Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Leptinotarsa decemlineata
(Colorado potato beetle)

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Datasheet

Leptinotarsa decemlineata (Colorado potato beetle)

Summary

  • Last modified
  • 20 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Host Animal
  • Preferred Scientific Name
  • Leptinotarsa decemlineata
  • Preferred Common Name
  • Colorado potato beetle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Colorado beetle principally attacks an introduced field crop grown as a monoculture, but not to an extent that has affected the area of the crop grown. It is not accordingly invasive in the usual environmental sense....

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Pictures

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PictureTitleCaptionCopyright
Leptinotarsa decemlineata (Colorado potato beetle); adult on potato stem. USA. Body length 8.5-11.5 mm.
TitleAdult
CaptionLeptinotarsa decemlineata (Colorado potato beetle); adult on potato stem. USA. Body length 8.5-11.5 mm.
CopyrightPublic Domain - released by the USDA-ARS/original image by Scott Bauer
Leptinotarsa decemlineata (Colorado potato beetle); adult on potato stem. USA. Body length 8.5-11.5 mm.
AdultLeptinotarsa decemlineata (Colorado potato beetle); adult on potato stem. USA. Body length 8.5-11.5 mm.Public Domain - released by the USDA-ARS/original image by Scott Bauer
Leptinotarsa decemlineata (Colorado potato beetle); body strongly convex dorsally with large abdomen; colour changing with development, first instar cherry-red with shiny, black head and legs; later instars becoming progressively carrot-red, then pale orange in final instar.
TitleLarvae
CaptionLeptinotarsa decemlineata (Colorado potato beetle); body strongly convex dorsally with large abdomen; colour changing with development, first instar cherry-red with shiny, black head and legs; later instars becoming progressively carrot-red, then pale orange in final instar.
Copyright©C. Trouvé/SRPV, Loos-en-Gohelle
Leptinotarsa decemlineata (Colorado potato beetle); body strongly convex dorsally with large abdomen; colour changing with development, first instar cherry-red with shiny, black head and legs; later instars becoming progressively carrot-red, then pale orange in final instar.
LarvaeLeptinotarsa decemlineata (Colorado potato beetle); body strongly convex dorsally with large abdomen; colour changing with development, first instar cherry-red with shiny, black head and legs; later instars becoming progressively carrot-red, then pale orange in final instar.©C. Trouvé/SRPV, Loos-en-Gohelle
Leptinotarsa decemlineata (Colorado potato beetle); head, pronotum and venter yellow-orange with black markings, legs and scutellum orange-yellow, elytra yellow-orange with five longitudinal black stripes. Body length 8.5-11.5 mm.
TitleAdult
CaptionLeptinotarsa decemlineata (Colorado potato beetle); head, pronotum and venter yellow-orange with black markings, legs and scutellum orange-yellow, elytra yellow-orange with five longitudinal black stripes. Body length 8.5-11.5 mm.
Copyright©C. Trouvé/SRPV, Loos-en-Gohelle
Leptinotarsa decemlineata (Colorado potato beetle); head, pronotum and venter yellow-orange with black markings, legs and scutellum orange-yellow, elytra yellow-orange with five longitudinal black stripes. Body length 8.5-11.5 mm.
AdultLeptinotarsa decemlineata (Colorado potato beetle); head, pronotum and venter yellow-orange with black markings, legs and scutellum orange-yellow, elytra yellow-orange with five longitudinal black stripes. Body length 8.5-11.5 mm. ©C. Trouvé/SRPV, Loos-en-Gohelle

Identity

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Preferred Scientific Name

  • Leptinotarsa decemlineata Say, 1824

Preferred Common Name

  • Colorado potato beetle

Other Scientific Names

  • Chrysomela decemlineata Say
  • Doryphora decemlineata Say
  • Leptinotarsa multitaeniata Stål
  • Polygramma decemlineata Say

International Common Names

  • English: Colorado beetle
  • Spanish: catarinita de la papa; dorifora; escarabajo de las hojas de patata; escarabajo del Colorado; mayata o catarinita de la papa; tortuguilla de la papa de Colorado
  • French: chrysomèle de la pomme de terre; doryphore de la pomme de terre
  • Russian: Koloradskii kartofel'nyi zhuk
  • Portuguese: doriforo; escaravelho da batateira

Local Common Names

  • Denmark: coloradobille; kartoffelbille
  • Finland: koloradokuoriainen
  • Germany: Kartoffelkäfer; Kolorado-käfer
  • Hungary: burgonyabogar
  • Iran: susske sibsamini
  • Italy: crisomela della patata; dorifora delle patate; scarabeo del Colorado
  • Netherlands: Coloradokever
  • Norway: koloradobille
  • Sweden: koloradoskalbagge
  • Turkey: patetes bocegi

EPPO code

  • LPTNDE (Leptinotarsa decemlineata)

Summary of Invasiveness

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Colorado beetle principally attacks an introduced field crop grown as a monoculture, but not to an extent that has affected the area of the crop grown. It is not accordingly invasive in the usual environmental sense. It has no effects on the environment.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Chrysomelidae
  •                             Genus: Leptinotarsa
  •                                 Species: Leptinotarsa decemlineata

Notes on Taxonomy and Nomenclature

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A comprehensive list of the synonyms of L. decemlineata is provided by Jacques (1988).

Description

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Eggs

The yellow or pale-orange, elongate-oval eggs, are ca 1.2 mm long. They are laid in groups of 12-25 on the underside of potato leaves. The females glue them to the leaf by one end using a special secretion. The long axis of the egg is almost perpendicular to the leaf. Eggs within a mass tend to form irregular rows and hatch simultaneously.

Larvae

Body strongly convex dorsally, with large abdomen. Head, bearing 6 ocelli behind the antenna on each side and a pair of 5-dentate mandibles. Three thoracic segments, each bearing a pair of 3-segmented legs, plus claw. Abdomen 9-segmented. Colour changing with development, first instar cherry-red with shiny, black head and legs; later instars becoming progressively carrot-red, then pale orange in final instar.

Head, legs and posterior part of pronotum black to deep brown; two conspicuous rows of dark spots occur on the lateral aspects of the mesothoracic and abdominal segments 1 to 7, the uppermost surrounding the spiracles, and also segments 8 and 9 with dark dorsal plates. Setae when present are very small, some occur on the head, legs, pronotum, on the pigmented areas and ventrally. Spiracles small, annular with black peritremes and situated on the mesothorax and first 8 abdominal segments. Body length of full-grown larva about 15 mm.

A detailed generic diagnosis of Leptinotarsa larvae is provided by Cox (1982) and the first instar is described by Peterson (1951). The weights of the four larval instars are given by Balachowsky (1963).

Pupae

Yellowish, bearing short setae on low, conical, brown tubercles. Head bearing several short setae, mandibles apically unidentate. Thorax with pronotum bearing about 100 setae; meso- and metathorax much more sparsely setose; apices of femora bearing about 3-5 setae and apical tarsal segment 1 seta. Abdominal segments 1-6 with lateral expansion dorsal to spiracle, dorsally bearing about 48 short setae, laterally about 9 setae on large papilla ventral to spiracle. Apical abdominal segment bearing a single, brown, median, sharply-pointed urogomphus or spine. Spiracles situated on mesothorax and abdominal segments 1-8; peritremes dark brown, but pale on abdominal segments 6-8. For further details, see Cox (1996).

Adults

Head, pronotum and venter yellow-orange with black markings, legs and scutellum orange-yellow, elytra yellow-orange with five longitudinal black stripes. Apical segment of maxillary palpi cylindrical, rounded apically, shorter than preceding segment. Elytra punctate-striate, epipleura glabrous. Mesosternum not raised above level of prosternum. Profemora normal, third tarsal segment entire, tarsal claws simple, divergent, not fused basally. Body length 8.5-11.5 mm.

The genus was revised by Jacques (1988). A key to the North American species is given by Wilcox (1972) and Jacques (1985).

Distribution

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The beetle was first discovered by Thomas Nuttall in 1811 and was described in 1824. Suddenly, in 1859, the Colorado potato began devastating potato crops 100 miles west of Omaha, Nebraska, USA (Pope and Madge, 1984). Whether the attacks stemmed from a change in food preference by the beetle, or were the result of its first meeting with the cultivated potato, remains uncertain. Over the next few years, the beetle caused crippling damage as it spread eastward to the Atlantic coast, which it reached in 1874 (Pope and Madge, 1984).

