Levin et al., 2004|
|-Madhya Pradesh||Present||IIE, 1993; EPPO, 2014|
|-Maharashtra||Present||IIE, 1993; EPPO, 2014|
|-Manipur||Present||Devi et al., 2002|
|-Nagaland||Present||Imosanen, and Singh, 2005|
|-Odisha||Present||IIE, 1993; EPPO, 2014|
|-Rajasthan||Present||IIE, 1993; EPPO, 2014|
|-Sikkim||Present||IIE, 1993; EPPO, 2014|
|-Tamil Nadu||Present||IIE, 1993; EPPO, 2014|
|-Uttar Pradesh||Present||IIE, 1993; EPPO, 2014|
|-Uttarakhand||Present||Kuldeep and Ram, 2007; EPPO, 2014|
|-West Bengal||Present||IIE, 1993; EPPO, 2014|
|Indonesia||Present||IIE, 1993; EPPO, 2014|
|-Irian Jaya||Present||IIE, 1993; EPPO, 2014|
|-Java||Present||IIE, 1993; EPPO, 2014|
|-Moluccas||Present||IIE, 1993; EPPO, 2014|
|-Nusa Tenggara||Present||EPPO, 2014|
|-Sulawesi||Present||IIE, 1993; EPPO, 2014|
|-Sumatra||Present||IIE, 1993; EPPO, 2014|
|Iran||Widespread||IIE, 1993; EPPO, 2014|
|Iraq||Present||IIE, 1993; EPPO, 2014|
|Israel||Widespread||IIE, 1993; EPPO, 2014|
|Japan||Widespread||IIE, 1993; EPPO, 2014|
|Jordan||Present||IIE, 1993; EPPO, 2014|
|Kazakhstan||Present||IIE, 1993; EPPO, 2014|
|Korea, DPR||Present||IIE, 1993; EPPO, 2014|
|Korea, Republic of||Present||IIE, 1993; EPPO, 2014|
|Kuwait||Present||IIE, 1993; EPPO, 2014|
|Kyrgyzstan||Present||IIE, 1993; EPPO, 2014|
|Laos||Widespread||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Lebanon||Present||IIE, 1993; EPPO, 2014|
|Malaysia||Present||IIE, 1993; EPPO, 2014|
|-Peninsular Malaysia||Present||EPPO, 2014|
|-Sabah||Present||IIE, 1993; EPPO, 2014|
|-Sarawak||Present||IIE, 1993; EPPO, 2014|
|Myanmar||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Nepal||Present||IIE, 1993; EPPO, 2014|
|Pakistan||Present||APPPC, 1987; IIE, 1993; EPPO, 2014|
|Philippines||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Saudi Arabia||Widespread||IIE, 1993; EPPO, 2014|
|Singapore||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Sri Lanka||Present||IIE, 1993; EPPO, 2014|
|Syria||Widespread||IIE, 1993; EPPO, 2014|
|Taiwan||Widespread||IIE, 1993; EPPO, 2014|
|Tajikistan||Widespread||IIE, 1993; EPPO, 2014|
|Thailand||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Turkey||Restricted distribution||****||IIE, 1993; EPPO, 2014|
|Turkmenistan||Present||IIE, 1993; EPPO, 2014|
|United Arab Emirates||Present||IIE, 1993; EPPO, 2014|
|Uzbekistan||Present||IIE, 1993; EPPO, 2014|
|Vietnam||Present||IIE, 1993; Waterhouse, 1993; EPPO, 2014|
|Yemen||Widespread||IIE, 1993; EPPO, 2014|
|Algeria||Widespread||IIE, 1993; EPPO, 2014|
|Angola||Present||IIE, 1993; EPPO, 2014|
|Benin||Present||IIE, 1993; EPPO, 2014|
|Botswana||Present||IIE, 1993; EPPO, 2014|
|Burkina Faso||Present||IIE, 1993; EPPO, 2014|
|Burundi||Present||IIE, 1993; EPPO, 2014|
|Cameroon||Present||IIE, 1993; EPPO, 2014|
|Cape Verde||Present||IIE, 1993; EPPO, 2014|
|Central African Republic||Present||IIE, 1993; EPPO, 2014|
|Chad||Present||IIE, 1993; EPPO, 2014|
|Congo||Present||IIE, 1993; EPPO, 2014|
|Congo Democratic Republic||Present||IIE, 1993; EPPO, 2014|
|Côte d'Ivoire||Present||IIE, 1993; EPPO, 2014|
|Egypt||Widespread||IIE, 1993; EPPO, 2014|
|Ethiopia||Present||IIE, 1993; EPPO, 2014|
|Gabon||Present||IIE, 1993; EPPO, 2014|
|Gambia||Present||IIE, 1993; EPPO, 2014|
|Ghana||Present||IIE, 1993; EPPO, 2014|
|Guinea||Present||IIE, 1993; EPPO, 2014|
|Kenya||Present||IIE, 1993; EPPO, 2014|
|Lesotho||Present||IIE, 1993; EPPO, 2014|
|Libya||Widespread||IIE, 1993; EPPO, 2014|
|Madagascar||Present||IIE, 1993; EPPO, 2014|
|Malawi||Present||IIE, 1993; EPPO, 2014|
|Mali||Present||IIE, 1993; EPPO, 2014|
|Mauritania||Present||IIE, 1993; EPPO, 2014|
|Mauritius||Present||IIE, 1993; EPPO, 2014|
|Morocco||Restricted distribution||IIE, 1993; EPPO, 2014|
|Mozambique||Present||IIE, 1993; EPPO, 2014|
|Namibia||Present||IIE, 1993; EPPO, 2014|
|Niger||Present||IIE, 1993; EPPO, 2014|
|Nigeria||Present||IIE, 1993; EPPO, 2014|
|Réunion||Present||IIE, 1993; EPPO, 2014|
|Rwanda||Present||IIE, 1993; EPPO, 2014|
|Saint Helena||Present||IIE, 1993; EPPO, 2014|
|Senegal||Present||IIE, 1993; EPPO, 2014|
|Seychelles||Present||IIE, 1993; EPPO, 2014|
|Sierra Leone||Present||IIE, 1993; EPPO, 2014|
|Somalia||Present||IIE, 1993; EPPO, 2014|
|South Africa||Widespread||IIE, 1993; EPPO, 2014|
|-Canary Islands||Present||IIE, 1993; EPPO, 2014|
|Sudan||Present||IIE, 1993; EPPO, 2014|
|Swaziland||Present||IIE, 1993; EPPO, 2014|
|Tanzania||Present||IIE, 1993; EPPO, 2014|
|Togo||Present||IIE, 1993; EPPO, 2014|
|Tunisia||Restricted distribution||IIE, 1993; EPPO, 2014|
|Uganda||Present||IIE, 1993; EPPO, 2014|
|Zambia||Present||IIE, 1993; EPPO, 2014|
|Zimbabwe||Widespread||IIE, 1993; EPPO, 2014|
|USA||Absent, formerly present||NAPPO, 2015; NAPPO, 2016|
|-Florida||Absent, formerly present||NAPPO, 2015; NAPPO, 2016|
Central America and Caribbean
|Puerto Rico||Restricted distribution||NAPPO, 2014|
|Argentina||Restricted distribution||EPPO, 2014; Murúa et al., 2014|
|Brazil||Restricted distribution||EPPO, 2014|
