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Datasheet

Helicoverpa zea
(American cotton bollworm)

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Datasheet

Helicoverpa zea (American cotton bollworm)

Summary

  • Last modified
  • 01 April 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Helicoverpa zea
  • Preferred Common Name
  • American cotton bollworm
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Helicoverpa zea (American cotton bollworm); a normal, 12-day-old cotton bollworm larva raised on a control diet. USA.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); a normal, 12-day-old cotton bollworm larva raised on a control diet. USA.
Copyright©Peggy Greb/USDA Agricultural Research Service/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); a normal, 12-day-old cotton bollworm larva raised on a control diet. USA.
LarvaHelicoverpa zea (American cotton bollworm); a normal, 12-day-old cotton bollworm larva raised on a control diet. USA.©Peggy Greb/USDA Agricultural Research Service/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larval damage under bloom. USA.
TitleLarval damage
CaptionHelicoverpa zea (American cotton bollworm); larval damage under bloom. USA.
Copyright©Ronald Smith/Auburn University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larval damage under bloom. USA.
Larval damageHelicoverpa zea (American cotton bollworm); larval damage under bloom. USA.©Ronald Smith/Auburn University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva and larval damage to tomato fruit. USA. June 2017.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); larva and larval damage to tomato fruit. USA. June 2017.
Copyright©C.Watts/via flickr - CC BY 2.0
Helicoverpa zea (American cotton bollworm); larva and larval damage to tomato fruit. USA. June 2017.
LarvaHelicoverpa zea (American cotton bollworm); larva and larval damage to tomato fruit. USA. June 2017.©C.Watts/via flickr - CC BY 2.0
Helicoverpa zea (American cotton bollworm); late instar larva on cotton boll (Gossypium hirsutum). USA.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); late instar larva on cotton boll (Gossypium hirsutum). USA.
Copyright©Ronald Smith/Auburn University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); late instar larva on cotton boll (Gossypium hirsutum). USA.
LarvaHelicoverpa zea (American cotton bollworm); late instar larva on cotton boll (Gossypium hirsutum). USA.©Ronald Smith/Auburn University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); full grown larva on cotton boll. USA.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); full grown larva on cotton boll. USA.
Copyright©Scott Bauer/USDA Agricultural Research Service/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); full grown larva on cotton boll. USA.
LarvaHelicoverpa zea (American cotton bollworm); full grown larva on cotton boll. USA.©Scott Bauer/USDA Agricultural Research Service/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva in the field, on cotton (Gossypium hirsutum). USA.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); larva in the field, on cotton (Gossypium hirsutum). USA.
Copyright©Russ Ottens/University of Georgia/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva in the field, on cotton (Gossypium hirsutum). USA.
LarvaHelicoverpa zea (American cotton bollworm); larva in the field, on cotton (Gossypium hirsutum). USA.©Russ Ottens/University of Georgia/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larval damage to corn ear (Zea mays). USA.
TitleLarval damage
CaptionHelicoverpa zea (American cotton bollworm); larval damage to corn ear (Zea mays). USA.
Copyright©Whitney Cranshaw/Colorado State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larval damage to corn ear (Zea mays). USA.
Larval damageHelicoverpa zea (American cotton bollworm); larval damage to corn ear (Zea mays). USA.©Whitney Cranshaw/Colorado State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva and larval damage to corn ear (Zea mays). USA. April 2017.
TitleLarval damage
CaptionHelicoverpa zea (American cotton bollworm); larva and larval damage to corn ear (Zea mays). USA. April 2017.
Copyright©Scot Nelson/via flickr - CC BY 2.0
Helicoverpa zea (American cotton bollworm); larva and larval damage to corn ear (Zea mays). USA. April 2017.
Larval damageHelicoverpa zea (American cotton bollworm); larva and larval damage to corn ear (Zea mays). USA. April 2017.©Scot Nelson/via flickr - CC BY 2.0
Helicoverpa zea (American cotton bollworm); larva and larval damage to a corn ear (Zea mays). USA.
TitleLarval damage
CaptionHelicoverpa zea (American cotton bollworm); larva and larval damage to a corn ear (Zea mays). USA.
Copyright©Eric R. Day/Virginia Polytechnic Institute & State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva and larval damage to a corn ear (Zea mays). USA.
Larval damageHelicoverpa zea (American cotton bollworm); larva and larval damage to a corn ear (Zea mays). USA.©Eric R. Day/Virginia Polytechnic Institute & State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva in the field, feeding on a panicle of pearl millet (Pennisetum glaucum). USA.
TitleLarva
CaptionHelicoverpa zea (American cotton bollworm); larva in the field, feeding on a panicle of pearl millet (Pennisetum glaucum). USA.
Copyright©Russ Ottens/University of Georgia/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); larva in the field, feeding on a panicle of pearl millet (Pennisetum glaucum). USA.
LarvaHelicoverpa zea (American cotton bollworm); larva in the field, feeding on a panicle of pearl millet (Pennisetum glaucum). USA.©Russ Ottens/University of Georgia/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); pupa, in field, on soyabean (Glycine max). USA. September 2011.
TitlePupa
CaptionHelicoverpa zea (American cotton bollworm); pupa, in field, on soyabean (Glycine max). USA. September 2011.
Copyright©Adam Sisson/Iowa State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); pupa, in field, on soyabean (Glycine max). USA. September 2011.
PupaHelicoverpa zea (American cotton bollworm); pupa, in field, on soyabean (Glycine max). USA. September 2011.©Adam Sisson/Iowa State University/Bugwood.org - CC BY 3.0 US
Helicoverpa zea (American cotton bollworm); adult. Cuivre River State Park, Missouri USA. September 2014.
TitleAdult
CaptionHelicoverpa zea (American cotton bollworm); adult. Cuivre River State Park, Missouri USA. September 2014.
Copyright©Andy Reago & Chrissy McClarren/via wikipedia - CC BY 2.0
Helicoverpa zea (American cotton bollworm); adult. Cuivre River State Park, Missouri USA. September 2014.
AdultHelicoverpa zea (American cotton bollworm); adult. Cuivre River State Park, Missouri USA. September 2014.©Andy Reago & Chrissy McClarren/via wikipedia - CC BY 2.0
Helicoverpa zea (American cotton bollworm); adult.
TitleAdult
CaptionHelicoverpa zea (American cotton bollworm); adult.
CopyrightPublic Domain - Released by the USGS Bee Inventory & Monitoring Lab.
Helicoverpa zea (American cotton bollworm); adult.
AdultHelicoverpa zea (American cotton bollworm); adult.Public Domain - Released by the USGS Bee Inventory & Monitoring Lab.
Helicoverpa armigera (cotton bollworm); a parasitic wasp attacks a caterpillar, laying a single egg inside and marking the caterpillar with her own chemical scent.
TitleNatural enemy
CaptionHelicoverpa armigera (cotton bollworm); a parasitic wasp attacks a caterpillar, laying a single egg inside and marking the caterpillar with her own chemical scent.
Copyright©USDA-ARS
Helicoverpa armigera (cotton bollworm); a parasitic wasp attacks a caterpillar, laying a single egg inside and marking the caterpillar with her own chemical scent.
Natural enemyHelicoverpa armigera (cotton bollworm); a parasitic wasp attacks a caterpillar, laying a single egg inside and marking the caterpillar with her own chemical scent.©USDA-ARS