L. decemlineata became established in Europe following its introduction from the USA to Bordeaux, France in 1922 (after several unsuccessful attempts from 1876). The beetle spread rapidly in Europe despite intensive control operations to contain it. It was first reported in Belgium and Spain in 1935, Luxembourg in 1936, the Netherlands and Switzerland in 1937, Austria in 1941, Hungary and the former Czechoslovakia in 1945, Poland and Romania in 1947, and Turkey in 1949. It is widespread in the European part of the former USSR, and has progressively spread eastwards to most potato-growing areas, reaching the Far Eastern provinces (and is no longer a quarantine pest for Russia for that reason). L. decemlineata was detected in Xinjiang, China in 1993 (Guo et al., 2010; Liu et al., 2012).

A record of L. decemlineata in Zhejiang (CABI/EPPO, 2003; EPPO, 2006) published in previous versions of the Compendium was based on a paper by Mo and Cheng (2003) which only refers to laboratory studies on the pest resistance to insecticides. L. decemlineata has not been observed in the field in Zhejiang. 

L. decemlineata has been reported from Denmark, Finland, Norway, Sweden and the UK, including the islands of Guernsey and Jersey (Thomas and Wood, 1980), but the beetle is not established in these countries.

See also CABI/EPPO (1998, No. 92).

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaPresentAzaryan et al., 1985; Nalbandyan, 1987; CABI/EPPO, 2003; EPPO, 2014
AzerbaijanPresentAkhmedhov, 1980; Salmanov, 1988; CABI/EPPO, 2003; EPPO, 2014
ChinaRestricted distributionCABI/EPPO, 2003; EPPO, 2014
-XinjiangPresentJolivet, 1991; CABI/EPPO, 2003; Guo et al., 2010; Liu et al., 2012; EPPO, 2014
-ZhejiangAbsent, invalid recordCABI/EPPO, 2003; EPPO, 2014
Georgia (Republic of)PresentMachavariani, 1987; Kipiyani et al., 1989; CABI/EPPO, 2003; EPPO, 2014
IranPresentGanbalani, 1989; CABI/EPPO, 2003; EPPO, 2014
IraqRestricted distributionEPPO, 2014
JapanAbsent, intercepted onlyEPPO, 2014
KazakhstanRestricted distributionVlasova, 1978; Kim and Vasil'ev, 1986; Jolivet, 1991; CABI/EPPO, 2003; EPPO, 2014
KyrgyzstanPresentVlasova, 1978; CABI/EPPO, 2003; EPPO, 2014
TajikistanRestricted distributionLebedev and Smetnik, 1983; CABI/EPPO, 2003; EPPO, 2014
TurkeyRestricted distribution****CABI, 1992; Gürkan and Bosgelmez, 1984; CABI/EPPO, 2003; EPPO, 2014; Gözel and Gözel, 2014
TurkmenistanPresentKaraev, 1974; Pilenkova et al., 1988; CABI/EPPO, 2003; EPPO, 2014
UzbekistanRestricted distributionLebedev and Smetnik, 1983; CABI/EPPO, 2003; EPPO, 2014

Africa

LibyaAbsent, invalid recordBen Saad & Bishop, 1976; EPPO, 2014

North America

CanadaRestricted distributionCABI/EPPO, 2003; EPPO, 2014
-AlbertaPresentMcDonald, 1976; Harrison and Mitchell, 1988; CABI/EPPO, 2003; EPPO, 2014
-British ColumbiaRestricted distributionJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-ManitobaPresentSenanayake and Holliday, 1988; CABI/EPPO, 2003; EPPO, 2014
-New BrunswickPresentBoiteau et al., 1987; CABI/EPPO, 2003; EPPO, 2014
-Nova ScotiaPresentAnon., 1950; CABI/EPPO, 2003; EPPO, 2014
-OntarioPresentTurnball et al., 1988; CABI/EPPO, 2003; EPPO, 2014
-Prince Edward IslandPresentAnon., 1950; CABI/EPPO, 2003; EPPO, 2014
-QuebecPresentJacques, 1988; Mailloux and Bostanian, 1989; CABI/EPPO, 2003; EPPO, 2014
-SaskatchewanPresentAnon., 1950; CABI/EPPO, 2003; EPPO, 2014
MexicoWidespreadJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
USAWidespread1811CABI/EPPO, 2003; EPPO, 2014
-AlabamaPresentChittenden, 1911; CABI/EPPO, 2003; EPPO, 2014
-ArizonaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-ArkansasPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-ColoradoPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-ConnecticutPresentHare, 1984; Hare and Moore, 1988; CABI/EPPO, 2003; EPPO, 2014
-DelawarePresentHough-Goldstein and Whalen, 1993; CABI/EPPO, 2003; EPPO, 2014
-FloridaPresentJacques, 1985; CABI/EPPO, 2003; EPPO, 2014
-GeorgiaPresentChalfant & Young, 1984; Ghidiu and Oetting, 1987; CABI/EPPO, 2003; EPPO, 2014
-IdahoPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-IllinoisPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-IndianaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-IowaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-KansasPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-KentuckyPresentAnon., 1938; CABI/EPPO, 2003; EPPO, 2014
-LouisianaPresentCABI/EPPO, 2003; EPPO, 2014
-MainePresentStorch and Dill, 1987; CABI/EPPO, 2003; EPPO, 2014
-MarylandPresentSchroder & Athanas, 1989a; Schroder & Athanas, 1989b; CABI/EPPO, 2003; EPPO, 2014
-MassachusettsPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-MichiganPresentJansson et al., 1989; Drummond et al., 1990; CABI/EPPO, 2003; EPPO, 2014
-MinnesotaPresentCutkomp et al., 1958; Jacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-MississippiPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-MissouriPresentRiley, 1869; CABI/EPPO, 2003; EPPO, 2014
-MontanaPresentCABI/EPPO, 2003; EPPO, 2014
-NebraskaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-New HampshirePresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-New JerseyPresentCarter and Ghidiu, 1988; Williams, 1988; CABI/EPPO, 2003; EPPO, 2014
-New MexicoPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-New YorkPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-North CarolinaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-North DakotaPresentPost, 1954; CABI/EPPO, 2003; EPPO, 2014
-OhioPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-OklahomaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-OregonPresentCABI/EPPO, 2003; EPPO, 2014
-PennsylvaniaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-Rhode IslandPresentDrummond et al., 1987; CABI/EPPO, 2003; EPPO, 2014
-South CarolinaPresentChittenden, 1911; CABI/EPPO, 2003; EPPO, 2014
-South DakotaPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-TennesseePresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-TexasPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-UtahPresentHackett and Clark, 1989; Hackett et al., 1992; CABI/EPPO, 2003; EPPO, 2014
-VirginiaPresentZehnder and Evanylo, 1989; Tisler and Zehnder, 1990; CABI/EPPO, 2003; EPPO, 2014
-WashingtonPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-West VirginiaPresentCABI/EPPO, 2003; EPPO, 2014
-WisconsinPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014
-WyomingPresentJacques, 1988; CABI/EPPO, 2003; EPPO, 2014

Central America and Caribbean

Costa RicaAbsent, formerly presentEPPO, 2014
CubaPresentWeiser and Hostounsky, 1967; CABI/EPPO, 2003; EPPO, 2014
GuatemalaPresentKing and Saunders, 1984; CABI/EPPO, 2003; EPPO, 2014