|-Bahia||Restricted distribution||EPPO, 2014|
|-Espirito Santo||Present||Pratissoli et al., 2015|
|-Goias||Present||Czepak et al., 2013; EPPO, 2014|
|-Mato Grosso||Present||Czepak et al., 2013; Wee et al., 2013; EPPO, 2014|
|-Sao Paulo||Present||Bueno et al., 2014|
|Paraguay||Restricted distribution||EPPO, 2014; Murúa et al., 2014|
|Uruguay||Present||Castiglioni et al., 2016|
|Albania||Widespread||IIE, 1993; EPPO, 2014|
|Austria||Present, few occurrences||EPPO, 2014|
|Belgium||Absent, intercepted only||EPPO, 2014|
|Bulgaria||Widespread||****||IIE, 1993; EPPO, 2014|
|Croatia||Absent, formerly present||EPPO, 2014|
|Cyprus||Restricted distribution||IIE, 1993; EPPO, 2014|
|Czech Republic||Eradicated||EPPO, 2014|
|Denmark||Absent, intercepted only||IIE, 1993; EPPO, 2014|
|Estonia||Absent, formerly present||EPPO, 2014|
|Finland||Present, few occurrences||EPPO, 2014|
|France||Restricted distribution||IIE, 1993; EPPO, 2014|
|Germany||Present, few occurrences||IIE, 1993; EPPO, 2014|
|Greece||Widespread||IIE, 1993; Demirumlaut~er, 2012; EPPO, 2014|
|Hungary||Restricted distribution||1951||IIE, 1993; Bozsik, 2007; EPPO, 2014|
|Italy||Restricted distribution||IIE, 1993; EPPO, 2014|
|-Sardinia||Present||IIE, 1993; EPPO, 2014|
|-Sicily||Present||IIE, 1993; EPPO, 2014|
|Latvia||Absent, formerly present||EPPO, 2014|
|Lithuania||Present||Ostrauskas et al., 2002|
|Malta||Restricted distribution||IIE, 1993; EPPO, 2014|
|Moldova||Present||Timus and Croitoru, 2006|
|Montenegro||Present||Radonjic and Hrncic, 2011|
|Netherlands||Eradicated||IPPC, 2007; EPPO, 2014||Absent, pest eradicated (incidental findings), confirmed by survey. Based on long-term annual surveys, 318 survey observations in 2012.|
|Norway||Absent, formerly present||IIE, 1993; EPPO, 2014|
|Poland||Present, few occurrences||IIE, 1993; EPPO, 2014|
|Portugal||Widespread||IIE, 1993; EPPO, 2014|
|-Azores||Present||IIE, 1993; EPPO, 2014|
|-Madeira||Present||IIE, 1993; EPPO, 2014|
|Romania||Widespread||IIE, 1993; EPPO, 2014|
|Russian Federation||Restricted distribution||EPPO, 2014|
|-Russian Far East||Present|
|-Southern Russia||Restricted distribution||EPPO, 2014|
|-Western Siberia||Present||EPPO, 2014|
|Serbia||Restricted distribution||EPPO, 2014|
|Slovakia||Restricted distribution||EPPO, 2014|
|Spain||Widespread||IIE, 1993; EPPO, 2014|
|Switzerland||Restricted distribution||IIE, 1993; EPPO, 2014|
|-Channel Islands||Absent, formerly present||EPPO, 2014|
|-England and Wales||Eradicated||EPPO, 2014|
|Ukraine||Widespread||****||IIE, 1993; EPPO, 2014|
|Yugoslavia (Serbia and Montenegro)||Restricted distribution||IIE, 1993|
|American Samoa||Present||IIE, 1993; EPPO, 2014|
|Australia||Widespread||****||IIE, 1993; EPPO, 2014|
|-Australian Northern Territory||Present||IIE, 1993; EPPO, 2014|
|-New South Wales||Present||IIE, 1993; EPPO, 2014|
|-Queensland||Present||IIE, 1993; EPPO, 2014|
|-South Australia||Present||EPPO, 2014|
|-Western Australia||Present||IIE, 1993; EPPO, 2014|
|Fiji||Present||IIE, 1993; EPPO, 2014|
|Kiribati||Present||IIE, 1993; EPPO, 2014|
|Marshall Islands||Present||IIE, 1993; EPPO, 2014|
|Micronesia, Federated states of||Present||EPPO, 2014|
|New Caledonia||Widespread||IIE, 1993; EPPO, 2014|
|New Zealand||Widespread||IIE, 1993; EPPO, 2014|
|-Kermadec Islands||Present||IIE, 1993|
|Norfolk Island||Present||Holloway, 1977; IIE, 1993; EPPO, 2014|
|Northern Mariana Islands||Present||IIE, 1993; EPPO, 2014|
|Papua New Guinea||Present||APPPC, 1987; IIE, 1993; EPPO, 2014|
|Samoa||Present||IIE, 1993; EPPO, 2014|
|Solomon Islands||Present||IIE, 1993; EPPO, 2014|
|Tonga||Present||IIE, 1993; EPPO, 2014|
|Tuvalu||Present||IIE, 1993; EPPO, 2014|
|Vanuatu||Present||IIE, 1993; EPPO, 2014|
Risk of IntroductionTop of page
H. armigera is listed as an A2 quarantine pest by EPPO (OEPP/EPPO, 1981). Although it is certainly a serious outdoor pest in Mediterranean countries, it has probably reached the limits of its natural distribution in the EPPO region. Quarantine status arises from the risk of introduction into glasshouse crops in northern Europe. EPPO recommends (OEPP/EPPO, 1990) that imported propagation material should derive from an area where H. armigera does not occur or from a place of production where H. armigera has not been detected during the previous 3 months.
Hosts/Species AffectedTop of page The most important crop hosts of which H. armigera is a major pest are cotton, pigeonpea, chickpea, tomato, sorghum and cowpea; other hosts include groundnut, okra, peas, field beans (Lablab spp.), soyabeans, lucerne, Phaseolus spp., other Leguminosae, tobacco, potatoes, maize, flax, a number of fruits (Prunus, Citrus), forest trees and a range of vegetable crops. A wide range of wild plant species support larval development: important species in India include Acanthospermum spp., Datura spp., Gomphrena celosioides and, in Africa, Amaranthus spp., Cleome sp. and Acalypha sp.