Identity

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Preferred Scientific Name

  • Helicoverpa zea (Boddie, 1850)

Preferred Common Name

  • American cotton bollworm

Other Scientific Names

  • Bombyx obsoleta Fabricius
  • Chloridea obsoleta Fabricius
  • Heliothis armigera auct.nec Huebner Hübner
  • Heliothis ochracea Cockerell
  • Heliothis umbrosa Grote
  • Heliothis zea Boddie
  • Phalaena zea (Boddie)

International Common Names

  • English: bollworm, American; corn earworm; cotton bollworm; tomato fruitworm
  • Spanish: bellotero; elotero; gusano bellotero del algodon; gusano de la bellota del algodón; gusano de la mazorca; gusano de la mazorca del maiz; gusano de las cápsulas; gusano del elote del maíz; gusano del fruto del tomate; gusano elotero; noctua del tomate; oruga de la mazorca
  • French: chenille des epis du mais; noctuelle de la tomate; noctuelle des tomates; ver de la capsule; ver de l'épi du maïs

Local Common Names

  • Argentina: isoca del maiz
  • Brazil: lagarta da espiga do milho; lagarta das espicas
  • Denmark: amerikansk bomuldsugle
  • Germany: Amerikanischer Baumwollkapselwurm; Wurm, Amerikanischer Baumwollkapsel-
  • Italy: elotide del cotone; elotide del granturco; elotide del pomodoro; elotide del tomato; nottua del granturco; nottua gialla del granturco
  • Netherlands: Mimosa-rups
  • Turkey: yesil kurt

EPPO code

  • HELIZE (Helicoverpa zea)

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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The taxonomic situation regarding H. zea is complicated and presents several problems. Hardwick (1965) reviewed the New World corn earworm species complex and the Old World African bollworms, 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. Specifically, 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; 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 (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 since dissection of genitalia is required for identification, there has been resistance to the name change (for example, Heath and Emmet, 1983), but Hardwick's work is generally accepted and so the name change must also be accepted (Matthews, 1991).

Description

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Egg

Eggs are subspherical, radially ribbed, 0.52 mm high and 0.59 mm in diameter, stuck singly to the plant substrate, green when laid, turning red and finally grey before hatching. Egg maturity takes 2-3 days at 20-30°C. Eggs are usually located on the silk of maize, or on fruiting structures.

Larva

On hatching, the tiny grey caterpillars have a black head; they grow through six instars usually, but five and seven instars are not uncommon, and the final body size is approximately 40 mm long. In the third instar, two colour phases can develop: brown (the predominant phase) and green (less frequent). Longitudinal lines of white, cream or yellow are present, and the spiracular band is the most distinct. As the larvae develop, the pattern becomes better defined, but in the final instar (sixth) the coloration changes abruptly into a bright pattern, often pinkish, and with extra striations. Larvae have 5 pairs of prolegs.

Pupa

Pupae are light to dark brown depending on maturity and approximately 20 mm long, with two distinct terminal cremaster spines. Pupae are located 5-15 cm below the soil surface in earthen cells.

Adult

A stout-bodied (20-25 mm long) brown moth of wing-span 38-43 mm; forewing pale brown (female) to greenish (male) with darker transverse markings, underwings pale with a broad dark marginal band. 

Distribution

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H. zea is confined to the New World. It occurs throughout the Americas from Canada to Argentina (IIE, 1993).

Distribution Table

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

Last updated: 17 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Europe

NetherlandsAbsent, Confirmed absent by survey
RussiaAbsent, Confirmed absent by survey
SloveniaAbsent
SwitzerlandAbsent, Intercepted only
United KingdomAbsent, Intercepted onlyOriginal citation: Seymour and (1978)

