Helicoverpa armigera

Morphology, life cycle, damage caused and control

Noctuidae: a family of owlet moths

‘Noctuca’ (‘nox’= night) latin = little owlet or night owl

family of dull coloured moths

active at night
 Helicoverpa armigera

A moth

• cotton bollworm
• corn earworm
• Old World (African) bollworm
Kingdom: Animalia
• scarce bordered straw
Phylum: Arthropoda
• gram pod borer
Class: Insecta
Order: Lepidoptera
Superfamily: Noctuoidea
Family: Noctuidae
Genus: Helicoverpa
Species: H. armigera
Distribution
Cosmopolitan

This species comprises two sub-species:

Helicoverpa armigera armigera is widespread in central and southern Europe, temperate

Asia and Africa;

Helicoverpa armigera conferta is found in Australia, and Oceania.

The former sub-species has also recently been confirmed to have successfully invaded Brazil
and has since spread across much of South America and reached the Caribbean. It is a
migrant species, able to reach Scandinavia and other northern territories.
Morphology
Adult

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.

Hind wings are pale-straw colour with a broad dark-brown border that contains a paler patch;
(this is a distinctive feature in adults)
they have yellowish margins and strongly marked veins and a dark, comma-shaped marking in the middle.
Antennae are covered with fine hairs.
Egg
Eggs are spherical-shaped with diameter of about 0.4mm to
0.6mm.
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
Newly laid eggs are characterised by yellowish white colour,
glistening at first After a few days colour changes from yellowish
white to dark brown before hatching.
When eggs are close to hatching, black head capsule is visible
through the eggshell.
laid on plants which are flowering, or are about to produce
flowers.
Larva

Larvae of H. armigera undergo 6 or 7 developmental growth stages before developing into pupal stage.
The neonate (newly hatched) larvae:
may grow up to 7mm long, with yellowish-white to reddish-brown colour and dark-spotted body.
The dark spots on the body is due to the dark coloured spiracles and tuberculate bases.
no prominent markings;
head, prothoracic shield, supra-anal shield and prothoracic legs are very dark-brown to black;
prolegs are present on the third to sixth, and tenth, abdominal segments.
Neonate larvae usually eat soft leaves, flowers or flower buds, moving about their feeding sites.
Moulting occurs when larvae are about to develop into following larval stage.
Subsequent larval instars may grow up to 40mm long,
showing distinctive characteristics:
head is brown and mottled;
prothoracic and supra-anal plates and legs are pale brown;
claws and spiracles are black;
the skin surface consists of close-set, minute tubercles. and
crochets on prolegs are arranged in 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.
Feed on a wide range of plants, including many important cultivated crops.

It is the late larval stage that causes damage to crops by consuming the affected plant parts. Larvae shift from
foliar feeders to developing seeds and fruits as larval instar development progresses (Reed & Pawar 1982).

It is a major pest in cotton and one of the most polyphagous and cosmopolitan pest species.
 Pupa

When the last larval instars are about to become pupae, the larvae will burrow and enter the soil to
pupate.

Pupae are characterised with:

14-18mm in length,
mahogany brown colour
smooth surface and
rounded anterior and posterior ends
2 tapering, parallel spines at posterior tip