See Matthews (1991) and Majunath et al. (1989) for full lists.
Host selection has been summarized by King (1994) and treated in some depth by Fitt (1991), and for all moths by Ramaswamy (1988).
Host Plants and Other Plants AffectedTop of page
|Abelmoschus esculentus (okra)||Malvaceae||Main|
|Acalypha (Copperleaf)||Euphorbiaceae||Wild host|
|Albizia procera (white siris)||Fabaceae||Main|
|Amaranthus (amaranth)||Amaranthaceae||Wild host|
|Arachis hypogaea (groundnut)||Fabaceae||Main|
|Avena sativa (oats)||Poaceae||Main|
|Brassica oleracea var. gemmifera (Brussels sprouts)||Brassicaceae||Other|
|Brassica oleracea var. italica (broccoli)||Brassicaceae||Main|
|Brassica rapa subsp. chinensis (Chinese cabbage)||Brassicaceae||Main|
|Brassicaceae (cruciferous crops)||Brassicaceae||Main|
|Broussonetia papyrifera (paper mulberry)||Moraceae||Main|
|Brugmansia candida (angel's trumpet)||Solanaceae||Other|
|Cajanus cajan (pigeon pea)||Fabaceae||Main|
|Callistephus chinensis (China aster)||Asteraceae||Other|
|Capsicum annuum (bell pepper)||Solanaceae||Main|
|Chenopodium album (fat hen)||Chenopodiaceae||Wild host|
|Cicer arietinum (chickpea)||Fabaceae||Main|
|Commelina benghalensis (wandering jew)||Commelinaceae||Wild host|
|Convolvulus arvensis (bindweed)||Convolvulaceae||Wild host|
|Datura (thorn-apple)||Solanaceae||Wild host|
|Datura metel (Hindu datura)||Solanaceae||Wild host|
|Datura stramonium (jimsonweed)||Solanaceae||Wild host|
|Gaillardia pulchella (Indian blanket)||Asteraceae||Other|
|Glycine max (soyabean)||Fabaceae||Main|
|Gomphrena (globe-amaranth)||Amaranthaceae||Wild host|
|Guizotia abyssinica (niger)||Asteraceae||Other|
|Helianthus annuus (sunflower)||Asteraceae||Main|
|Hordeum vulgare (barley)||Poaceae||Main|
|Hyoscyamus niger (black henbane)||Solanaceae||Wild host|
|Hyptis suaveolens (pignut)||Lamiaceae||Other|
|Lablab purpureus (hyacinth bean)||Fabaceae||Main|
|Linum usitatissimum (flax)||Main|
|Mangifera indica (mango)||Anacardiaceae||Main|
|Medicago sativa (lucerne)||Fabaceae||Main|
|Nicotiana tabacum (tobacco)||Solanaceae||Main|
|Papaver somniferum (Opium poppy)||Papaveraceae||Other|
|Pennisetum glaucum (pearl millet)||Poaceae||Main|
|Phaseolus vulgaris (common bean)||Fabaceae||Main|
|Pisum sativum (pea)||Fabaceae||Main|
|Prunus (stone fruit)||Rosaceae||Main|
|Rosa damascena (Damask rose)||Rosaceae||Other|
|Solanum lycopersicum (tomato)||Solanaceae||Main|
|Solanum melongena (aubergine)||Solanaceae||Main|
|Solanum tuberosum (potato)||Solanaceae||Main|
|Sonchus arvensis (perennial sowthistle)||Asteraceae||Wild host|
|Sorghum bicolor (sorghum)||Poaceae||Main|
|Triticum aestivum (wheat)||Poaceae||Main|
|Vigna unguiculata (cowpea)||Fabaceae||Main|
|Zea mays (maize)||Poaceae||Main|
Growth StagesTop of page Flowering stage, Fruiting stage, Vegetative growing stage
SymptomsTop of page On Cotton
Bore holes are visible at the base of flower buds, the latter being hollowed out. Bracteoles are spread out and curled downwards. Leaves and shoots may also be consumed by larvae. Larger larvae bore into maturing green bolls; young bolls fall after larval damage. Adults lay fewer eggs on smooth-leaved varieties.
Young fruits are invaded and fall; larger larvae may bore into older fruits. Secondary infections by other organisms lead to rotting.
Eggs are laid on the silks, larvae invade the cobs and developing grain is consumed. Secondary bacterial infections are common.
Larvae feed on the developing grain, hiding inside the head during the daytime. Compact-headed varieties are preferred.
Foliage, sometimes entire small plants consumed; larger larvae bore into pods and consume developing seed. Resistant cultivars exist.
Flower buds and flowers bored by small larvae, may drop; larger larvae bore into locules of pods and consume developing seed. Short duration and determinate varieties are subject to greater damage. Less-preferred varieties exist.
Leaves, sometimes flowers attacked by larvae; severe infestations cause defoliation. Less preferred varieties exist.
List of Symptoms/SignsTop of page
|Fruit / external feeding|
|Fruit / internal feeding|
|Fruit / lesions: black or brown|
|Fruit / premature drop|
|Growing point / external feeding|
|Inflorescence / external feeding|
|Inflorescence / internal feeding|
|Leaves / external feeding|
Biology and EcologyTop of page In southern Bulgaria, there are two complete generations a year and a partial third, winter being passed in the pupal stage in the soil. Adults emerge in the first 3 weeks of May and, 2-6 days later (rarely 10), oviposition begins. This period lasts 5-24 days and, within this time, a female may lay up to 3180 eggs (up to 457 in 24 h), singly and mainly at night, on chickpeas, cotton, maize, okras, tobacco, tomatoes, Phaseolus and certain weeds. At 25°C, they hatch in 3 days, but can take 10-11 days in colder weather. The first generation larvae (i.e. the larval progeny of the overwintering generation) appear in May and feed for 24-36 days; those of the second generation feed for 16-30 days, and those of the third generation (at 25-26°C) develop in 19-26 days. When fully fed, the larvae descend to the soil and, after 1-7 days, pupate in an earthen cell, 2-8 cm below the surface. The overwintering pupae remain in the soil for 176-221 days, whereas this stage lasts 13-19 days in the first generation, 8-15 days in August and up to 44 days in colder weather in September. Longevity of adults is about 3 weeks.
In southern France, adults appear from May until the end of October. Some are thought to be migrants and others to have overwintered there. A second generation occurs during summer, and third-generation adults appear in September. Second-generation adults from more northern regions migrate towards the south and Mediterranean Basin in autumn. The principal host on which eggs are laid are maize in south-western France and tomatoes in the Rhone Valley.