North America

Antigua and BarbudaPresent
BahamasPresent
BarbadosPresent
BermudaPresent
CanadaPresent, Localized
-British ColumbiaPresent
-ManitobaPresent
-New BrunswickPresent
-Nova ScotiaPresent
-OntarioPresent
-QuebecPresent
-SaskatchewanPresent
Costa RicaPresent
CubaPresent
DominicaPresent
Dominican RepublicPresent
El SalvadorPresent
GuadeloupePresent
GuatemalaPresent
HaitiPresent
HondurasPresent
JamaicaPresent
MartiniquePresent, Widespread
MexicoPresent, Widespread
MontserratPresent
NicaraguaPresent
PanamaPresent
Puerto RicoPresent
Saint Kitts and NevisPresent, Localized
Saint LuciaPresent
Saint Vincent and the GrenadinesPresent
Trinidad and TobagoPresent, Widespread
U.S. Virgin IslandsPresent
United StatesPresent, Widespread
-AlabamaPresent
-ArizonaPresent
-ArkansasPresent
-CaliforniaPresent
-ColoradoPresent
-ConnecticutPresent
-DelawarePresent
-FloridaPresent
-GeorgiaPresent
-HawaiiPresent
-IdahoPresent
-IllinoisPresent
-IndianaPresent
-IowaPresent
-KansasPresent
-KentuckyPresent
-LouisianaPresent
-MainePresent
-MarylandPresent
-MassachusettsPresent
-MichiganPresent
-MinnesotaPresent
-MississippiPresent
-MissouriPresent
-MontanaPresent
-NebraskaPresent
-NevadaPresent
-New HampshirePresent
-New JerseyPresent
-New MexicoPresent
-New YorkPresent
-North CarolinaPresent
-North DakotaPresent
-OhioPresent
-OklahomaPresent
-OregonPresent
-PennsylvaniaPresent
-Rhode IslandPresent
-South CarolinaPresent
-South DakotaPresent
-TennesseePresent
-TexasPresent
-UtahPresent
-VermontPresent
-VirginiaPresent
-WashingtonPresent
-West VirginiaPresent
-WisconsinPresent
-WyomingPresent

South America

ArgentinaPresent
BoliviaPresent
BrazilPresent, Widespread
-BahiaPresent
-CearaPresent
-Distrito FederalPresent
-GoiasPresent
-Mato GrossoPresent
-Mato Grosso do SulPresent
-Minas GeraisPresent
-ParaPresent
-ParanaPresent
-PernambucoPresent
-Rio de JaneiroPresent
-Rio Grande do SulPresent
-RoraimaPresent
-Santa CatarinaPresent
-Sao PauloPresent
ChilePresent, Widespread
-Easter IslandPresent
ColombiaPresent
EcuadorPresent, Widespread
Falkland IslandsPresent, Few occurrences
French GuianaPresent
GuyanaPresent
ParaguayPresent, Widespread
PeruPresent
SurinamePresent
UruguayPresent, Widespread
VenezuelaPresent

Risk of Introduction

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H. zea was recently added to the EPPO A1 list of quarantine pests, and is also considered as a quarantine pest by APPPC. Originally, H. zea was considered as practically synonymous with H. armigera, an A2 quarantine pest (EPPO/CABI, 1996). The addition to the EPPO list harmonizes it with EU Directive Annex I/A1.

Phytosanitary Measures

For the related H. armigera, EPPO (EPPO, 1990) makes recommendations on phytosanitary measures which would also be suitable for H. zea. According to these, 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.

Bibliographies are included in the monograph by Hardwick (1965) (2000 titles on H. zea), and the reviews by Fitt (1989) (194 titles), and King and Coleman (1989) (159 references). Most of the basic research on H. zea was done in the early 1900s and published under early synonyms. Many references to H. zea are made in publications relating to the cultivation/protection of specific crops, for example, Chiang (1978), COPR (1983), and Pitre (1985).

Hosts/Species Affected

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H. zea is polyphagous in feeding habits but it shows a definite preference in North America for young maize cobs and tassels, and particularly for the cultivars grown as sweetcorn and popcorn, and also for Sorghum. Most hosts are recorded from the Poaceae, Malvaceae, Fabaceae and Solanaceae; in total more than 100 plant species are recorded as hosts. A feeding preference is shown for flowers and fruits of host plants.

For further information see Barber (1937), Neunzig (1963), Davidson and Peairs (1966) and Matthews (1991).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Abelmoschus esculentus (okra)MalvaceaeMain
    Abutilon theophrasti (velvet leaf)MalvaceaeOther
      Amaranthus (amaranth)AmaranthaceaeOther
        Arachis hypogaea (groundnut)FabaceaeOther
          Brassica oleracea (cabbages, cauliflowers)BrassicaceaeOther
            Brassica oleracea var. botrytis (cauliflower)BrassicaceaeOther
              Brassica oleracea var. capitata (cabbage)BrassicaceaeOther
                Cajanus cajan (pigeon pea)FabaceaeMain
                  Capsicum (peppers)SolanaceaeOther
                    Capsicum annuum (bell pepper)SolanaceaeMain
                      Chenopodium quinoa (quinoa)ChenopodiaceaeOther
                        Cicer arietinum (chickpea)FabaceaeOther
                          CitrusRutaceaeOther
                            Cucumis melo (melon)CucurbitaceaeOther
                              Cucumis sativus (cucumber)CucurbitaceaeOther
                                Fragaria (strawberry)RosaceaeOther
                                  Fragaria ananassa (strawberry)RosaceaeOther
                                    Geranium carolinianum (Carolina geranium)GeraniaceaeOther
                                      Gerbera (Barbeton daisy)AsteraceaeOther
                                        Glycine max (soyabean)FabaceaeMain
                                          Gossypium (cotton)MalvaceaeMain
                                            Helianthus annuus (sunflower)AsteraceaeMain
                                              Ipomoea purpurea (tall morning glory)ConvolvulaceaeOther
                                                Lactuca sativa (lettuce)AsteraceaeOther
                                                  Lamium amplexicaule (henbit deadnettle)LamiaceaeOther
                                                    Lespedeza juncea var. sericea (Sericea lespedeza)FabaceaeOther
                                                      Lonicera japonica (Japanese honeysuckle)CaprifoliaceaeWild host
                                                        Medicago lupulina (black medick)FabaceaeOther
                                                          Medicago sativa (lucerne)FabaceaeOther
                                                            Nicotiana tabacum (tobacco)SolanaceaeOther
                                                              Panicum miliaceum (millet)PoaceaeOther
                                                                Phaseolus (beans)FabaceaeMain
                                                                  Phaseolus vulgaris (common bean)FabaceaeMain
                                                                    Salix (willows)SalicaceaeOther
                                                                      Securigera varia (crown vetch)FabaceaeOther
                                                                        Solanum lycopersicum (tomato)SolanaceaeMain
                                                                          Solanum melongena (aubergine)SolanaceaeMain
                                                                            Sorghum bicolor (sorghum)PoaceaeMain
                                                                              Spinacia oleracea (spinach)ChenopodiaceaeOther
                                                                                Trifolium (clovers)FabaceaeOther
                                                                                  Trifolium incarnatum (Crimson clover)FabaceaeOther
                                                                                    Vicia sativa (common vetch)FabaceaeOther
                                                                                      Vicia villosa (hairy vetch)FabaceaeWild host
                                                                                        Vigna unguiculata (cowpea)FabaceaeOther
                                                                                          Zea mays (maize)PoaceaeMain
                                                                                            Zea mays subsp. mays (sweetcorn)PoaceaeMain