Europe

AlbaniaPresentCABI/EPPO, 2003; EPPO, 2014
AndorraPresentPetitpierre, 1988; CABI/EPPO, 2003; EPPO, 2014
AustriaWidespread1944Cate et al., 1985; CABI/EPPO, 2003; EPPO, 2014
BelarusWidespreadKorol' et al., 1994; CABI/EPPO, 2003; EPPO, 2014
BelgiumWidespread1935CABI/EPPO, 2003; EPPO, 2014
Bosnia-HercegovinaPresentCABI/EPPO, 2003; EPPO, 2014
BulgariaWidespread1958Nedyalkov, 1984; Bozhkov, 1985; CABI/EPPO, 2003; EPPO, 2014
CroatiaWidespreadCABI/EPPO, 2003; EPPO, 2014
CyprusAbsent, confirmed by surveyEPPO, 2014
Czech RepublicWidespread1945Jelinek, 1993; CABI/EPPO, 2003; EPPO, 2014
DenmarkEradicatedAnon., 1979; Bejer and Esbjerg, 1980; EPPO, 2014
EstoniaWidespreadCABI/EPPO, 2003; EPPO, 2014
FinlandRestricted distributionEkbom, 1982; CABI/EPPO, 2003; EPPO, 2011; EPPO, 2014
FranceWidespread1922Davauchelle, 1984; Ribault, 1984; CABI/EPPO, 2003; EPPO, 2014
-CorsicaPresentEPPO, 2014
GermanyWidespread1936Hilbert & Kurth,1987; Kurth, 1987; Kurth, 1988; CABI/EPPO, 2003; EPPO, 2014
GreeceRestricted distribution1963Soultanopoulou-Mantaka, 1981; CABI/EPPO, 2003; EPPO, 2014
GuernseyAbsent, confirmed by surveyEPPO, 2014
HungaryWidespread1947Kaisler, 1988; Fodor et al., 1989; CABI/EPPO, 2003; EPPO, 2014
IrelandAbsent, intercepted onlyEPPO, 2014
ItalyWidespread1947Pucci and Dominici, 1988; CABI/EPPO, 2003; EPPO, 2014
-SardiniaAbsent, invalid recordEPPO, 2014
-SicilyPresentEPPO, 2014
LatviaPresentGebauer and Shilova, 1968; CABI/EPPO, 2003; EPPO, 2014
LithuaniaWidespreadZeimantiene, 1973; Bartinkaite and Babonas, 1985; CABI/EPPO, 2003; EPPO, 2014
LuxembourgRestricted distribution1936Anon., 1968; CABI/EPPO, 2003; EPPO, 2014
MacedoniaPresentCABI/EPPO, 2003; EPPO, 2014
MaltaAbsent, no pest recordEPPO, 2014
MoldovaRestricted distributionCABI/EPPO, 2003; EPPO, 2014
NetherlandsWidespread1936NPPO of the Netherlands, 2013; Anon., 1978; CABI/EPPO, 2003; EPPO, 2014
NorwayAbsent, intercepted onlyAnon., 1977; EPPO, 2014
PolandWidespread1946Pruszynski et al., 1987; Godzinska, 1989; CABI/EPPO, 2003; EPPO, 2014
PortugalWidespread1943Anon., 1968; CABI/EPPO, 2003; EPPO, 2014
-AzoresAbsent, confirmed by surveyEPPO, 2014
-MadeiraAbsent, confirmed by surveyEPPO, 2014
-Portugal (mainland)WidespreadCABI/EPPO, 2003
RomaniaWidespreadBob et al.,1984; Stef & Buzinovschi,1982; CABI/EPPO, 2003; EPPO, 2014
Russian FederationRestricted distribution1949CABI/EPPO, 2003; EPPO, 2014
-Central RussiaPresentCABI/EPPO, 2003; EPPO, 2014
-Eastern SiberiaPresentCABI/EPPO, 2003; EPPO, 2014
-Russian Far EastPresentCABI/EPPO, 2003; EPPO, 2014
-Southern RussiaPresentCABI/EPPO, 2003; EPPO, 2014
-Western SiberiaPresentCABI/EPPO, 2003; EPPO, 2014
SerbiaWidespreadEPPO, 2014
SlovakiaPresent1945CABI/EPPO, 2003; EPPO, 2014
SloveniaWidespreadCABI/EPPO, 2003; EPPO, 2014
SpainWidespread1935Anon., 1968; Del Rivero et al., 1969; CABI/EPPO, 2003; EPPO, 2014
-Balearic IslandsRestricted distributionJolivet, 1953; EPPO, 2014
-Spain (mainland)WidespreadCABI/EPPO, 2003
SwedenEradicated1972Grõnsbo, 1980; Wiktelius, 1985; CABI/EPPO, 2003; EPPO, 2014
SwitzerlandPresent1937Murbach, 1975; CABI/EPPO, 2003; EPPO, 2014
UKEradicated1901Seymour et al., 1985; CABI/EPPO, 2003
-Channel IslandsEradicatedEPPO, 2014
-England and WalesEradicatedEPPO, 2014
-Northern IrelandAbsent, intercepted onlyEPPO, 2014
-ScotlandAbsent, intercepted onlyEPPO, 2014
UkraineWidespreadKoval, 1986; Kovtun, 1966; Kharsun et al., 1990; CABI/EPPO, 2003; EPPO, 2014
Yugoslavia (Serbia and Montenegro)WidespreadZabel and Kostic, 1988; CABI/EPPO, 2003

Risk of Introduction

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Because of its capacity for adaptation to different climatic conditions (Ushatinskaya and Ivanchik, 1982) and different host plants (Hsiao, 1982), L. decemlineata is constantly moving into fresh areas and crossing international borders. The beetle has obviously not reached the extent of its possible geographic range in the EPPO region but its spread has slowed considerably in recent years, almost entirely due to international collaborative action, for example, between France and the Channel Islands, with EPPO support (Portier, 1980). The British Isles, the Nordic countries, and some other European islands, maintain themselves free through the EU system of 'protected zones'. In Russia and other CIS countries, where L. decemlineata has spread eastwards to reach the Pacific, an attempt was made (Vlasova, 1978) to estimate the potential final distribution; it was assumed that the requirement for one full generation would be a period in summer of at least 60 days of temperature over 15°C and winter temperatures not falling below -8°C. Establishment is not likely in colder areas of the EPPO region where only one partial generation could develop. Similarly, Worner (1988) tried to predict where L. decemlineata could establish in New Zealand. Potential distribution has been discussed by Jolivet (1991) for Asia and by Sutherst (1991) for the world.

A cost/benefit analysis performed by Aitkenhead (1981) indicated that the cost of the measures used to exclude L. decemlineata from the UK was less than the likely costs of control if introduced.

It should be stressed that L. decemlineata readily, though slowly, spreads by its own natural means across land areas where it has become established, and cannot be stopped by phytosanitary measures. It is mainly in this way that it has colonized Europe and northern Asia. It cannot normally cross such natural barriers as seas (except over relatively short distances, cf. Means of Movement and Dispersal), mountains and deserts. Outbreaks of Colorado beetle are relatively easily detected (it is a conspicuous insect) and eradicated. The insect does not survive on natural vegetation. As a consequence, uninfested areas which are protected by geographical barriers have excellent prospects of permanently excluding the pest by the regular application of phytosanitary measures. This applies to most of the uninfested areas in the world, with the exception of eastern Asia, where there is a significant risk of invasion by overland movement from northern Asia.

L. decemlineata is regulated by the European Union (EU, 2000) and by other EPPO countries (Belarus, Turkey). Although the pest is already widespread in Europe, certain areas where it does not occur are designated as 'protected zones'. L. decemlineata is treated as a quarantine pest by practically all regions of the world (Africa, Asia, Central America and Caribbean, South America, South Pacific), with the exception of North America, its area of origin.

Hosts/Species Affected

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L. decemlineata attacks potatoes and various other cultivated crops including tomatoes and aubergines. It also attacks wild solanaceous plants, which occur widely and can act as a reservoir for infestation. The adults feed on the tubers of host plants in addition to the leaves, stems and growing points.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Hyoscyamus niger (black henbane)SolanaceaeOther
Nicotiana tabacum (tobacco)SolanaceaeOther
SolanaceaeSolanaceaeWild host
Solanum lycopersicum (tomato)SolanaceaeOther
Solanum melongena (aubergine)SolanaceaeOther
Solanum tuberosum (potato)SolanaceaeMain

Growth Stages

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Symptoms

Top of page Adults and larvae feed on the edges of leaves and may quickly defoliate young plants. They eventually strip all leaves from the haulm; exceptionally, tubers exposed at the soil surface are also eaten. Characteristic black and sticky excrement is left on the stem and leaves by the larvae and adults.

List of Symptoms/Signs

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SignLife StagesType
Leaves / external feeding
Stems / external feeding
Vegetative organs / external feeding

Biology and Ecology

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The beetles overwinter as diapausing adults in the soil, typically at depths of 7.6 to 12.7 cm (Lashomb et al., 1984). Overwintered adult beetles emerge from the ground over a period of several weeks in spring or early summer, depending on the climate and their physiological condition (Hare 1990). Following emergence in the spring overwintered adults disperse to find suitable host plants by walking and by flight. Unfed beetles display greater flight activity than those that locate a suitable host and begin feeding soon after emergence (Ferro et al., 1999). Host plants are located largely by chance, through random searching. However, potato plant odour is attractive to the beetles and potato plants damaged by feeding are more attractive than undamaged plants, under laboratory conditions (Bolter et al., 1997; Landolt et al., 1999). Maximum food consumption occurs at 25°C.