In Tunisia, Capsicum, tomatoes, maize and cauliflowers are most frequently attacked. The eggs are laid on plants at or near flowering.
In the former USSR, eggs are laid on weeds during spring and early summer; the developing larvae attack cultivated crops and then cotton flowers in August. Larvae rarely move from one plant to another. About 80% of pupae enter diapause at the beginning of October and overwinter in this state.
In Iran, H. armigera also overwinters as the pupal stage, under the soil surface. At the beginning of May, adults emerge and mate promptly. The females lay eggs on weeds and host plants of economic importance, but normally the first generation feeds on weeds. The oviposition period lasts for about 20 days, during which time each female lays 500-2700 eggs. The incubation period takes 3-4 days in summer and about a week during spring and autumn. The larval period lasts 14-18 days in summer and 17-21 days in autumn. During the growing season, H. armigera produces two to six generations according to the climatic conditions. In the northern part of Iran, the most important cotton-growing area in the country, there are four to six generations annually.
In South Africa, the oviposition period is 10-23 days, with an average of 730 eggs per female (total 1600; maximum per night 480). Hairy surfaces are preferred for oviposition, which is closely linked with the period of bud burst and flower production in most host plants. Eggs hatch in 3 days at 22.5°C, and in 9 days at 17.0°C. The larval period lasts 18 days at 22.5°C and 51 days at 17.5°C, development thresholds being 14 and 36°C; rate of development is also affected by food. Fully grown larvae leave the plant to pupate in the soil at a depth of 3-15 cm. In Zimbabwe, pupation may occur in the tip of a maize cob. The pupa may undergo a facultative diapause, which considerably extends the pupal period. In southern Africa, the minimum pupal period in summer is 12 days, increasing as temperature falls to about 57 days. Emerging female moths must feed before their ovarioles are mature. Average life spans for males and females in South Africa are 9 and 14 days, respectively (8 and 11 days in Zimbabwe).
In South and South-East Asia, development times are generally similar to those in South Africa. Egg incubation has been recorded as 2-5, usually 3, days in India and Western Tanganyika (Tanzania). On eclosion the neonate larva usually eats some or all of the empty eggshell before wandering for some distance and starting to feed on the plant, usually in a secluded place such as a flower, flower bud, or the underside of a leaf. Larger larvae prefer to feed on immature fruiting bodies - these are often hollowed out as in cotton and pulses - but will feed on leaves in their absence; larvae often move about between feeding sites on or between adjacent plants. Moulting, particularly of larval stages 2-4, often takes place on the upper surface of a leaf in full sunlight. The number of larval instars varies from five to seven, with six being most common (Hardwick, 1965). The duration of larval development depends on the temperature (to a maximum of 35°C) and on the quality of the host food. Duration varied from 12.8-21.3 days on maize at temperatures of 24-27.2°C; 15.2-23.8 days on tomato at an average of 24.3°C, and averaged 21.1 days (including a prepupal period of 2.7 days) on cotton flower buds at 21.0-27.0°C. Duration of the larval stage was generally shorter on pigeonpea and soyabean than on cotton and tomato, although there is some inconsistency between authors' results (see King, 1994). The heaviest larvae had fed on cotton (Jayaraj, 1982).
On completion of growth the fully fed larva enters the soil to pupate, at a depth which depends upon the hardness of the soil. It is generally formed at a depth of 2.5 to 17.5 cm, but occasionally in surface litter or at the last feeding site on the plant. After a prepupal stage of 1-4 days, during which the larva becomes shorter and more uniform in colour, it moults into a pupa which turns chestnut-brown after about 24 hours.
H. armigera has a facultative pupal diapause which is induced by short daylengths (11-14 hours/day) and low temperatures (15-23°C) experienced as a larva. The duration of the non-diapausing pupal stage varies from 6 days at 35°C to 30 days at 15°C, about 10-14 days in the field in central India. Diapausing pupae may remain in that state for several months, and durations of over 1 year have been recorded in the laboratory. A summer diapause, in which pupae enter a state of arrested development during prolonged hot, dry conditions, has been recorded in the Sudan (Hackett and Gatehouse, 1982).
The duration of the adult stage depends upon the availability of food, as sucrose or nectar; pupal weight (as fat body content); temperature; and activity, with female moths generally living longer than males. In captivity, longevity varied from 1-23 days for male and 5-28 days for female H. armigera in South Africa (Pearson, 1958).
For further information, see Ditman and Cory (1931), Dominguez Garcia-Tejero (1957), Pearson (1958), Hardwick (1965), Cayrol (1972), Delatte (1973), Ibrahim et al. (1974), Roome (1979), Hackett and Gatehouse (1982), Jayaraj (1982) and King (1994).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Bacillus thuringiensis||Pathogen||Larvae||Sudan; Haryana; Shandong; Uttar Pradesh; Tamil Nadu; Delhi|
|Bacillus thuringiensis aizawai||Pathogen||Larvae|
|Bacillus thuringiensis entomocidus||Pathogen||Larvae|
|Bacillus thuringiensis galleriae||Pathogen||Larvae|
|Bacillus thuringiensis kenyae||Pathogen||Larvae|
|Bacillus thuringiensis kurstaki||Pathogen||Larvae|
|Bacillus thuringiensis shandongiensis||Pathogen||Larvae|
|Bacillus thuringiensis subsp. dendrolimus||Pathogen||Larvae|
|Bacillus thuringiensis thuringiensis||Pathogen||Larvae|
|Bracon brevicornis||Parasite||Larvae||Chad; South Africa||cotton; field crops|
|Bracon hebetor||Parasite||Larvae||Israel; USSR|
|Campoletis chlorideae||Parasite||Larvae||Australia; China; India; India; Andhra Pradesh; Pakistan||Amaranthus viridis; Cicer arietinum; cotton; Papaver hybridum; polyphagous|
|Chelonus insularis||Parasite||Larvae||South Africa||fruit trees|
|Chrysopa formosa||Predator||China; Shandong||cotton|
|Chrysopa intima||Predator||China; Shandong||cotton|
|Chrysoperla carnea||Predator||Andhra Pradesh; China; Portugal; USSR; Tamil Nadu||cotton|
|Coccinella septempunctata||Predator||China; Shandong||cotton|
|Coccinella transversalis||Predator||Australia; Queensland||Citrus|
|Copidosoma truncatellum||Parasite||Cape Verde||beans|