                                                                                              Growth Stages

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                                                                                              Flowering stage, Fruiting stage, Vegetative growing stage

                                                                                              Symptoms

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                                                                                              Fruiting structures are consumed or damaged and feeding damage also facilitates entry by diseases and other insect pests. In cotton the square (flower bud), flowers and young bolls are attacked and larvae excavate the interior. Young shoots and leaves can also be damaged, especially in the absence of fruiting structures.

                                                                                              Young maize plants have serial holes in the leaves following whorl feeding on the apical leaf. On larger plants the silks are grazed and eggs can be found stuck to the silks. As the ears develop, the soft milky grains in the top few centimeters of the cobs are eaten; usually only one large larva per cob can be seen. Ear damage is often localized to the tip but can increase the incidence of disease.

                                                                                              Sorghum heads are grazed. Legume pods are holed and the seeds eaten. Bore holes can be seen in tomato fruits, cotton bolls, cabbage and lettuce hearts, and flower heads.

                                                                                              List of Symptoms/Signs

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                                                                                              SignLife StagesType
                                                                                              Fruit / external feeding
                                                                                              Fruit / internal feeding
                                                                                              Growing point / external feeding
                                                                                              Growing point / internal feeding; boring
                                                                                              Inflorescence / external feeding
                                                                                              Inflorescence / internal feeding
                                                                                              Leaves / external feeding
                                                                                              Seeds / external feeding
                                                                                              Seeds / internal feeding

                                                                                              Biology and Ecology

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                                                                                              Eggs are laid mostly on the silks of maize plants in small numbers (one to three), stuck to the plant tissues. Choice of oviposition site by the female seems to be governed by a combination of physical and chemical cues. Female fecundity can be dependent upon the quality and quantity of larval food, and also on the quality of adult nutrition. Up to 3000 eggs have been laid by a single female in captivity, but 1000-1500 per female is more usual in the wild. Hatching occurs after 2-4 days and the eggs change colour from green through red to grey. The tiny grey larvae first eat the eggshell and after a short rest they wander actively for a while before starting to feed on the plant. They usually feed on the silks initially and then on the young tender kernels after entering the tip of the husk. By the third instar the larvae become cannibalistic and usually only one larva survives per cob. Feeding damage is typically confined to the tip of the cob. Larval development usually takes 14-25 (mean 16) days, but under cooler conditions 60 days are required. In the final instar (usually sixth) feeding ceases and the fully fed caterpillar leaves the cob and descends to the ground. It then burrows into the soil for some 10-12 cm and forms an earthen cell, where it rests in a prepupal state for a day or two, before finally pupating. Two basic types of pupal diapause are recognized, one in relation to cold and the other in response to arid conditions. In the tropics pupation takes 10-14 (mean 13) days; the male takes 1 day longer than the female. Diapausing pupae are viable as far north as 40-45°N in the USA.

                                                                                              Adults are nocturnal in habit and emerge in the evenings. Maize fields in the USA regularly produce 40,000 to 50,000 adult moths per hectare. Flying adults respond to light radiation at night and are attracted to light traps (Hardwick, 1968), especially the ultraviolet type, in company with many other local noctuids. Sex aggregation pheromones have been identified and synthesized for most of the Heliothis/Helicoverpa pest species, and pheromone traps can be used for population monitoring. Adult longevity is recorded as being about 17 days in captivity; they drink water and feed on nectar from both floral and extra-floral nectaries. The moths fly strongly and are regular seasonal migrants, flying hundreds of kilometres from the USA into Canada. They migrate by flying high with prevailing wind currents.

                                                                                              The life cycle can be completed in 28-30 days at 25°C and in the tropics there may be up to 10-11 generations per year. All stages of the insect are to be found throughout the year if food is available, but development is slowed or stopped by either drought or cold. In the northern USA there are only two generations per year, in Canada only one generation.

                                                                                              For more information, see Hardwick (1965), Beirne (1971), Balachowsky (1972), Allemann (1979), King and Saunders (1984) and Fitt (1989).

                                                                                              Means of Movement/Dispersal

                                                                                              H. zea is a facultative seasonal nocturnal migrant, and adults migrate in response to poor local conditions for reproduction, when weather conditions are suitable. Three types of movement are practiced by Helicoverpa moths: short-range, long-range, and migration. Short-range dispersal is usually within the crop and low over the foliage, and largely independent of wind currents. Long-range flights are higher (up to 10 m), further (1-10 km), and usually downwind, from crop to crop. Migratory flights occur at higher altitudes (up to 1-2 km) and may last for several hours. The moths can be carried downwind hundreds of kilometres; 400 km is not uncommon for such a flight. There is now evidence that many of them originate in Mexico as young adults and migrate northwards into the USA in the early spring. Probably three generations are required to effect the annual displacement from Mexico up to southern Ontario. Transatlantic dispersal is clearly a possibility for this moth, although it has not yet been demonstrated.

                                                                                              Air-freight transportation of agricultural produce from the New World to Europe is an ever increasing commercial enterprise, especially with vegetables and ornamentals. Almost every year, caterpillars of H. zea are intercepted on this produce in the UK (Seymour, 1978).