Beetles typically mate before entering hibernal diapause, and also mate repeatedly in the spring, often within 24 h of emergence from the soil (Ferro et al., 1999). Sperm precedence in multiply mated females is incomplete (Alyokhin and Ferro, 1999). Oviposition begins 5-10 days after emergence at 15-30°C. Eggs are laid in masses, containing 10-30 eggs, on the lower surface of the leaf. Egg laying usually continues over several weeks, with each female laying up to 2000 eggs. The larvae hatch using egg bursters or oviruptors situated on the meso- and metathorax and abdominal segment 1 (Cox, 1988). They hatch in 4-14 days. After freeing themselves from the chorion, the larvae partly or entirely consume the chorion before feeding on leaf tissue. Larvae moult four times, the last of which is the larval/pupal moult. Larval development requires as little as 8 days or as long as 28 days at average temperatures of 29 and 14°C, respectively. Mature fourth-instar larvae burrow into the soil where they pupate. The pupal stage typically lasts 8 to 18 days, depending on temperature. Developmental thresholds, which range from 8 to 12°C, vary among populations and life stages. At constant temperatures, development is most rapid between 25 and 33°C; at higher temperatures larval growth is slowed and mortality increases (Hare, 1990).

The larvae are hardy and resistant to unfavourable weather, although heavy rain and strong winds may cause high mortality, especially in earlier instars. Cannibalism of eggs by adults can be considerable, averaging 19% in one study (Schrod et al., 1996). Cannibalism during the first instar is particularly common at high temperatures with low humidity but negligible under moderate conditions when suitable foliage is present.

Larvae from the same egg mass hatch synchronously and tend to remain grouped on the lower leaf surface until the first moult, after which they migrate to immature foliage on the plant. Larvae are voracious foliage feeders. Although total consumption depends on host plant, first instars consume ca 3% of the total foliage consumed during development and second, third and fourth instars consume ca 5, 15 and 77% of the total (Ferro et al., 1985). By the final or fourth instar, they may feed on petioles and stems, if the plants have become severely defoliated. The fully developed larvae descend to the ground and bury themselves in the soil at varying depths (usually to several centimetres) according to soil conditions. Pupation occurs in smoothly-lined cells and the pupal period lasts 10-20 days, after which the first-generation adults emerge. These adults may walk or fly to the nearest host plant to feed. After feeding for several days, adults of the first and subsequent generations may mate and reproduce, or cease feeding and enter diapause, depending on temperature, photoperiod and host plant condition. A portion of the population may produce some offspring before entering diapause (Tauber et al., 1988; Voss et al., 1988; Voss and Ferro, 1990a, b; Nault and Kennedy, 1999).

The number of complete generations varies between one, near the colder extremes, to about four, in the warmest areas where development from egg to adult is completed in 30 days. The minimum requirements for completion of one full generation are at least 60 days during summer when the temperatures exceed 15°C and winter temperatures that remain above -8°C. There are some cold areas in which only a partial generation is produced and the beetles cannot permanently become established. In general, sunny weather with a mean daily temperature of 17-20°C results in growth and spread of populations, but if temperatures do not exceed 11-14°C and humidity is high, the populations may actually decrease (Svikle, 1976).

In the northern part of the beetles' range in Europe, newly emerged first generation beetles feed and then burrow 25-40 cm into the soil, where they enter diapause and hibernate over winter. Mortality during hibernation averaged 30% in the Ukraine, but may be as high as 83%, mainly due to fungal and bacterial infections (Koval, 1984). A significant portion of pre-diapause adults migrate to field margins near tree lines before burrowing into the soil, although large numbers of beetles also enter the soil and overwinter within potato fields (Weber and Ferro, 1993; Weber et al., 1994; Nault et al., 1997).

In temperate regions, photoperiod is the most important factor inducing 'hibernal diapause' in teneral adults of L. decemlineata, but ambient temperatures and food quality may have modifying effects (Hsiao, 1988). This species is a typical 'long-day' insect entering diapause after exposure to a critically short photoperiod, which varies with latitude. In general, populations from the south require a shorter photoperiod for diapause induction than those from the north (de Wilde and Hsiao, 1981).Critical photoperiods approach 16 hours for northern populations (latitude 45°N) (Tauber et al., 1988) and decline to about 12 hours for southern populations (latitude 32°N).

The influence of age or physiological condition of potato foliage on diapause induction and the activity of the corpora allata was demonstrated by De Wilde et al. (1969). The corpora allata of diapausing beetles were small and inactive, as expected, but were 26% smaller in beetles fed mature foliage than in beetles fed young potato foliage. In adults fed mature foliage, the difference in size of the corpora allata was correlated with a >50% reduction in mean daily egg production.

High temperatures reduce the beetle's sensitivity to photoperiod; a shorter photoperiod will induce diapause at higher temperatures (de Wilde and Hsiao, 1981). This allows the exploitation of high-latitude areas that have relatively mild conditions. Response to temperature varies among populations. For example, the critical photoperiod for diapause induction in an upstate New York population was longer, and the induction of diapause by low temperature was greater, than for populations from the warmer coastal areas of Long Island (Tauber et al., 1988). The removal of host plants seems to be a primary factor in the induction of diapause in some southern populations. For some potato beetle populations in North Carolina, potato harvest, which occurs during during late June when daylengths are at their longest, causes the beetles to burrow into the soil where they remain in a state of diapause until the following spring. The survival of these beetles until the following spring is positively related to the amount of time that they have access to host foliage before entering the soil (Nault and Kennedy, 1999).

Populations from warmer, arid regions exhibit an 'aestival diapause,' which is induced by shortage or deterioration of their host plants. These populations hibernate above ground and resume feeding and reproduction once moisture and host-plant conditions become favourable. The incidence of aestival diapause was significantly greater when adults were fed foliage of Solanum dulcamara, a less suitable host plant. Of the first-generation females reared entirely on this wild host, 66% entered diapause without producing a second generation, and all first-generation females reproduced when reared entirely on potato, S. tuberosum. This diapause induction on S. dulcamara is thought to have occurred in response to the generally lower and declining nutritional quality of the wild host as the season progressed (Hare, 1983).

Because diapause is induced by a combination of photoperiod, temperature and food quality, there can be substantial variation in the proportion of adults entering diapause within generations and within populations between seasons. Overwintered females oviposit over a period of weeks; hence, first-generation adults do not emerge synchronously. The earliest may emerge under conditions favouring immediate reproductive development (long days and high temperatures), whilst their siblings, a few weeks younger, may emerge under conditions inducing diapause. Consequently, early-emerging first-generation adults may produce a second generation, all of which will subsequently enter diapause, while the later emerging first-generation adults may enter diapause either immediately or after depositing a few eggs (Tauber et al., 1988; Voss et al., 1988).

Burrowing in the ground during hibernal diapause enables L. decemlineata to escape severe cold temperatures and contributes to its success as a persistent pest in temperate regions (Boiteau and Coleman, 1996). Adult beetles are capable of perennial diapause in extreme climates. These two strategies of adaptation provide optimal synchronization of the species to its host plants in diverse geographical regions.