|Cotesia kazak||Parasite||Larvae||Australia; Cape Verde; New Zealand; Queensland||beans; Cajanus cajan; polyphagous; tomatoes|
|Cotesia marginiventris||Parasite||Larvae||Australia; Cape Verde; Fiji; India||Cajanus cajan; polyphagous; tomatoes|
|Cotesia ruficrus||Parasite||Larvae||Australia; Cape Verde||beans; polyphagous|
|cytoplasmic polyhedrosis viruses||Pathogen||Larvae|
|Drino imberbis||Parasite||Larvae||Chad; Mauritius||cotton; tobacco; tomatoes; vegetables|
|Eocanthecona furcellata||Predator||Nebapure and Meena, 2011|
|Erigonidium graminicolum||Predator||China; Shandong||cotton|
|Exorista xanthaspis||Parasite||Larvae||India; Gujarat||sunflowers|
|Glabromicroplitis croceipes||Parasite||Larvae||India; New Zealand|
|Goniophthalmus halli||Parasite||Larvae||Cape Verde; Chad; Kenya; Zimbabwe||Cajanus cajan; Citrus; cotton; tomatoes|
|Harmonia axyridis||Predator||China; Shandong||cotton|
|Helicoverpa armigera nuclear polyhedrosis virus||Pathogen||Larvae|
|Heliothis nucleopolyhedrosis virus||Pathogen|
|Hippodamia variegata||Predator||China; Shandong||cotton|
|Hyposoter didymator||Parasite||Australia; Cape Verde||beans; Cajanus cajan; polyphagous; tomatoes|
|Meteorus laphygmarum||Parasite||Larvae||Cameroon; Chad||cotton|
|Microchelonus blackburni||Parasite||Eggs/Larvae||Haryana; India; Maharashtra||Cajanus cajan; cotton|
|Misumenops tricuspidatus||Predator||China; Shandong||cotton|
|Propylea japonica||Predator||Eggs/Larvae||China; Shandong||cotton|
|Sinophorus xanthostomus||Parasite||New Zealand|
|Sycanus indagator||Predator||Mauritius||tobacco; tomatoes; vegetables|
|Telenomus remus||Parasite||Eggs||Cape Verde||Cajanus cajan; tomatoes|
|Trichogramma brasiliense||Parasite||Eggs||India; Karnataka|
|Trichogramma chilonis||Parasite||Eggs||Cape Verde; China; India; India; Gujarat; South Africa; Shandong||beans; cotton; field crops; Medicago sativa; potatoes|
|Trichogramma dendrolimi||Parasite||Eggs||China; China; Shanxi||cotton|
|Trichogramma evanescens||Parasite||Eggs||Spain; Uzbekistan||cotton|
|Trichogramma fasciatum||Parasite||Eggs||South Africa||field crops|
|Trichogramma perkinsi||Parasite||Eggs||India; South Africa||field crops; polyphagous|
|Trichogramma pretiosum||Parasite||Eggs||India; Indonesia; South Africa; Karnataka|
|Trichogramma semifumatum||Parasite||Eggs||South Africa||field crops|
|Trichogrammatoidea armigera||Parasite||Eggs||Cape Verde||Cajanus cajan; tomatoes|
|Xysticus croceus||Predator||China; Shandong||cotton|
Notes on Natural EnemiesTop of page The important species of natural enemies vary from crop to crop and from country to country. Many more parasitoids have been recorded by workers in a range of countries than it has been possible to include here; those included in the List of Natural Enemies are noted as having been of significant importance, although not necessarily on all crops, in all seasons or locations. Levels of parasitism are in many cases host-related, particularly in the Trichogrammatidae, parasitism generally being higher, and by more species, on sorghum than on other crops. There was a notable lack of transfer of parasitoids from sorghum to pigeonpea where these two crops were intercropped (Manjunath et al., 1989).
The impact of parasitoids on the seasonal abundance of H. armigera is still poorly understood. Few quantitative details from life tables (e.g. Titmarsh, 1985), other than of percentage parasitism, have been published and, except for egg parasitoids, rates are usually low. Intermediate rates of parasitism have been recorded for some of the Tachinidae, but these generally occur too late in the larval stage to reduce host damage. Reviews for Europe and a number of areas in Asia are provided by King and Jackson (1989), for Africa by van den Berg et al. (1988), and for Sri Lanka and Australia by Waterhouse and Norris (1987). In most areas, species of Telenomus and Trichogrammatidae (Trichogramma and Trichogrammatoidea) are important egg parasitoids, and larvae are parasitized by at least one species each of Braconidae, Ichneumonidae and Tachinidae.
The relative importance of parasitoids and predators varies betwen localities and crops. For example in Kenya, van den Berg (1993) found that predators, chiefly Anthocoridae and Formicidae, suppressed H. armigera on sunflower, maize, sorghum and cotton, but parasitism was low. In contrast, in northern Tanzania, parasitism was the major cause of mortality on sorghum, cotton and a weed (Cleome sp.), but the importance of the different species of parasitoid varied with host plant (van den Berg et al., 1990). The predators of H. armigera have generally been inadequately studied (exceptions include Bishop and Blood, 1977, 1980, 1981 and Room, 1979 in Australia; Cock et al., 1989, 1991 in Kenya) and there is little quantitative information on their impact on populations. Predators include Anthocoridae and Chrysopidae feeding on eggs and predatory Hemiptera and Formicidae on eggs and larvae. Those predators included in this summary either have been widely reported to prey on H. armigera or have been noted as of particular local importance. As in the case of native species, the highly mobile and polyphagous habits of this pest militate against the establishment and impact of all natural control agents. Augmentative releases of Chrysopa carnea against H. zea and Heliothis virescens have been attempted in the USA (Ridgeway et al., 1977) but the cost of mass-producing the predator, as with parasitoids bred for mass release programmes, has not hitherto been economically viable for H. armigera.
Records of nematode parasites, usually Mermithidae, are available from all regions where inventories of natural enemies are available, however high rates of parasitism occur only sporadically when conditions are favourable. There is some evidence that, in India, they may be important in suppressing early season populations on wild hosts (e.g. Acanthospermum hispidum) and low-growing crops such as groundnut on alfisols (Bhatnagar et al., 1985).
There has been some success in the use of pathogens, Bacillus thuringiensis and Helicoverpa armigera nuclear polyhedrosis virus (HaNPV) preparations, applied like insecticides to manage larval populations of H. armigera. However the relatively high cost, rapid inactivation by ultraviolet light, often slow or poor field performance and, in the case of HaNPV, difficulty in obtaining consistently high levels of purity and virulence necessary to achieve satisfactory control, have limited their usefulness. The increasing prevalence of resistance to insecticides and awareness of environmental concerns has given a new impetus to the development of suitable microbials to include in IPM strategies for H. armigera (King, 1994).