                                                                                              Natural enemies

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                                                                                              Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                                                              Agelaius phoeniceus Predator
                                                                                              Archytas marmoratus Parasite Larvae/Pupae
                                                                                              Aspergillus niger Antagonist
                                                                                              Bacillus circulans Pathogen Larvae
                                                                                              Bacillus subtilis Pathogen Larvae
                                                                                              Bacillus thuringiensis Pathogen Larvae
                                                                                              Bacillus thuringiensis alesti Pathogen Larvae
                                                                                              Bacillus thuringiensis israelensis Pathogen Larvae
                                                                                              Bacillus thuringiensis kurstaki Pathogen Larvae
                                                                                              Bacillus thuringiensis thuringiensis Pathogen Larvae
                                                                                              baculo-like virus Pathogen
                                                                                              Baculovirus heliothis Pathogen
                                                                                              Beauveria bassiana Pathogen
                                                                                              Brachymeria ovata Parasite Pupae
                                                                                              Calleida decora Predator Larvae
                                                                                              Calosoma sayi Predator Larvae
                                                                                              Campoletis flavicincta Parasite
                                                                                              Campoletis grioti Parasite
                                                                                              Campoletis sonorensis Parasite
                                                                                              Cardiochiles nigriceps Parasite Larvae
                                                                                              Cardiochiles seminiger Parasite Larvae
                                                                                              Chelonus curvimaculatus Parasite Larvae
                                                                                              Chelonus insularis Parasite Larvae
                                                                                              Chelonus narayani Parasite Larvae Hawaii
                                                                                              Chrysoperla carnea Predator
                                                                                              Chrysoperla rufilabris Predator
                                                                                              Coleomegilla maculata Predator
                                                                                              Collops quadrimaculatus Predator
                                                                                              Cotesia congregata Parasite Larvae
                                                                                              Cotesia kazak Parasite Larvae
                                                                                              Cotesia marginiventris Parasite Larvae
                                                                                              Cytoplasmic polyhedrosis virus (CPV) Pathogen Larvae
                                                                                              cytoplasmic polyhedrosis viruses Pathogen Larvae
                                                                                              Diapetimorpha introita Parasite
                                                                                              Encarsia porteri Parasite Eggs
                                                                                              Entomophaga aulicae Pathogen
                                                                                              Erythemis simplicicollis Predator
                                                                                              Eucelatoria bryani Parasite Larvae Hawaii
                                                                                              Eucelatoria rubentis Parasite
                                                                                              Euphorocera edwardsii Parasite Larvae
                                                                                              Euplectrus comstockii Parasite
                                                                                              Euplectrus platyhypenae Parasite
                                                                                              Geocoris punctipes Predator Eggs
                                                                                              Geocoris uliginosus Predator Eggs
                                                                                              Glabromicroplitis croceipes Parasite Larvae
                                                                                              Helicoverpa armigera nuclear polyhedrosis virus Pathogen Adults/Larvae
                                                                                              Heliothis nucleopolyhedrosis virus Pathogen
                                                                                              Hippodamia convergens Predator
                                                                                              Hyposoter exiguae Parasite
                                                                                              Ichneumon promissorius Parasite
                                                                                              Iridovirus Pathogen Larvae
                                                                                              Lebia analis Predator Larvae
                                                                                              Lespesia aletiae Parasite Larvae
                                                                                              Lespesia archippivora Parasite Larvae
                                                                                              Leuconostoc mesenteroides Pathogen
                                                                                              Metagonistylum minense Parasite Larvae
                                                                                              Metaphidippus galathea Predator
                                                                                              Metarhizium anisopliae Pathogen
                                                                                              Meteorus autographae Parasite Larvae
                                                                                              Microchelonus blackburni Parasite Larvae
                                                                                              Microplitis demolitor Parasite Larvae
                                                                                              Microplitis rufiventris Parasite Larvae
                                                                                              Nabis alternatus Predator
                                                                                              Nabis roseipennis Predator
                                                                                              Nemorilla pyste Parasite Larvae
                                                                                              Nomuraea rileyi Pathogen Larvae USA; South Carolina soyabeans
                                                                                              Notoxus monodon Predator
                                                                                              Nucleopolyhedrosis virus Pathogen Larvae
                                                                                              Orius insidiosus Predator Eggs Hawaii
                                                                                              Orius tristicolor Predator
                                                                                              Oxyopes salticus Predator
                                                                                              Paecilomyces tenuipes Pathogen
                                                                                              Palexorista laxa Parasite Larvae
                                                                                              Paratriphleps laeviusculus Predator USA cotton; maize; tomatoes
                                                                                              Peucetia viridana Predator
                                                                                              Phidippus audax Predator
                                                                                              Philonthus alumnus Predator
                                                                                              Phyllobaenus pubescens Predator
                                                                                              Podisus connexivus Predator
                                                                                              Podisus maculiventris Predator
                                                                                              Podisus placidus Predator
                                                                                              Polistes metricus Predator
                                                                                              Pristomerus spinator Parasite
                                                                                              Pseudatomoscelis seriatus Predator
                                                                                              Rogas perplexus Parasite Larvae
                                                                                              Solenopsis invicta Predator
                                                                                              Spanagonicus albofasciatus Predator
                                                                                              Staphylococcus epidermidis Pathogen
                                                                                              Steinernema carpocapsae Parasite
                                                                                              Steinernema feltiae Parasite
                                                                                              Steinernema riobravis Parasite
                                                                                              Telenomus remus Parasite
                                                                                              Telenomus spodopterae Parasite Eggs
                                                                                              Trichogramma atopovirilia Parasite Eggs
                                                                                              Trichogramma brevicapillum Parasite Eggs
                                                                                              Trichogramma chilonis Parasite Eggs
                                                                                              Trichogramma deion Parasite Eggs
                                                                                              Trichogramma evanescens Parasite Eggs
                                                                                              Trichogramma exiguum Parasite Eggs
                                                                                              Trichogramma maltbyi Parasite Eggs
                                                                                              Trichogramma minutum Parasite Eggs
                                                                                              Trichogramma nagarkattii Parasite Eggs
                                                                                              Trichogramma parkeri Parasite Eggs
                                                                                              Trichogramma perkinsi Parasite Eggs
                                                                                              Trichogramma pretiosum Parasite Eggs California; Nicaragua; Nova Scotia; Texas; USA; Texas cotton
                                                                                              Trichogramma thalense Parasite Eggs
                                                                                              Vairimorpha necatrix Pathogen
                                                                                              Vespula pensylvanica Predator
                                                                                              virus-like particles
                                                                                              Winthemia rufiventris Parasite Larvae
                                                                                              Xanthomonas maltophilia Pathogen
                                                                                              Zelus tetracanthus Predator