Natural spread of the beetle over large areas occurs by windborne dispersal, particularly of the spring generation. Adults can be also carried over long distances in sea water. In addition, adults and larvae are readily transported on potato plants and tubers, and in all forms of packaging and transport. The international trade of fresh vegetables grown on land harbouring overwintering beetles is a common means of transporting L. decemlineata.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Amphimermis avoluta Parasite
Amphimermis volubilis Parasite
Anisodactylus signatus Predator
Apateticus cynicus Predator Larvae
Apateticus lineolatus Predator Larvae
Apiomerus pinctipes Predator
Ardea cinerea Predator
Arma custos Predator
Bacillus thuringiensis Pathogen Germany; Maryland
Bacillus thuringiensis galleriae Pathogen
Bacillus thuringiensis kurstaki Pathogen
Bacillus thuringiensis san-diego Pathogen
Bacillus thuringiensis tenebrionis Pathogen Larvae
Bacillus thuringiensis thuringiensis Pathogen
Bacillus thuringiensis tolworthi Pathogen
Beauveria bassiana Pathogen Adults/Larvae Maryland; New Jersey
Beauveria brongniartii Pathogen
Calathus halensis Predator
Carabidae Predator Adults/Eggs/Larvae/Pupae
Carabus hampei Predator
Chrysomelobia labidomerae Parasite Adults Mexico Solanum angustifolium; Solanum rostratum
Chrysoperla carnea Predator Bulgaria; Moldova; USSR
Chrysoperla plorabunda Predator
Chrysoperla rufilabris Predator
Chrysoperla sinica Predator
Coccinella septempunctata Predator
Coleomegilla maculata Predator
Coleomegilla maculata Predator
Collops quadrimaculatus Predator Eggs
Edovum puttleri Parasite Eggs Colombia; Italy; Maryland; Mexico; Russia; USA; USA; New Jersey; USSR aubergines; potatoes
Endoreticulatus fidelis Pathogen
Formica polyctena Predator Poland potatoes
Formica pratensis Predator Poland potatoes
Formica rufa Predator Poland potatoes
Formica rufibarbis Predator Poland potatoes
Harpalus rufipes Predator
Heterorhabditis bacteriophora Parasite
Heterorhabditis heliothidis Parasite
Hexamermis albicans Parasite
Hexamermis angustata Parasite
Hexamermis brevis Parasite
Hexamermis cornuta Parasite
Hexamermis pusilla Parasite
Hippodamia convergens Predator Eggs/Larvae
Hippodamia variegata Predator
Iridovirus Pathogen
Klebsiella ozaenae Pathogen
Klebsiella pneumonea Pathogen
Klebsiella rhinoscleromatis Pathogen
Lebia analis Predator Eggs
Lebia cyane Predator Eggs/Pupae
Lebia grandis Predator Eggs/Larvae/Pupae France potatoes
Lebia guatemalena Predator/parasite Eggs/Pupae
Lebia quadrinotata Predator/parasite Eggs/Pupae
Lebia rufosutura Predator/parasite Eggs/Pupae
Linobia coccinellae Predator Adults
Megacephala virginica Predator
Megaselia rufipes Parasite Adults
Meigenia mutabilis Parasite
Merops apiaster Predator
Metarhizium anisopliae Pathogen
Micrococcus lysodeikticus Pathogen
Myiopharus aberrans Parasite Adults USSR
Myiopharus americanus Parasite Adults USSR
Myiopharus doryphorae Parasite Larvae Mexico; Poland; USSR potatoes; Solanum angustifolium; Solanum rostratum
Myiopharus macellus Parasite Larvae
Nosema melasomae Pathogen
Oplomus dichrous Predator Eggs/Larvae Mexico Solanum angustifolium; Solanum rostratum
Paecilomyces farinosus Pathogen
Passer domesticus Predator
Perdix perdix Predator Adults/Larvae/Pupae
Perichaeta unicolor Parasite
Perillus bioculatus Predator Adults/Eggs/Larvae Belgium; Delaware; France; Germany; Greece; Italy; Mexico; Moldova; Poland; USSR; Yugoslavia potatoes; Solanum angustifolium; Solanum rostratum
Perillus circumcinctus Predator Larvae
Perillus confluens Predator Larvae
Phalangium opilio Predator
Phasianus colchicus Predator Adults/Larvae/Pupae
Pica pica Predator
Picromerus bidens Predator
Podisus connexivus Predator
Podisus maculiventris Predator Larvae
Pristionchus uniformis Parasite
Protacanthus milberti Predator Adults
Pselliopus cinctus Predator Larvae
Pterostichus chalcites Predator
Pterostichus cupreus Predator
Pterostichus melanarius Predator
Pterostichus sericeus Predator
Silpha obscura Predator
Sinea diadema Predator Larvae
Steinernema carpocapsae Parasite Hungary
Steinernema feltiae Parasite
Stiretrus anchorago Predator Larvae
Stiretrus fimbriatus Predator Larvae
Strongygaster triangulifera Parasite
Xenorhabdus nematophilus Pathogen
Xysticus kochii Predator
Zicrona caerulea Predator

Notes on Natural Enemies

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L. decemlineata is attacked by several hymenopterous and dipterous parasitoids; many predators including pentatomid, reduviid and mirid bugs; beetles, including ground beetles and ladybird beetles; lacewing larvae; and parasitic nematodes, protozoans, fungi and bacteria.

Two hymenopteran egg parasitoids, the Nearctic eulophid Edovum puttleri and the Palaearctic mymarid Anaphes flavipes, are reported from L. decemlineata, but the former is potentially more important with regard to biocontrol. The New World E. puttleri parasitizes eggs of L. decemlineata (Grissell, 1981; Jansson et al., 1987; Logan et al., 1987; Ruberson et al., 1987; Williams, 1987; Schroder and Athanas, 1989a), which become less suitable for parasitization and parasitoid development as they age (Ruberson et al., 1985; Hu et al., 1999). Although it has been released for biological control of the potato beetle in the potato-growing areas of the USA, E. puttleri has not been effective in potato because it lacks a hibernal diapause and is poorly adapted to the low temperatures that prevail during the period when first generation Colorado potato beetle eggs are present (Obrycki et al., 1985). It has, however, been used in inundative release programmes for biological control in aubergine (Lashomb et al., 1987).

Several Tachinidae are known to parasitize the Colorado beetle. Meigenia mutabilis attacks the larvae of L. decemlineata in Russia (Bjegovic, 1967) and Myiopharus doryphorae (Geismer, 1920) and M. macellus (Arnaud, 1978) attack the larvae in the USA. M. abberans (Trouvelot, 1932) and M. australis [M. americanus] (Reinhard, 1935) attack adults in the USA. M. doryphorae is larviparous and first to final-instar larvae of Colorado beetle are attacked (Tamaki et al., 1983a, b). Maximum parasitism rate in this host reached almost 75% in late August to September when the crop had already suffered damage. The effectiveness of the parasite was limited by its low abundance during the first generation of L. decemlineata. The development and behavioural ecology of M. doryphorae and M. aberrans are described by Lopez et al. (1997, 1998).

In Germany, the phorid Megaselia rufipes parasitized 0.2-19.0% of L. decemlineata adults depending on location; hibernation of the phorid occurs in the pupal state (Eisenschmidt, 1958).

The pentatomid heteropteran Podisus maculiventris (the spined soldier bug) is an indigenous, generalist predator commonly found throughout North America, east of the Rocky Mountains (Baker and Lambdin, 1985). This bug preys on eggs and larvae of the Colorado beetle in Illinois, Missouri, Iowa (Anon., 1868) and Delaware (Heimpel and Hough-Goldstein, 1992; Hough-Goldstein and McPherson, 1996). P. maculiventris is attracted to host plant volatiles produced in response to prey feeding (Dickens, 1999). In the field, P. maculiventris employs a search strategy that causes it to consume a constant number of prey, which is independent of prey density (O'Neil, 1997).

The pentatomid Perillus bioculatus (the double-eyed assassin bug) preys on the adults, eggs and larvae of L. decemlineata (Chittenden, 1911; Tamaki and Butt, 1978; Drummond et al., 1984). Prey consumption by an individual of P. bioculatus during its development from egg to adult averages 285 eggs, or 3.7 fourth-instar larvae or 5.1 adults of L. decemlineata (Franz and Szmidt, 1960). After hibernation, adults generally consume 0.7-0.8 adult potato beetles or 0.5 fourth-instar L. decemlineata per day. Like P. maculiventris, P. bioculatus responds to host plant volatiles produced in response to feeding by Colorado potato beetle (Dickens, 1999; Weissbecker et al., 1999). P. bioculatus contributes to beetle mortality but natural populations are generally ineffective in suppressing Colorado potato beetle densities, especially when beetle densities are high (Harcourt, 1971;Tamaki and Butt, 1978).

The generalist predator Euthyrhynchus floridanus, a pentatomid occurring widely in the southern states of the USA, Central and South America, was reported feeding on larvae of L. decemlineata in South Carolina (Chittenden, 1911). E. floridanus apparently is of limited importance as a predator of Colorado potato beetle (Oetting and Yonke, 1975).

The potential for classical biological control of L. decemlineata using natural enemies is suggested by recent collections of natural enemies from a wide range of climates and habitats in Mexico (Logan et al., 1987). A total of 18 species of natural enemies were encountered feeding on the Colorado potato beetle. The densities of Mexican populations of the beetle were typically low, suggesting that natural enemies may be limiting factors. Oplomus dichrous, an egg/larval predator, was the most common predator. O. dichrous is native to Central America, Mexico and south-western USA.

Drummond et al. (1987) attained 95% control in the field with a release ratio of about one O. dichrous adult to 40-50 Colorado potato beetle eggs. However, initial studies of O. dichrous undertaken in Rhode Island, USA, indicated that the bug was poorly adapted to temperate potato production regions because its populations were poorly synchronized with those of the Colorado potato beetle, its population growth rate was low under cool temperatures and it appeared to lack the ability to overwinter in the north-eastern USA.

Recent studies in the USA demonstrated that the Neuropteran Chrysoperla rufilabris can effectively prey on the eggs and larvae of Colorado potato beetle in both laboratory and field-cage tests. In field-cage experiments, releases of C. rufilabris larvae at a rate of 80940/ha resulted in an 84% reduction of the Colorado potato beetle population (Nordlund et al., 1991).