More comprehensive lists of parasitoids and predators by country or region are given by the authors who contributed papers to the Workshop on the Biological Control of Heliothis held in New Delhi in 1985 (King and Jackson, 1989). The role of natural enemies in the control of H. armigera, mainly in cotton, has been reviewed by King (1994). The worldwide distribution, abundance and potential for biocontrol of the natural enemies of economically important Heliothis and Helicoverpa spp. have been reviewed by King et al. (1982), King and Coleman (1989) and King and Jackson (1989).
Means of Movement and DispersalTop of page The importance and success of H. armigera is in large measure due to its well-developed survival strategies, diapause and dispersal, which enable it to exploit food sources separated both by unfavourable times and by distance, and thereby also to escape its natural enemies. H. armigera is effectively a facultative migrant, not displaying typical migratory behaviour, but responding largely to local environmental cues and undertaking either short or longer distance flight in directions largely governed by prevailing weather systems (Fitt, 1989). Innately, the disposition to disperse is governed by reproductive maturity, so that in more transient habitats where dispersal has greater survival value, the length of the pre-reproduction period is greater than in less extreme habitats (Colvin, 1990); in these habitats, such as in India, the tendency to fly was moderated chiefly by feeding which reduced the pre-maturation period.
Adults can migrate over long distances, borne by wind, for example from southern Europe to the UK (Pedgley, 1985). Movement in international trade is mainly on ornamental plants and on cut flowers; also in cotton bolls and in tomato fruits.
For references and further information refer to Pedgley (1985), Farrow and Daly (1987), Pedgley et al. (1987), Fitt (1989), Colvin (1990), Riley et al. (1992), King (1994).
ImpactTop of page Introduction
H. armigera, like its close relatives H. zea and Heliothis virescens in the New World, is a pest of major importance in most areas where it occurs, damaging a wide variety of food, fibre, oilseed, fodder and horticultural crops. Its considerable pest significance is based on the peculiarities of its biology - its mobility, polyphagy, rapid and high reproductive rate and diapause make it particularly well adapted to exploit transient habitats such as man-made ecosystems. Its predilection for the harvestable flowering parts of high-value crops including cotton, tomato, sweetcorn and the pulses confers a high economic cost, and socio-economic cost in subsistence agriculture, due to its depredations. However, regional and even relatively local differences in host preference can give rise to differences in pest status on particular crops; this was shown by populations in northern and southern India where severe infestations of cotton are only a relatively recent event.
H. armigera has been reported causing serious losses throughout its range, in particular to cotton, tomatoes and maize. For example, on cotton, two to three larvae on a plant can destroy all the bolls within 15 days; on maize, they consume grains; and on tomatoes, they invade fruits, preventing development and causing falling.
Monetary losses result from the direct reduction of yields and from the cost of monitoring and control, particularly the cost of insecticides. In Australia, Wilson (1982) estimated total Australian losses at $A 23.5 million; with increases in the prices of insecticides and the replacement of the cheaper pyrethroids with more expensive alternatives to counter pyrethroid resistance, Twine (1989) has estimated that costs in Queensland alone would have increased to about $A 25 million annually.
In India, where H. armigera commonly destroys over half the yield of pulse crops, pigeon pea and chickpea, losses were estimated at over $US 300 million per annum (Reed and Pawar, 1982), while in the late 1980s losses of both pulses and cotton were estimated to exceed $US 500 million, with an additional $US 127 million spent on insecticides on these two crops annually (KN Mehrotra, Indian Agricultural Research Institute, New Delhi, unpublished data, 1987/88). Following the rapid upsurge of pyrethroid resistance, and reduced effectiveness of other insecticide groups in H. armigera (Dhingra et al., 1988; McCaffery et al., 1989) these figures will certainly need to be revised upwards.
Oerke et al. (1994) reported that H. armigera is an economically important pest or a key pest in Africa, Asia, Europe and the former USSR, and Oceania. Previously, Ridgway et al. (1984) had reported also that H. armigera was partly responsible for a major portion of cotton crop losses.
In Africa, H. armigera can reduce yields substantially. In the Côte d'Ivoire, between 1978 and 1983, cotton crop loses in the south of the country were primarily due to H. armigera and were ca 60% (Moyal, 1988). In Zimbabwe, potential crop losses due to H. armigera were 1175 kg/ha (Gledhill, 1976). While H. armigera has now been contained as a pest on cotton in Zimbabwe, it is important in Tanzania where the economic loss of cotton was estimated at over $US 20 million (Reed and Pawar, 1982).
In Andhra Pradesh, India, problems in controlling H. armigera were first encountered in 1987. More than 30 insecticide treatments were applied, yet the average yield fell from 436 kg/ha in 1986/87 to 186 kg/ha in 1987/88. This was a reduction of 61% (Armes et al., 1992). In Thailand, H. armigera has been the principal cotton pest since the mid-1960s. Losses due to H. armigera were at least 31% in 1975-79 (Mabbett et al., 1980). In China, losses due to H. armigera larvae increased with plant age. Crop losses were substantial regardless of soil fertility (Sheng, 1988). The damage threshold, 7.5 kg/ha, was reached at 35 egg clusters/100 plants. Integrated pest management reduced H. armigera infestations from 1.6 to 0.1% in Jiangsu between 1976 and 1982 (Jin, 1986).
In the EPPO region, H. armigera is of great economic importance in Israel, Morocco, Portugal, former USSR and Spain, and of lesser importance in the other countries where it is established. Despite extensive spread in Greece, H. armigera only causes periodic damage to cotton.
Chickpeas and Other Crops
In India, chickpea is the most important pulse crop and is grown on 7.3 million hectares in various agro-climatic conditions. Although its yield potential is 2.5-3 t/ha, the average yield is only ca 0.8 t/ha. The extent of losses caused by H. armigera varies from region to region and depends upon climate and crop intensity. However, a monetary loss of 203 crore rupees annually is estimated.
Changes in sowing date have had a considerable influence on pod damage and seed yield of chickpea. Pod damage due to H. armigera increased as sowing dates grew later. At five different sowing dates, % pod damage was 5.8, 8.1, 14.9, 18.2 and 26.2% while corresponding seed yields of 2452, 2409, 1859, 1439 and 1010 kg/ha, respectively, were recorded. The co-efficients of correlation between sowing date and pod damage and between pod damage and seed yield were significant (Saxena et al., 1998). The larval population of H. armigera on chickpea was ca four times higher at dense spacing (33 plants/m²) than at wide spacing (3 plants/m²) (Yadava et al., 1998).
Chickpea yields have been shown to increase following control treatments. The application of nuclear polyhedrosis virus reduced larval populations by 26.8% and pod damage by 36.6% and increased yields by 72% compared with untreated plots (Bhagwat and Wightman, 1998).