                                                                                              Notes on Natural Enemies

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                                                                                              Kogan et al. (1989) provides a full list of natural enemy records.

                                                                                              Impact

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                                                                                              In North America it is reported that H. zea is the second most important economic pest species (preceded by codling moth, Cydia pomonella) (Hardwick, 1965). Fitt (1989) quotes the estimated annual cost of damage by H. zea and H. virescens together on all crops in the USA as more than US$ 1000 million, despite the expenditure of US$ 250 million on insecticide application.

                                                                                              Reasons for the success and importance of this agricultural pest include its high fecundity, polyphagous larval feeding habits, high mobility of both larvae locally and adults with their facultative seasonal migration, and a facultative pupal diapause.

                                                                                              Damage is usually serious and costly because of the larval feeding preference for the reproductive structures and growing points rich in nitrogen (for example, maize cobs and tassels, sorghum heads, cotton bolls and buds, etc), and they have a direct influence on yield. Many of the crops attacked are of high value (cotton, maize, tomatoes). If this pest should become established in protected cultivation economic damage could be widespread.

                                                                                              Infestations of maize grown for silage or for grain are not of direct economic importance; losses are typically about 5% and no control measures are taken, but they serve as a focus, or reservoir of infestation. In many areas the first generation is not regarded as a pest (often on Trifolium) and it does not become an economic pest on cultivated crops until the second, third or even fourth generation.

                                                                                              Detection and Inspection

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                                                                                              Feeding damage is usually visible and the larvae can be seen on the surface of plants but often they are hidden within plant organs (flowers, fruits, etc). Bore holes may be visible, but otherwise it is necessary to cut open the plant organs to detect the pest. Because of morphological similarity, it is impossible to distinguish the larvae of H. zea from those of Heliothis armigera, already present in the EPPO region. Positive identification is achieved by rearing the larvae and examining the genitalia of the adult.

                                                                                              Similarities to Other Species/Conditions

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                                                                                              The adults are similar in appearance to Heliothis armigera, but differ in several details in their genitalia (Hardwick, 1965); dissection and slide-mounting are required for specific determination, and some aspects are comparative so that a series of closely related species have to be available for comparison.

                                                                                              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

                                                                                              Control of H. zea has been advocated in the USA since the middle of the nineteenth century, and measures fall into two broad categories: those aimed at an overall pest population reduction, and others aimed at the protection of a particular crop. In most situations it is now recommended that integrated pest management be used (Bottrell, 1979).

                                                                                              Cultural Control

                                                                                              Various cultural practices can be used to kill the different instars, including deep ploughing, discing and other methods of mechanical destruction, manipulation of sowing dates and use of trap crops.

                                                                                              Biological Control

                                                                                              In many areas, natural control of this pest may be quite effective for most of the time. Insect parasitoids attack the eggs (especially Trichogramma spp.) and larvae, and some predators can be important in reducing pest populations. King and Coleman (1989) discuss the prospects for long-term biological control of Heliothis/Helicoverpa spp., and clearly this should be an important component of any regional IPM programme.

                                                                                              The most frequently tried method of achieving biological control has been by augmentative releases of artificially reared parasites or predators, especially using Trichogramma spp. However, releases in cotton have not been consistently effective against heliothine populations. Microplitis croceipes could be more effective because it is less affected by organophosphate pesticides and synthetic pyrethroids.

                                                                                              There has also been interest in exploiting entomophagous pathogens such as Bacillus thuringiensis and Heliothis NPV. In cotton, maize and tomato, transgenic crop varieties expressing the active Bacillus thuringiensis toxin have been used commercially.

                                                                                              A commercial formulation of the nuclear polyhedrosis virus Baculovirus heliothis gave control that was equal to chemical methods. However, the cost of virus applications was higher than chemical control methods. (Martinez and Swezey, 1988).

                                                                                              Host-Plant Resistance

                                                                                              The development of crop cultivars resistant or tolerant to damage by Heliothis and Helicoverpa spp. has major potential in their management, particularly for communities with few resources. Many crops possess some genetic potential that can be exploited by breeders to produce varieties less subject to pest damage. Resistance can take three basic forms: antixenosis, antibiosis and tolerance. Varieties of crop hosts showing resistance to Heliothis or Helicoverpa have been identified or developed in cotton, chickpeas, soyabean, tomato, maize, sorghum, millet and tobacco.

                                                                                              In maize, resistant genotypes have been identified which have a high concentration of maysin (rhamnosyl-6-C-(4-ketofucosyl)-5,7,3',4'-tetrahydroxyflavone), a C-glycosyl flavone, in silk tissue. Quantitative trait loci for maysin production were identified on chromosomes 1 (p1) and 9 (umc105a) (Byrne et al., 1996).

                                                                                              In cotton, gossypol glands on the calyx crowns of flower buds confers considerable resistance to H. zea (Calhoun et al., 1997).