The carabid beetle Lebia grandis is thought to feed almost exclusively on the Colorado potato beetle (Lindroth, 1969). The adults forage nocturnally on potato foliage, where they prey on eggs and larvae (Hazzard et al., 1991). The larvae exhibit a parasitoid-like lifestyle by developing to maturity on a single larva or pupa of L. decemlineata. This carabid may be the most important endemic predator of the Colorado potato beetle (Heimpel and Hough-Goldstein, 1992).

The coccinellids C. maculata and Hippodamia convergens are polyphagous predators that feed on eggs and small larvae of Colorado potato beetle in North America (Riley, 1869; Franz, 1957; Groden et al., 1990; Cappaert et al., 1991). C. maculata is perhaps the most abundant predator of L. decemlineata in many potato production areas, and the only one consistently present wherever the beetle eggs were found in the field (Hazzard et al., 1991; Hilbeck and Kennedy 1996). It caused 40% mortality in the second generation in 1986, 38% in the first generation in 1987, and 58% in the second generation in 1987. In a series of greenhouse and field experiments, consumption of potato beetle eggs by adult C. maculata was inversely related to the population density of egg masses (Arpaia et al., 1997). However, under commercial potato production conditions the abundance of C. maculata and daily predation rate appeared to be independent of prey density (Hilbeck and Kennedy, 1996). The high mobility of C. maculata, relative to the insect herbivores on which it feeds, may contribute to its effectiveness as a biological control agent in agricultural ecosystems (Coll et al., 1994). However, the structure of the agroecosystem is an important determinant of the contribution of C. maculata to biological control within specific crop in an agroecosystem (Groden et al., 1990; Nault and Kennedy, 2000).

L. decemlineata is the natural host for the microsporidia Nosema leptinotarsae and Endoreticulatus (=Pleistophora) fidelis (Lipa, 1968; Brooks et al., 1988). N. leptinotarsae occurs in the haemolymph but infections of E. fidelis are limited to the epithelial cells of the midgut (Hostounsky and Weiser, 1975; Brooks et al., 1988). E. fidelis has been observed to cause a gradual decline in laboratory colonies of Colorado potato beetle. Most work has been conducted on L. decemlineata as an experimental host for other Nosema species.

Experimental infection by N. gastroideae causes a reduction in growth rate, apparently due to inadequate food intake (Hostounsky and Weiser, 1973). A high death rate was reported in L. decemlineata infected with N. gastroideae and N. polygrammae (Hostounsky and Weiser, 1975). Chrysomelid microsporidians can be transmitted not only from one insect to another of the same species (horizontal transmission) but also from one generation to another (vertical transmission). Horizontal transmission is oral, with the hosts becoming infected after swallowing spores (Hostounsky and Weiser, 1973; Hostounsky, 1984).

The fungi Beauveria bassiana and Scopulariopsis have been reported from L. decemlineata (Humber, 1992). Natural outbreaks of B. bassiana cause impressive levels of mortality in the Colorado potato beetle in the absence of pesticides. In Poland, B. bassiana was the dominant fungal pathogen infecting 21% of diapausing Colorado potato beetle (Mietkiewski et al., 1996). B. bassiana does not infect Colorado potato beetle eggs, although all other stages are susceptible (Long et al., 1998). Another fungus, Paecilomyces farinosus parasitizes a wide range of insect hosts, often causing epizootics. The potential of P. farinosus as a microbial insect pathogen was evaluated using L. decemlineata (Samsinakova and Kalakova, 1978).

The parasitic nematodes Steinernema (= Neoaplectana) carpocapsae and Heterorhabditis heliothidis have a wide host range and can kill their host within 48 hours after contact. They can also grow on artificial media, have a durable infective stage for storage and application, and are environmentally safe (Poinar, 1986). Significant control of L. decemlineata has been achieved using both S. carpocapsae and H. heliothidis (Wright et al., 1986) and up to 71% mortality was observed when S. carpocapsae was used alone (Toba et al., 1983). The DD-136 strain of S. carpocapsae was more effective against larval than adult L. decemlineata, suggesting that the physiological condition of the insects was an important factor (Webster, 1972).

The parasitic podapolipid mite Chrysomelobia labidomerae caused increased mortality and reduced fecundity of L. decemlineata and may be of value in biological control programmes (Eickwort, 1982). However, Drummond et al. (1984) reported up to 100% parasitization by C. labidomerae on several Leptinotarsa species in Mexico with no deleterious effects. The parasitic hemisarcoptid mite Linobia coccinellae has the same potential for control of the Colorado potato beetle as does C. labidomerae, even though it is not the natural host (O'Connor, 1982).

Spiroplasmas have been isolated from L. decemlineata. These host-specific, gut-inhabiting commensal organisms are being considered for development as a genetically engineered pathogen for biocontrol. The Colorado potato beetle spiroplasma (CPBS) was first isolated from larvae and adults of L. decemlineata, collected in Maryland, USA (Clark, 1982). This pathogen is widespread in Canada, the USA (Hackett and Clark, 1989; Hackett et al., 1992) Poland and Germany, and the Ukraine and Byelorussia (Lipa, 1991a, b). The incidence of infection in larvae and adults in the field varied from 0-100% in North America (Clark, 1982; Hackett et al., 1992) and 0-40% in Europe (Lipa, 1991a). The average incidence of infection in field- and insectary-reared L. decemlineata was 17 and 43%, respectively, in an extensive survey in North America; thus reflecting greater transmission among insectary-reared beetles, a density-related factor (Hackett et al., 1992).

The incidence of infection among first-generation larvae remained low (5%) throughout June in Maryland, USA, before increasing to 100% (Clark, 1982; Hackett and Clark, 1989) as vagile adults spread the spiroplasmas to adjacent plants. There is no evidence that the incidence of infection declines once populations are infected with the spiroplasma. However, the low prevalence of CPBS early in the season may increase the opportunity for an engineered microbe to establish because there would be less competition from wild-type CPBS.

Means of Movement and Dispersal

Top of page The main means of natural spread of the beetle over large areas is by wind-borne migration, particularly of the spring generation. Adults flying over sea water will eventually fall in the sea, but can survive in sea water for several days, and be washed onto beaches by the tide (e.g. in Sweden, Channel Islands).

Adults and larvae can readily be transported on potato plants and tubers, and in all forms of packaging and transport. Fresh vegetables (of non-host crops) grown on land harbouring overwintered beetles are a common means of transport in international trade. For a review of detections in imported consignments and in the field in the UK, see Bartlett (1980).

Colorado beetle contaminates means of transport (e.g. lorries) by walking, or flying, on board. As a result, it will most likely be found on the outside of packages. However, there is no special association with the packing material.

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsPossible, but difficult in practice to differentiate from the means of transport. Travellers do not Yes
Land vehiclesAdults are occasionally found in all sorts of road vehicles, which pass or park near infested fields Yes
Plants or parts of plantsIn Europe, adults are more often found contaminating non-host vegetables than host. In a crop rotati Yes
Soil, sand and gravelAdults can contaminate almost any bulk material moved in trade Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes adults Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants adults Pest or symptoms usually visible to the naked eye
Leaves adults; eggs; larvae Yes Pest or symptoms usually visible to the naked eye
Stems (above ground)/Shoots/Trunks/Branches adults; eggs; larvae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Roots
Seedlings/Micropropagated plants
True seeds (inc. grain)
Wood

Wood Packaging

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Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood
Solid wood packing material with bark
Solid wood packing material without bark

Impact

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Introduction

The Colorado beetle, L. decemlineata, is one of the most economically damaging insect pests of potato in the many countries where it now occurs (Hare, 1990). L. decemlineata adults and larvae indirectly reduce potato tuber yields by devouring foliage. If plants become entirely defoliated prior to tuber initiation, total crop loss will result. L. decemlineata will also attack tomato and aubergine. In many areas, it is the only pest of ware potato crops against which insecticides have to be applied.

L. decemlineata originated in southwestern North America where it utilized a variety of solanaceous weed species (Hsiao, 1981). In the first half of the 19th century, its host range was expanded to include potato, which was grown east of the Rocky Mountains as the western USA became settled. By the late 19th century, the distribution of L. decemlineata continued eastward throughout North America and by the early 20th century it had eventually spread into Europe and Asia (Hsiao, 1981). L. decemlineata is currently distributed between latitudes 15° and 60°N. It does not occur in general in tropical countries, nor in most of eastern Asia, Korea, Japan, India, northern Africa, or the temperate Southern Hemisphere (Vlasova, 1978; Worner, 1988; Jolivet, 1991).