Not all pupae will develop into adults without pupal diapause. The period of pupal diapause varies and
highly dependent on temperature and photoperiod (Hackett & Gatehouse 1982).
 Life cycle of Helicoverpa armigera
Life cycle of H. armigera takes about 73 days (in absence of pupal diapause) in spring or autumn
with average temperature of 16 – 18°C from eggs to adults, whereas in summer the life cycles take
about 34 days with average temperature of 28°C from eggs to adults (Zalucki et al. 1986).
Life cycle
The female cotton bollworm can lay several hundred eggs, distributed on various parts of the plant. Under favourable
conditions, the eggs can hatch into larvae within three days and the whole lifecycle can be completed in just over a
month.
The eggs are spherical and 0.4 to 0.6 mm in diameter, and have a ribbed surface. They are white, later becoming
greenish.
The larvae take 13 to 22 days to develop, reaching up to 40 mm long in the sixth instar. Their colouring is variable,
but mostly greenish and yellow to red-brown. The head is yellow with several spots. Three dark stripes extend along
the dorsal side and one yellow light stripe is situated under the spiracles on the lateral side. The ventral parts of the
larvae are pale. They are rather aggressive, occasionally carnivorous and may even cannibalise each other. If
disturbed, they fall from the plant and curl up on the ground.
The pupae develop inside a silken cocoon over 10 to 15 days in soil at a depth of 4–10 centimetres (1.6–3.9 in), or in
cotton bolls or maize ears.
Host plants
The cotton bollworm is a highly polyphagous species.
The most important crop hosts are:
tomato,
cotton,
pigeon pea,
chickpea,
rice,
sorghum, and
cowpea.
Other hosts include groundnut, okra, peas, field beans, soybeans, lucerne, Phaseolus spp., other Leguminosae,
tobacco, potatoes, maize, flax, Dianthus, Rosa, Pelargonium, Chrysanthemum, Lavandula angustifolia, a number
of fruit trees, forest trees, and a range of vegetable crops.
In Russia and adjacent countries, the larvae populate more than 120 plant species, favouring Solanum, Datura,
Hyoscyamus, Atriplex, and Amaranthus genera.

Economic significance[
The greatest damage is caused to cotton, tomatoes, maize, chick peas, alfalfa, and tobacco. The economic
threshold of harmfulness in central Asia is three to five larvae per hundred plants of long-staple cotton and
eight to 12 larvae per hundred plants on medium-staple cotton.[4] In cotton crops, blooms that have been
attacked may open prematurely and stay fruitless. When the bolls are damaged, some will fall off and others
will fail to produce lint or produce lint of an inferior quality. Secondary infections by fungi and bacteria are
common and may lead to rotting of fruits. Injury to the growing tips of plants may disturb their development,
maturity may be delayed, and the fruits may be dropped.

Control measures include the use of NOCTOVI adulticide attract and kill formulation, growing of resistant
varieties, weeding, inter-row cultivation, removing crop residues, deep autumn ploughing, winter watering to
destroy the pupae, the use of insecticides or biological control through the release of entomophages such as
Trichogramma spp. and Habrobracon hebetor. Monitoring is possible by the use of sex pheromone traps.
Development of Bt cotton (genetically modified to produce Bacillus thuringiensis toxin) improved yields of lint
Larva on Helicoverpa armigera (cotton bollworm); young tomatoes are invaded and fall, larger
tomato larvae may bore into older fruits.
Larvae on Helicoverpa armigera (cotton bollworm); larvae (two colour forms) on pearl millet.
pearl
millet
Larvae Helicoverpa armigera (cotton bollworm); larvae on pigeon pea pod.
on
Larva on Helicoverpa armigera (cotton bollworm); larva feeding at base of cotton flower.
cotton
flower
Management
Sampling: Traps with the female sex pheromone are examined from mid-April and on, the signal for chemical
treatments being four days after 3-4 males are trapped during two consecutive examinations.
Plant resistance: Partial resistance to the pest has been noted in some varieties of the major crops but none appear to
be completely immune.
Horticultural control: The removal of susceptible crop residues and weeds and deep plowing in autumn or winter,
intended to destroy the overwintering insects.

Genetically improved crops: The sowing of Bt cotton (genetically modified cotton that produces the Bacillus
thuringiensis (Bt) toxins (“Bollgard” and “Ingard”) can greatly reduce pesticide applications. However, the pest is
developing resistance to Bt in Australia.

Chemical control: Organochlorides, pyrethroids, carbamates and organophosphates were formerly in use, but the pest
has developed considerable resistance to them. One approach is to rotate these chemicals every few weeks during
the season. Preparations of B. thuringiensis are effective during the early part of the season. Spinosad is still
effective. A novel method is to apply “Noctovi”, a product that attracts bollworms from up to 50 meters, integrated with
a pesticide, to kill the pests.