                                                                                              Transgenic maize containing genes encoding delta-endotoxins from Bacillus thuringiensis (Bt) kurstaki have been commercialized in the USA. Feeding studies using Cry1A(c) toxins demonstrated transformed cotton plants are highly toxic to first-fourth instars of H. zea, but not to fifth instar larvae. Movement of fifth instar larvae from non-Bt plants to Bt-cotton plants in mixed stands could result in feeding and injury to Bt plants (Halcomb et al., 1996). A new type of toxin called vegetative insecticidal proteins (vip) has been isolated from Bacillus thuringiensis during the vegetative phase of growth which shows a wide spectrum of activities against lepidopteran insects, especially noctuids such as H. zea (Estruch et al., 1996).

                                                                                              There is little knowledge of the interactions between natural enemies of Heliothis or Helicoverpa and host-plant resistance, but it cannot be assumed that resistance will always be compatible with natural control. For example, laboratory tests using resistant tomato plants containing an alkaloid (alpha-tomatine) were found to be toxic to Hyposoter exiguae, a parasite of H. zea. The parasite acquired the alkaloid from its host after the host had ingested the alkaloid (Campbell and Duffey, 1979).

                                                                                              Chemical Control

                                                                                              Chemical control of the larvae has been the most widely used and generally successful method of pest destruction on most crops, but it is not easy because of larvae feeding within plant structures. The early history of chemical control of corn earworms is given by Hardwick (1965), while COPR (1983) includes a list of 29 insecticides effective for the control of Heliothis/Helicoverpa spp. at the recommended rates given. Pesticide resistance has been known for some years and is quite widespread (Fitt, 1989) especially on cotton crops.

                                                                                              For cotton, chemicals recommended for control include sulprofos, profenofos, thiodicarb, chlorpyrifos, acephate, amitraz and pyrethroids. Several Bacillus thuringiensis sprays are also recommended (Anon., 1997).

                                                                                              For maize, oil applied to silks reduces H. zea damage to the ear. Applications are generally made 3 days after silk emergence and applied on a weekly basis until silk dry down.

                                                                                              Sterile Backcrosses

                                                                                              Sterile male offspring are produced when certain species are crossed, for example, Heliothis subflexa and H. virescens. This fact has been exploited and evaluated on the island of St Croix, Virgin Islands, where after a 3-year release program, suppression was achieved.

                                                                                              Pheromonal Control

                                                                                              Mating of H. zea was reduced by 50% in a 12 ha maize field treated with hollow fibres containing (Z)-9-tetradecenyl formate (Mitchell and McLaughlin, 1982). Likewise, (Z)-11-hexadecenal, a component of the Heliothis virescens pheromone, reduced the mating of females of H. zea by 85% (Mitchell et al., 1976).

                                                                                              References

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                                                                                              Allemann DV, 1979. Maize pests in the USA. In: Hafliger E, ed. Maize. Basle, Switzerland: CIBA-GEIGY Ltd., 58-63.

                                                                                              Anon., 1997. Insect Control Guide. Ohio, USA: Meister Publishing Co., 442 pp.

                                                                                              Balachowsky AS, ed. , 1972. Entomology applied to agriculture. Tome II. Lepidoptera. Second volume. Zygaenoidea-Pyraloidea-Noctuoidea. Entomologie appliquee a l'agriculture. Tome II. Lepidopteres. Deuxieme volume. Zygaenoidea-Pyraloidea-Noctuoidea. Paris, France: Masson et Cie, pp 1059-1634.

                                                                                              Barber GW, 1937. Seasonal availability of food plants of two species of Heliothis in eastern Georgia. Journal of Economic Entomology, 30:150-158.

                                                                                              Beirne BP, 1971. Pest insects of annual crop plants in Canada. I. Lepidoptera. II. Diptera. III. Coleoptera. Memoirs of the Entomological Society of Canada, 78:124 pp.

                                                                                              Bottrell DG, 1979. Guidelines for integrated control of maize pests. FAO, Plant Production and Protection Paper No. 91. Rome, Italy: FAO.

                                                                                              Byrne PF, McMullen MD, Snook ME, Musket TA, Theuri JM, Widstrom NW, Wiseman BR, Coe EH, 1996. Quantitative trait loci and metabolic pathways: genetic control of the concentration of maysin, a corn earworm resistance factor, in maize silks. Proceedings of the National Academy of Sciences of the United States of America, 93(17):8820-8825; 38 ref.

                                                                                              CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp.

                                                                                              Calhoun DS, Jones JE, Dickson JI, Caldwell WD, Burris E, Leonard BR, Moore SH, Aguillard W, 1997. Registration of 'H1244' cotton. Crop Science, 37(3):1014-1015; 6 ref.

                                                                                              Campbell BC, Duffey SS, 1979. Tomatine and parasitic wasps: potential incompatibility of plant antibiosis with biological control. Science, 205(4407):700-702.

                                                                                              Chiang HC, 1978. Pest management in corn. Annual Review of Entomology, 23:101-123.

                                                                                              COPR, 1983. Pest Control in Tropical Tomatoes. London, UK: COPR.

                                                                                              Davidson RH, Peairs LM, 1966. Insect Pests of Farm, Garden and Orchard (6th edition). New York, USA: Wiley.

                                                                                              EPPO, 1990. Specific quarantine requirements. EPPO Technical Documents, No. 1008. Paris, France: European and Mediterranean Plant Protection Organization.

                                                                                              EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

                                                                                              Estruch JJ, Warren GW, Mullins MA, Nye GJ, Craig JA, Koziel MG, 1996. Vip3A, a novel Bacillus thuringiensis vegetative insecticidal protein with a wide spectrum of activities against lepidopteran insects. Proceedings of the National Academy of Sciences of the United States of America, 93(11):5389-5394; 23 refs.

                                                                                              Fitt GP, 1989. The ecology of Heliothis species in relation to agroecosystems. Annual Review of Entomology, 34:17-52.

                                                                                              Halcomb JL, Benedict JH, Cook B, Ring DR, 1996. Survival and growth of bollworm and tobacco budworm on nontransgenic and transgenic cotton expressing a CryIA insecticidal protein (Lepidoptera: Noctuidae). Environmental Entomology, 25(2):250-255; 29 ref.