Perhaps the greatest economic impact that L. decemlineata has had on agriculture has been since its development of resistance to insecticides. L. decemlineata has become resistant to >25 insecticides belonging to the traditional chemical classes (Forgash, 1981, 1985; Gauthier et al., 1981; Heim et al., 1990; Roush et al., 1990; Tisler and Zehnder, 1990; Bishop and Grafius, 1991; Georgiou and Lagunes-Tejeda, 1991). Factors responsible for insecticide resistance development in this pest are described in detail in Zehnder et al. (1994). In the late 1980s and early 1990s, potato growers in the eastern USA used as many as 12 insecticide applications averaging $145 per ha per season to manage L. decemlineata (Zehnder and Evanylo, 1989; Zehnder et al., 1995). Currently, the cost of controlling L. decemlineata infestations in the eastern USA averages between $138 and 368 per ha (Grafius, 1997). Because insecticide resistance in L. decemlineata populations is inevitable, agribusiness industries continue to invest millions of dollars into developing new insecticides and genetically modified crops that produce insecticidal toxins.

Crop yield and financial losses attributed to L. decemlineata are not frequently published nor are they discussed in major reviews of L. decemlineata biology and management (Lashomb and Casagrande, 1981; Radcliffe, 1982; Ferro and Voss, 1985; Hare, 1990; Zehnder et al., 1994). This may in part be due to the lack of controlled, replicated experiments in commercial fields required to document such information. Further, crops are often attacked by multiple pests and the contribution of yield loss due to a single pest species is often impossible to determine. One of the few examples in which losses due to L. decemlineata have been documented was in Michigan (Grafius, 1997). In 1994, Michigan potato growers spent $6.8 million to control L. decemlineata and still experienced losses in tuber yield totalling $7.0 million, which together was nearly 14% of the overall crop value. With the exception of Grafius 1997 study, there is a paucity of quantitative crop loss and financial loss information. Therefore, the remaining review of L. decemlineata economic impact on its hosts will include major findings that have advanced our knowledge in (1) predicting crop yield loss due to feeding injury and (2) developing economic injury levels and thresholds for use in managing this pest.

Economic injury levels and economic thresholds are major components in the decision-making process of pest management (Pedigo and Higley, 1992). Developing economic injury levels and thresholds requires knowledge of the market value of the crop, the cost of managing the pest and the crop yield response to pest density or damage. Since the late 1970s, considerable effort has been made in identifying tuber yield responses of potato to defoliation and density of L. decemlineata adults and larvae. Fewer studies have examined the yield/damage relationship in aubergine and tomato.

Crop Response to Defoliation by L. decemlineata

The response of solanaceous crops to defoliation by L. decemlineata varies considerably with the phenological stage of the plant. For example, tomato seedlings cannot recover from extensive feeding by L. decemlineata adults, but as plant canopy increases, the level of defoliation that can be tolerated also increases (Schalk and Stoner, 1979). Potato has been shown to be least susceptible to yield loss when defoliated very early or late in the season (Hare, 1980; Ferro et al., 1983). Hare (1980) and Zehnder and Evanylo (1989) demonstrated that potato could withstand high levels of defoliation within a few weeks before harvest. Many studies have shown that potato plants are least tolerant of defoliation during the bloom stage (Cranshaw and Radcliffe, 1980; Hare, 1980; Wellik et al., 1981; Ferro et al., 1983; Shields and Wyman, 1984; Dripps and Smilowitz, 1989; Senanayake and Holliday, 1990; Nault and Kennedy, 1996). The only exceptions were reported by Zehnder and Evanylo (1989) and Zehnder et al. (1995) who showed that potato was most sensitive to yield loss when defoliated during prebloom.

Igrc et al. (1999) stated that the level of yield loss provoked by L. decemlineata defoliation depends upon the correspondence between severe leaf damage and tuberization process. In multiple year field trials they reported yield increase on treated plots from 18% up to 817% in the year when complete defoliation occurred at the beginning of tuberization (i.e. pre-bloom). The initiation of tubers is the key developmental occurrence in the life of potato crop. If the leaf damage occurs in the early stage of tuberization, the yield loss can be very high and vice versa. In most cultivars tuberization corresponds with floral initiation. Significant variation in yield losses caused by L. decemlineata in field trials among years was also observed by Igrc Barcic et al. (2006).

Several approaches have been taken to describe potato response to defoliation in order to predict tuber yield losses. Some have modelled potato growth over an entire season in which defoliation was either continuous or occurred during a specific growth period (Logan and Casagrande, 1980; Logan, 1981; Elkington et al., 1985; Dripps and Smilowitz, 1989; Johnson et al., 1996). Others have described the relationship between potato tuber yield and defoliation when defoliation occurred during a specific plant growth stage (Nault and Kennedy, 1996; Nault and Kennedy, 1998). Cotty and Lashomb (1982) determined the relationship between yield of aubergine fruit with densities of L. decemlineata, whereas Cantelo and Cantwell (1983) described the response of tomato to simulated L. decemlineata feeding.

Development of Economic Injury Levels and Thresholds

Economic injury levels and thresholds have been developed for optimizing the use of insecticide applications to control L. decemlineata infestations. Economic injury levels and thresholds have been based on levels of defoliation as well as densities of larvae and adults. Defoliation-based thresholds have been reported to vary depending on the phenological stage of the plant. Zehnder et al. (1995) developed defoliation-based thresholds of 20% during plant emergence to early bloom, 30% during early to late bloom, and 60% during late bloom to harvest. Defoliation-based thresholds also have been reported to vary within a phenological stage. For example, Zehnder et al. (1995) recommended a threshold during the bloom stage as high as 30% for the cultivar Superior in Virginia, whereas Shields and Wyman (1984) recommended a threshold of only 10% for Superior and Russet Burbank in Wisconsin (Shields and Wyman, 1984). In North Carolina, the level of defoliation that Atlantic potato was shown to tolerate during bloom varied depending on the statistical approach used to develop the threshold (Nault and Kennedy, 1998). The level of defoliation deemed tolerable by potato was usually much higher using mean separation analysis than using regression. For example, the same data set analyzed using mean separation analysis indicated that potato could withstand 44% defoliation during bloom, whereas non-linear regression analysis indicated that potato could only tolerate 13% defoliation during bloom. Both types of analyses have limitations (see Nault and Kennedy, 1998).

The relationship between potato yield and insect density also has been modelled to develop economic injury levels (Hare and Moore, 1988; Senanayake and Holliday, 1990; Mailloux et al., 1991, 1995). Based on this approach, economic injury levels of 5.8 overwintered adults and 10 summer-generation adults per plant have been reported (Mailloux et al., 1995). For larvae, economic injury levels have been reported to be as high as 12 per stalk (Mailloux et al., 1991) and as low as 0.14 to 0.82 per plant (Senanayake and Holliday, 1990). In processing tomato cultivars in Maryland, yield loss did not result until the number of L. decemlineata adults and larvae exceeded 1.25 per plant and 2 per plant, respectively (Linduska and Dively, 1990).

Use of Economic Thresholds for Managing L. decemlineata

There are few studies in which the benefits of using economic thresholds for managing L. decemlineata have been reported. In several instances, however, potato tuber yield did not significantly differ between an economic-threshold-based management approach and the conventional management program. Yet, fewer insecticide applications were needed to manage L. decemlineata infestations when an action threshold was used (Wright et al., 1987; Stewart and Dornan, 1990; Zehnder et al., 1995).

Despite the success of using action thresholds to optimize the use of insecticides for managing L. decemlineata, preventative control approaches have been commonly used in the USA. For example, in-furrow applications of imidacloprid and use of transgenic potato containing the Bacillus thuringiensis subsp. tenebrionis endotoxin have been used to provide near season-long and season-long protection of L. decemlineata, respectively. Similarly, because L. decemlineata adults may completely devour tomato as soon as plants emerge or immediately after seedlings are transplanted, a preventative approach using insecticides at planting or transplanting is taken to prevent loss (Ghidiu, 1984; Ghidiu and Oetting, 1987).

Environmental Impact

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L. decemlineata invaded North America, and then Europe and Western Asia, in a classic pattern of regular geographical spread, hardly impeded by measures taken against it. However, this spread could not be called invasive because it occurred in an introduced crop planted over large areas as a monoculture. Though L. decemlineata attacks other Solanaceae, there are no indications that it affects wild plants in the natural environment to any significant extent. Control of the pest leads to the use of insecticides in potato crops, which most probably would not require such treatment in its absence. So to a certain degree L. decemlineata is responsible for an increased pesticide load in the environment.

Social Impact

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Though a serious pest of potato, L. decemlineata never caused such losses as those due to Phytophthora infestans, with their attendant social consequences. By the time of its introduction, peasant farmers were not so dependent on the single potato crop. In the early days, Colorado beetle could be partly controlled by hand removal and destruction of larvae and adults. Effective cheap insecticides fairly rapidly became available.