Biological conttrol: Several hymenopterous parasitoids, including Hyposoter didymator and Bracon hebetor attack
the pest in Israel. Other antagonists include Hemipteran predators like Chrysopa carnea Stephens Chrysopidae and
Orius spp. (Anthocoridae), but their effectiveness is often negated by pesticide applications. Coccinellidae are
important predators in Egypt. The egg parasitoids Trichogramma spp. (Trichogrammatidae) and H. hebetor attack the
pest in Syria. In Australia the Heliothis nuclear polyhedrosis virus and B. thuringiensis formulations control H.
armigera, when their action complement high levels of egg parasitism by Trichogramma. A densovirus was recently
isolated from H. armigera, but its effect is unknown.

Control: The pest can be controlled by spraying 0.02% malathion, endosulphan, endrin, monocrotophos or carbaryl at 15 day interval commencing
flowering stage. Dusting with BHC or carbaryl 10-15% also helps.
synthetic insecticides including organophosphates, synthetic pyrethroids and biorational compounds are the main method for H.
armigera control in different parts of the world. This wide use of pesticides is of environmental concern and has repeatedly led to the
development of pesticide resistance in this pest. Furthermore, the deleterious effects of insecticides on nontarget organisms including
natural enemies are among the major causes of pest outbreaks. It is therefore necessary to develop a novel strategy to manage
population of H. armigera and reduce the hazardous of synthetic chemicals.
The common trend towards reducing reliance on synthetic insecticides for control of insect pests in agriculture, forestry, and human
health has renewed worldwide interest in integrated pest management (IPM) programmes. IPM is the component of sustainable
agriculture with the most robust ecological foundation. IPM not only contributes to the sustainability of agriculture, it also serves as a
model for the practical application of ecological theory and provides a paradigm for the development of other agricultural system
components. The concept of IPM is becoming a practicable and acceptable approach among the entomologists in recent past all over
the world and focuses on the history, concepts, and the integration of available control methods into integrated programmes. However,
this approach advocates an integration of all possible or at least some of the known natural means of control with or without insecticides
so that the best pest management in terms of economics and maintenance of pest population below economic injury level (EIL) is
achieved.
Fundamental of effective IPM programmes is the development of appropriate pest management strategies and tactics that best interface
with cropping system-pest situations. Depending on the type of pest, however, some of the primary management strategies could be
selected. In the case of H. armigera, several management tactics should be considered to implement a comprehensive integrated
management. Potential of some of the control tactics to reduce population density of H. armigera in different cropping systems were
evaluated by several researchers and attempts have been made to develop integrated management approach for H. armigera using
host plant resistance [2, 4, 6, 11] including transgenic Bt crops [16], biological control (predators and parasitoids) [17], interference
methods including sex pheromones [18], biopesticides (especially commercial formulations of Bacillus thuringiensis) [19], cultural
practices (including appropriate crop rotations, trap crops, planting date and habitat complexity) [20] and selective insecticides
[21]. Likewise there remains a need for ongoing research to develop a suite of control tactics and integrate them into IPM systems for
sustainable management of H. armigera in cropping systems. Keeping this in view, integration of these methods based on the ecological
data especially thermal requirements of this pest and its crucial role in forecasting programme of H. armigera could lead a successful
integrated management for this pest in soybean cropping systems.
As discussed above, integrated management is typically problematic in cropping systems, especially in the case of H. armigera on
soybean. However, our intent in this section is not to develop an exhaustive review of all resources that may possibly contribute to more
effective pest management for the future, but to select several topic areas that will make essential contributions to sustainable soybean
cropping systems. Although the past research focused on developing various pest management tactics that would be packaged into an
integrated pest management strategy, we have selected several types of resources for our discussions, realizing that there are other
resources can be used in developing IPM programmes. Furthermore, to generate a comprehensive management programme, we
present our perspectives on future research needs and directions for sustainable management of this pest in soybean cropping systems
such as tri-trophic interactions [22], importance of modeling of insect population [23], crucial role of forecasting and monitoring
programmes in IPM [24], interactions among different management tactics in IPM [25] and significance of biotechnology and genetically
modified plants in IPM. Therefore, considering the importance of H. armigera in successful production of soybean, this review intends to
provide an appropriate document to the scientific community for sustainable management of H. armigera in soybean cropping systems.
Hosts/Species Affected
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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.