                                                                                              Hardwick DF, 1965. The corn earworm complex. Memoirs of the Entomological Society of Canada, 40:1-247.

                                                                                              Hardwick DF, 1968. A brief review of the principles of light trap design with a description of an efficient trap for collecting noctuid moths. Journal of the Lepidopterists' Society, 22:65-75.

                                                                                              Hardwick DF, 1970. A generic revision of the North American Heliothidinae (Lepidoptera: Noctuidae). Memoirs of the Entomological Society of Canada, 73:1-59.

                                                                                              Heath J, Emmet AM, ed. , 1983. The moths and butterflies of Great Britain and Ireland. Volume 10. Noctuidae (Cuculliinp to Hypeninp) and Agaristidae. The moths and butterflies of Great Britain and Ireland. Volume 10. Noctuidae (Cuculliinp to Hypeninp) and Agaristidae. Harley Books Colchester UK, 459 pp.

                                                                                              IIE, 1993. Distribution Maps of Pests, Series A No. 373 (1st revision). Wallingford, UK: CAB International.

                                                                                              King ABS, Saunders JL, 1984. The invertebrate pests of annual food crops in Central America. A guide to their recognition and control. London, UK: Overseas Development Administration.

                                                                                              King FG, Coleman RJ, 1989. Potential for biological control of Heliothis species. Annual Review of Entomology, 34:53-75.

                                                                                              Kogan M, Helm CG, Kogan J, Brewer E, 1989. Distribution and economic importance of Heliothis virescens and Helicoverpa zea in North, Central, and South America and of their natural enemies and host plants. In: King EG, Jackson RD, eds. Proceedings of the workshop on the biological control of Heliothis: increasing the effectiveness of natural enemies. New Delhi, India: USDA, Far East Regional Office, 241-297.

                                                                                              Lu YongYue, Liang GuangWen, 2002. Spatial pattern of cotton bollworm (Helicoverpa zea) eggs with geostatistics. Journal of Huazhong Agricultural University, 21(1):13-17.

                                                                                              Marchiori CH, Oliveira AMS, Costa MCR, 2002. Insects collected in maize crop in Itumbiara, south of Goiás state, Brazil. (Insetos coletados em cultivar de milho em Itumbiara, sul de Goiás, Brasil.) Arquivos do Instituto Biológico (São Paulo), 69(Suplemento, Resumos expandidos):233-234. http://www.biologico.sp.gov.br/ARQUIVOS/V69_supl_RE/marchiori.PDF

                                                                                              Martinez R, Swezey SL, 1988. Control of Heliothis zea (Boddie) larvae with a nuclear polyhedrosis virus (Baculovirus heliothis) in cotton, Leon, Nicaragua 1983. Revista Nicaraguense de Entomologia, 2:13-18.

                                                                                              Matthews M, 1991. Classification of the Heliothinae. NRI Bulletin No. 44. Chatham, Kent: Natural Resources Institute.

                                                                                              Mitchell ER, Baumhover AH, Jacobson M, 1976. Reduction of mating potential of male Heliothis spp. and Spodoptera frugiperda in field plots treated with disruptants. Environmental Entomology, 5(3):484-486.

                                                                                              Mitchell ER, McLaughlin JR, 1982. Suppression of mating and oviposition by fall armyworm and mating by corn earworm in corn, using the air permeation technique. Journal of Economic Entomology, 75(2):270-274.

                                                                                              Neunzig HH, 1963. Wild host plants and parasites. Journal of Economic Entomology, 52:135-139.

                                                                                              Olivares TS, Angulo AO, Badilla Q R, 2011. Taxonomic notes and new register of moths for Easter Island (Lepidoptera: Noctuoidea). Entomological News, 122(2):157-164. http://www.bioone.org/loi/entn

                                                                                              Pitre HN, 1985. Insect problems on sorghum in the USA. Proceedings of the international sorghum entomology workshop, 15-21 July 1984, Texas A & M University, College Station, Texas, USA. Patancheru, Andhra Pradesh, India: International Crops Research Institute for the Semi-Arid Tropics, 73-81.

                                                                                              Seymour PR, ed. , 1978. Insects and other invertebrates intercepted in check inspections of imported plant material in England and Wales during 1976 and 1977. Insects and other invertebrates intercepted in check inspections of imported plant material in England and Wales during 1976 and 1977. Plant Pathology Laboratory. Harpenden, Herts. UK, [2+] 54 pp.

                                                                                              Smith IM, McNamara DG, Scott PR, Holderness M, 1997. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization., Ed. 2:vii + 1425 pp.; many ref.

                                                                                              Distribution References

                                                                                              CABI, Undated. Compendium record. Wallingford, UK: CABI

                                                                                              EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. https://gd.eppo.int/

                                                                                              Marchiori C H, Oliveira A M S, Costa M C R, 2002. Insects collected in maize crop in Itumbiara, south of Goiás state, Brazil. (Insetos coletados em cultivar de milho em Itumbiara, sul de Goiás, Brasil.). Arquivos do Instituto Biológico (São Paulo). 69 (Suplemento, Resumos), 233-234. http://www.biologico.sp.gov.br/ARQUIVOS/V69_supl_RE/marchiori.PDF

                                                                                              Murúa M G, Scalora F S, Navarro F R, Cazado L E, Casmuz A, Villagrán M E, Lobos E, Gastaminza G, 2014. First record of Helicoverpa armigera (Lepidoptera: Noctuidae) in Argentina. Florida Entomologist. 97 (2), 854-856. http://www.fcla.edu/FlaEnt/ DOI:10.1653/024.097.0279

                                                                                              Olivares T S, Angulo A O, Badilla Q R, 2012. Taxonomic notes and new register of moths for Easter Island (Lepidoptera: Noctuoidea). Entomological News. 122 (2), 157-164. http://www.bioone.org/loi/entn DOI:10.3157/021.122.0208

                                                                                              Links to Websites

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                                                                                              GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

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