Damage has been reported in India on potatoes, sunflowers, Guizotia abyssinica, pigeon peas and cotton. Crop losses of 10-100% have been estimated for potatoes in India. In studies over three seasons, between 1982 and 1985, on four varieties average losses of 0.34% were recorded. Based on the average potato yield for India of 15.8 t/ha, the loss rate was 2.1% (Parihar and Singh, 1988).
An outbreak of this noctuid occurred on young Pinus radiata in New Zealand in 1969 and 1970, when the larvae consumed more than 50% foliage of about 60% of trees.
Detection and InspectionTop of page The feeding larvae can be seen on the surface of plants but they are often hidden within plant organs (flowers, fruits etc.). Bore holes and heaps of frass (excreta) may be visible, but otherwise it is necessary to cut open the plant organs to detect the pest.
Similarities to Other Species/ConditionsTop of page In Asia, H. armigera may sometimes be confused with H. assulta (a smaller, yellower species) on pulses, although the latter is seldom seen on pigeonpea and never on chickpea in India. In Sudan, H. armigera may be confused with H. fletcheri (which has a row of pale flecks in the forewing postmedial) on sorghum and some other crops. On rearing to adult, the species may be clearly distinguished.
In Europe, identification of all stages will be difficult should very similar American (H. zea) or Australian (H. punctigera) species be introduced and become established. Separation of the adult from similar species is most reliably done by reference to the male genitalia (Hardwick, 1965): the middle spine on the most basal coil of the everted aedeagus vesica is larger than all other spines.
Prevention and ControlTop of page
H. armigera is a pest of major importance in most areas where it occurs, damaging a wide variety of food, fibre, oilseed, fodder, commodity and horticultural crops. Its major pest status is rooted in its mobility, polyphagy, high reproductive rate and diapause, all of which make it particularly well adapted to exploit transient habitats such as man-made agro-ecosystems. Its predilection for harvestable parts of essential food and high-value crops like cotton, tomato, pulses and tobacco confers a high economic cost to its depredations. The high level of control required under these circumstances, and the absence, in most situations, of adequate natural control means that chemical, or at best integrated control methods usually need to be adopted.
In view of the need to make use of and exploit the existing spectra of natural enemies and to reduce excessive dependence on chemical control, particularly where there is resistance to insecticides, various IPM programmes have been developed in which different control tactics are combined to suppress pest numbers below a threshold. These vary from the judicious use of insecticides, based on economic thresholds and regular scouting to ascertain pest population levels, to sophisticated systems, almost exclusively for cotton, using computerized crop and population models to assess the need, optimum timing and product for pesticide application. The SIRATAC system, developed in Australia during the 1980s, and its subsequent derivatives fall into this category (Room, 1979, 1983; Hearn et al., 1981). A major constraint to the development of IPM for H. armigera, particularly on cotton, has been the need to deal with a complex of pests where control needs may be irreconcilable, as for example in the characteristics of the cotton plant which can either be unfavourable to H. armigera or to jassid pests in terms of leaf hairiness, and in the withholding of early season applications to encourage the build-up of natural enemies against the need to control sucking pests which can be severe on young plants.
Owing to its strongly dispersive habit, efforts to regulate the influx of H. armigera into crops is generally not a viable option. Some cultural methods, such as an enforced 'close' season, may be regarded as regulatory, but to be effective these will depend on strict compliance, geographical isolation and the absence of a significant alternative wild host population in the area.
Another aspect of regulatory control is in the use of insecticides against which H. armigera has severe incipient resistance, and of 'hard' insecticides which are particularly damaging to natural enemies. An example of this is the resistance management strategy developed in Australia, where the use of pyrethroids was confined to particular phases in the cotton-growing season, principally to minimize selection for resistance.
Cultural Control and Sanitary Methods
Cultural manipulations of the crop or cropping system and land management have been tried as tactics to manage H. armigera populations. Trap cropping and planting diversionary hosts have been widely applied and recommended in the past, although with limited success. In the case of cotton, the diversionary hosts maize and sorghum had too short an attractive period to sustain populations; the tendency of these and earlier-planted crops to augment or create infestations were major disadvantages. The importance of ploughing cotton stubble to reduce overwintering populations of pyrethroid-resistant H. armigera was stressed by Fitt and Forrester (1987), and post-harvest cultivation to destroy pupae of bollworms has received considerable attention in the USA. However, all in situ cultural control tactics (including area-wide management of early season populations on wild hosts, as advocated by several workers in the USA for American species; Stadelbacher, 1982), and the concept of a close season during which food plants are denied for over one generation, would seem to be largely invalid where the immigration of adults into the protected habitats is the key consideration.
One indirect cultural method which could be included under this heading is the regulation of crop agronomy, variety (such as the okra-leaved varieties of cotton), spacing and fertilizer regimes to render the crop, and thus target larvae, more accessible to insecticides or microbial formulations applied by conventional means.
The planting of crop varieties that are resistant or tolerant to H. armigera has received major attention, particularly for cotton, pigeonpea and chickpea. This is a tactic of considerable importance within IPM systems. Many crop species possess some genetic potential which can be exploited by breeders to produce varieties less subject to pest damage; this can take the form of antibiosis (unpalatability), antixenosis (non-preference) and tolerance. However, where there is a pest complex, interactions may not always be favourable. For example, fewer eggs were laid on plants having the glabrous leaf character in cotton, however both larval survival and susceptibility to jassid attack were higher. Varieties of chickpea, groundnut and pigeonpea showing varying degrees of resistance have been developed at ICRISAT in India, some of which have been successfully used by farmers.
In recent years, genetic engineering techniques have enabled genes carrying the toxic element of Bacillus thuringiensis to be introduced into crops such as cotton and tomato. Although the technique is still very much in its early stages, transgenic crop varieties offer considerable promise for use in IPM systems against H. armigera. As with the use of all resistant crop varieties, however, care still needs to be taken to avoid excessive selection pressure against the resistance factor, so that in such systems a mixture of both resistant and susceptible varieties is often recommended to lessen this.
While IPM strategies are generally geared to provide a regime in which maximum feasible advantage is taken of local biological control agents, their unassisted suppression of H. armigera populations to below an economic threshold without the use of insecticides would be a major advantage, both in ecological and economic terms, particularly if this was sustainable. To this end, substantial efforts have been made either to introduce exotic natural enemies or to augment existing populations of parasitoids and predators to achieve satisfactory levels of control. Because of the need to produce very large numbers of parasitoids or predators simultaneously and economically, emphasis has been placed on Trichogramma spp. which are most amenable to mass rearing. Although these and a number of other parasitic species have been field evaluated against H. armigera, results have not so far been encouraging, especially in agrosystems where insecticide applications against H. armigera or other pests are consistently necessary.