Symptoms
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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.

On Tomatoes

Young fruits are invaded and fall; larger larvae may bore into older fruits. Secondary infections by other organisms lead to rotting.

On Maize

Eggs are laid on the silks, larvae invade the cobs and developing grain is consumed. Secondary bacterial infections are common.

On Sorghum

Larvae feed on the developing grain, hiding inside the head during the daytime. Compact-headed varieties are preferred.

On Chickpea

Foliage, sometimes entire small plants consumed; larger larvae bore into pods and consume developing seed. Resistant cultivars exist.

On Pigeonpea

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.

On Groundnut

Leaves, sometimes flowers attacked by larvae; severe infestations cause defoliation. Less preferred varieties exist.
Impact
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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.

Crop Losses

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.

Cotton

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 Inspection
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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/Conditions

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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.
Helicoverpa armigera (Hübner)
Taxonomic placing: Insecta, Holometabola, Lepidoptera, Noctuidae.
Common name: African bollworm, cotton bollworm, corn earworm, Old World bollworm.
Geographical distribution: Between north and south 45° latitudes, recently also found in Brazil. CIE Map #15 (revised), 1993.

Morphology: Adult body is 12-20 mm in length, female forewings yellow to brown, green-gray in males, with a brown spot
near the anterior margin. Hindwings pale yellow with a brown band at the posterior edges and a dark round spot in the middle.
The larva is 35-40 mm in length, its pale-green to dark-brown color depends on the host plant, and with a dark, dorsal stripe.
The head, pronotum, the thoracic legs and the five pairs of abdominal prolegs are dark brown or black.
Host plants: Highly polyphagous, feeding on about 200 plant species, mainly annuals, developing on a wide range of food,
fiber, oil and fodder crops as well as on many wild plants and perennial horticultural crops.

Life cycle: Like other Noctuidae, the adults emerge after sunset, being active at night. The females mate several times and lay
300-3000 eggs during their lifetime, which lasts up to 3 weeks in summer. The eggs are laid singly on stems, leaves, and fruit.
The hatching larvae feed and reach a length of 40 mm, then leaving the plants to enter the soil, where they pupate. In
summer the pest develops in 25-40 days, in the cold season within 6-7 months. In the Middle East the pest completes 4-5
annual generations, some of which overlap. Most bollworm populations in the Middle East undergo a pupal diapause during
winter, emerging in early spring. This pest may migrate over long distances, borne by winds.
Economic importance: The damage caused by H. armigera is annually estimated, world-wide, to exceed US$2 billion, and
the bollworm is listed as an A2 quarantine pest by EPPO. In the Middle East it is a major pest of cotton, tomatoes and other
solanaceous crops, legumes like peas and beans, and maize (corn). Attacking cotton, the larvae initially feed on leaves, later
boring into flowers and bolls. As bolls are infested while too small to sustain the larvae, they move from boll to boll; the
affected bolls fail to develop and the quality of the lint is severely spoilt. Slightly damaged bolls are also damaged due to
being invaded by rot microorganisms. In tomatoes, the larvae bore into immature, ripening and ripe fruit, preferring the latter,
and cause rot. In maize, larvae bore into stems and can cause serious plant lodging. Infesting the developing cobs, larvae
penetrate mainly through the “silk” and feed on the seeds. In all cases, the economic value of the crops, for commercial or for
industrial use, is much reduced.
Prevention and Control
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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the
product's label.
Introduction

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.

IPM Programmes

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.

Regulatory Control

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.

Host-Plant Resistance

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.

Biological Control

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).

Chemical Control

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.

Modelling

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).