There have been attempts to enhance mortality due to natural enemies by the introduction of species that might complement existing natural enemies or be superior to them (reviewed by Waterhouse and Norris, 1987). Attempted introductions have included parasitoids of Heliothis virescens and Helicoverpa zea from the Americas as well as species from other parts of the range of H. armigera. Few of these have been successful. Trichogramma pretiosum and T. perkinsi from the USA are reported to have become established in Indonesia and South Africa, respectively. Other successful establishments are: India (Chelonus blackburni, Eucelatoria bryani, both from the USA, and Bracon kirkpatricki from Kenya); Fiji (Cotesia marginiventris, also from the USA); New Zealand (Glabrobracon croceipes from the USA); Western Australia (Cotesia kazak and Hyposoter didymator, both from Europe). None of these introductions appears to have had a significant beneficial impact. However, the introduction of Cotesia kazak from Greece into New Zealand, where there were no native parasitoids of this pest, resulted in substantial parasitism but because of the low tolerance for insect damage in tomato crops, insecticides are still needed.
The relative specificity, potential activity, environmental safety and immunity to insecticides have made microbial pesticides a favoured component of IPM strategies, and considerable efforts have been made to develop the most promising agents, Bacillus thuringiensis and Helicoverpa armigera nuclear polyhedrosis virus (HaNPV) into commercially viable products. Present and active under natural conditions, both these agents, but particularly HaNPV, have some impact on H. armigera populations, although seldom reaching the epizootic proportions necessary to achieve effective control. Field tests with artificially produced Bt and HaNPV have so far had only limited success, mainly because of rapid degradation by UV light, insufficient titres ingested by larvae, and lack of virulence. However work is continuing to overcome these constraints stimulated by increasing resistance to insecticides and awareness of the environmental threats they pose.
The whole subject of biological control of H. armigera is treated in considerable detail in King and Jackson (1989).
In most cases where H. armigera attacks high-value or staple crops, its control with insecticides, alone or within the context of an IPM programme, will be necessary. While it is clear that economic thresholds need to be carefully applied for best results, in many countries where resources are limiting or the advantages of IPM are poorly understood, insecticides are applied on an ad hoc basis with ensuing poor results and often entry onto the 'insecticide treadmill', where increasing numbers of applications achieve diminishing returns on their investment.
Most insecticide applications are targeted at the larval stages, but as these are only really effective when larvae are small, the need to scout for eggs and spray soon afterward is paramount. Young larvae are difficult to find, and older larvae soon burrow into the floral organs where they become less accessible to contact insecticides, require higher doses to kill and cause direct economic loss. Moreover, resistant larvae were still susceptible while less than 4 days old, so that targeting of neonates is essential in areas where resistant populations are present (Daly, 1988).
The considerable selection pressure which H. armigera has experienced, particularly to the synthetic pyrethroids which were used predominantly in the early 1980s, has resulted in the development of resistance to the major classes of insecticides in many of the areas where these have been used. Field failures resulting from pyrethroid resistance have been reported from Australia, Thailand, Turkey, India, Indonesia and Pakistan. Insecticide resistance management strategies have been aimed either at preventing the development of resistance, or containing it. All rely on a strict temporal restriction in the use of pyrethroids and their alternation with other insecticide groups to minimize selection for resistance. And while the strong propensity of H. armigera to disperse confers the advantage of diluting resistant populations through the influx of susceptible insects from unsprayed hosts, the same tendency ensures that the genes for resistance are spread more widely than their area of origin (Forrester et al., 1993).
Pyrethroid resistance in H. armigera may be conferred through three separate mechanisms: detoxification by mixed-function oxidases (metabolic resistance), nerve insensitivity, and delayed penetration. Metabolic resistance may be inhibited by piperonyl butoxide and other synergists, providing a (costly) means whereby the use of pyrethroids might be prolonged in populations where this is the principal mechanism.
Early Warning Systems
The importance of dispersive and migratory behaviour in the biology of H. armigera suggests that monitoring of these movements could provide an early warning of its invasion of an area or crop. Although work on long-distance movement using radar, backtracking and other techniques indicated that moths were able to (and often did) cover large distances, their occurrence in significant numbers at a particular location could seldom be predicted with any certainty. Changes in catch numbers in light and pheromone traps showed characteristic patterns of abundance for different locations in India (Srivastava et al., 1992), but the relationship between trap catch and subsequent egg or larval populations in a susceptible crop was usually variable to poor, with numbers captured differing markedly between traps separated by only a few tens of metres, although it was closest when moth densities were low and at the beginning of the seasonal cycle. Trapping H. armigera is thus only useful as a qualitative measure indicating the start of an infestation or a migratory 'wave front', indicating the need to begin scouting for immature stages in the crop.
Models are conceptual or mathematical devices which aim to simulate natural processes. As pest management tools they are used to predict or establish the optimal tactics required to achieve economic control of that pest, within the constraints of the model. Models for the management of H. armigera have been mostly restricted to cotton in Australia (and in the USA against related bollworms in cotton). They include the SIRATAC system (see IPM Programmes), and later, more sophisticated models such as HEAPS, which gives greater attention to biological parameters of H. armigera including adult movement, and take account of the presence of non-crop hosts in a region. The model informs of the optimum timing and type of insecticide to be applied (Zalucki et al., 1986; Dillon and Fitt, 1990). Because of their specificity to particular, uniform cropping environments, sophisticated models have been built for H armigera only as a pest of cotton, where the extensive scale and high value of the crop means that farmers are most willing and economically able to abide by their strictures and gain most advantage from their use.
Field Monitoring and Economic Threshold Levels
The ascertaining and utilization of economic thresholds is implicit in the evolution of an IPM programme. Field monitoring of pest populations is necessary to determine whether the threshold has been exceeded and control measures should be taken. The economic threshold of pest density, where the value of expected benefit derived from it exceeds the cost of implementation, depends on a knowledge of the relationship between population density and economic loss. However, it is often difficult to obtain precise data on this relationship because it is rarely simple, and many extraneous factors, both socio-economic and environmental, may influence it.
Action thresholds based on egg numbers have been used successfully as the basis for control decisions in cotton since 1961 in Malawi and Zimbabwe, where spraying was recommended at an average of one egg per two plants in twice-weekly counts (Matthews and Tunstall, 1968), while in the Sudan Gezira over two eggs or larvae per 18 plants (Haggis, 1982) and in Australia two eggs per metre row (Wilson, 1981) were used as thresholds. These thresholds are low and it has been argued by Kabissa (1989) that some damage may actually increase yields.
Trapping of adult moths has sometimes been used to assess the need to subsequent spraying, although for H. armigera this has been at best supplementary to scouting for eggs or larvae, as the relationship between catch and later larval populations is often poor (e.g. Rothschild et al., 1982).
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