Detection and Inspection

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Adults and larvae are easily seen because of their large size. L. decemlineata has a tendency to release its hold on plants that are shaken and this characteristic can be used to detect insects hidden among foliage. Visual sampling of potato fields was as efficient for estimating population density as the whole-plant bag-sampling method, and more efficient than sweep netting (Senanayake and Holliday, 1988). Soil sampling at harvest for buried beetles in diapause provides reliable results in area surveys (Glez, 1983). A sequential sampling plan has been reported for estimating populations of Colorado potato beetle egg masses and of adults and larvae (Hamilton et al., 1997a).

Similarities to Other Species/Conditions

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The Central and South American L. undecimlineata is similar to L. decemlineata, but the scutellum, underside and legs (except the claws) are black. This species is more common at altitudes above 1000 m, feeding on various Solanum species, but is of less economic importance.

Prevention and Control

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Cultural Control

Delayed colonization of the crop reduces the time available for second and subsequent generations to develop (Voss et al., 1988). Delayed colonization could be achieved by various means, such as crop rotation, manipulation by planting date, setting up different barriers (such as plastic line trenches, portable trench barriers, mulching), and mechanical collection of beetles on overwintering sites or immediately after colonization.

Crop rotation delays colonization of the crop by overwintered adults and reduces the size of the population that subsequently develops within the crop. Population reductions of 90% or more have been reported in potato (Lashomb and Ng, 1984; Wright, 1984). The effect of crop rotation is increased with increased distance from the source of overwintering beetles. Follett et al. (1996) suggested 0.5 km as the minimum distance necessary to fully benefit from field rotation. Because rotated potato fields require fewer insecticides to control L. decemlineata (Speese and Sterrett, 1998), crop rotation is an important tool in delaying the development of resistance to insecticides (Roush et al., 1990). Saxon and Wyman (2005), in their suggestion of developing of an area-wide L. decemlineata pest management strategy, reported that long-distance rotations of more than 400 m were an effective cultural control management strategy to limit adult beetle infestations in the spring. This strategy can be optimized when collaborating growers are able to maximize their rotational distances by coordinating their rotational schemes over large areas. Deploying long-distance rotations over a large area over many years would limit L. decemlineata populations and could result in a signifcantly reduced L. decemlineata populations entering fields in the spring.

The planting date of a potato crop can be manipulated to reduce the population of second generation larvae produced in the crop. Early planting of short season varieties allows the crop to mature before the second larval generation is produced. In contrast, colonization of late planted, rotated potato plantings occurs later in the season causing most summer generation adults to emerge after the critical photoperiod for diapause induction has been reached. Consequently, these adults do not produce a second generation of larvae (Weber and Ferro, 1993).

Plastic-lined trenches, which serve as pitfall traps, and trap crops of early-planted potato have both been shown to effectively intercept overwintered adults in the spring, before they colonize the potato crop. Plastic-lined trenches that were V-shaped with an average width at the top of 740±7 mm and depth of 223±2 mm retained 95% of the beetles they trapped in controlled field experiments (Misener et al., 1993; Boiteau and Osborn, 1999). Because 50-75% of overwintered beetles disperse into a nearby potato crop by walking, properly designed trenches, positioned to intercept the dispersing beetles can provide a significant level of crop protection. At one location in New York, USA, more than 100,000 overwintered beetles were trapped in 91 m of trench (Moyer, 1993).

Overwintering beetles are frequently found in windbreaks and hedgerows adjacent to the crop at densities of hundreds per square metre, whereas densities within the fields average only 3-7 beetles per square metre (Weber and Ferro 1993, Hunt and Tan 2000). This distribution of overwintering beetles clearly has important implications for the management of L. decemlineata, particularly when another solanaceous crop is planned for the same or an adjacent field in the following year. For some growers crop rotation is not a viable option due to a shortage of land. Hunt and Vernon (2001) suggested that barriers placed along the margins of tomato and potato fields adjacent to preferred overwintering sites may be effective in preventing or slowing the entry of beetles into the crop in the spring. They designed an above-ground trench composed of an extruded, UV retarded PVC plastic trough, designed to allow L. decemlineata and other pests to enter the device and become trapped and killed inside. The above-ground trench can capture thousands of newly emerged L. decemlineata as they walk from their overwintering habitat into an adjacent crop, and also reduce crop damage. This trap is easily installed in the spring and removed in the fall, and is reusable for several years. The trench is black to raise the temperature, which results in an increase in beetle mortality, and is designed to release water and beneficial insects.

Mulching potato and aubergine plantings with straw has been shown to significantly reduce L. decemlineata infestations and defoliation, reduce the number of insecticide treatments needed and increase yield (Zehnder and Hough-Goldstein, 1990; Stoner, 1993; 1997). The mulch delays colonization and increases predation on eggs and larvae (Ng and Lashomb, 1983; Brust, 1994). Lower potato beetle populations and less damage to tomato has been reported in reduced tillage plantings than in conventional tilled plantings (Hunt, 1998).

Mechanical collection, use of propane flamers, use of pneumatic thermal machines or bio-collectors can prevent adult colonization and reduce larval damage (Boiteau et al., 1992; Karalus, 1994; Pelletier et al., 1995; Lague et al., 1999a, b; Derafshi, 2006). The bio-collector is a novel control method approved for use in organic potato cultivation in Germany. The collector, which is attached to a tractor, blows chrysomelids off the potato plants and collects them in trays. Collections are made two to three times per year depending on the level of infestation (Karalus, 1994). However, some authors stated that mechanical control causes undesirable damage and its efficacy should be improved (Sablon et al., 2013).

Although Boiteau et al. (2012) found in laboratory trials that wood ash is toxic to adult and larval stages of the L. decemlineata, the significant control observed in the laboratory did not extend to field application.

Semiochemical-based strategies

The chemical ecology of L. decemlineata is not yet completely understood and this incomplete knowledge makes semiochemical-based approaches inefficient when compared to traditional insecticide treatments (Sablon et al., 2013). According to Sablon et al. (2013a). Nevertheless, alternative strategies have potential to control L. decemlineata populations and include: (1) disorientation of L. decemlineata adults by masking potato volatile organic compounds (VOC) with intercropping cultures; (2) use of synthetic mixtures of volatiles and/or aggregation pheromone to trap beetles; (3) antifeedant sprays on potatoes; (4) increase, with genetic manipulations, the natural capacity of the plant to recognize the presence of L. decemlineata through chemical signals, thereby triggering defence mechanisms.

(1) Intercropping or companion planting may represent an efficient method to repel and/or confuse L. decemlineata foraging for host plants. Matthews et al. (1982) reported that the use of tansy as an intercrop in potato fields may result with 60-100% of decrease in the number of beetles present in the fields. The initial results reported by Thierry and Visser (1986; 1987) have shown that potato VOCs mixed with VOCs from tomato or cabbage disrupted searching behaviour of L. decemlineata females for host plants. Alghali et al. (2000) reported that L. decemlineata larvae were more frequently encountered in the pure potato plots compared with the potato/cabbage intercropped crops (either alternate rows of potato and cabbage or alternate row strips of potato and cabbage). Moreau et al. (2006) tested bush beans (Phaseolus vilgaris cv. Provider), flax (cv. Natasia), French marigold (Tagetes patula cv. Bolero), horseradish and tansy (Tanacetum vulgare) as companion plants to potatoes. In addition, some botanical and microbiological preparations (capsicain extract, garlic extract, neem extract, Bt product, adjuvant and pine extract) were evaluated. The results showed that companion planting and garlic and capsicain extracts did not reduce the densities of L. decemlineata on potatoes.

(2) The basic strategy of trap crops is aimed to attract L. decemlineata in a specific part of the field where potato plants are planted earlier or where potato plants are treated with semiochemicals. The next step is to treat only this part of field to eliminate attracted beetles.

In a small plot study in Massachusetts, USA, the colonization of potato fields by adults that overwintered in wooded borders of potato fields was reduced by about 60% when beetles in the trap crop were collected daily or killed with an insecticide (Weber et al., 1994). Trap crops have been shown to reduce L. decemlineata infestations and increase yields in tomato by 61-87% (Hunt and Whitfield, 1996).

L. decemlineata as a chewing insect is very sensitive to VOCs released by host plants, because the damage they induce in plant tissues increases the release of these compounds (Szendrei and Rodriguez Saona, 2010). Dickens (2000) investigated in laboratory trials behavioural responses of the L. decemlineata to volatiles emitted from solanaceous host plants, non-host legume plants and 13 synthetic blends of three individual chemicals emitted by potato plants. He found that L. decemlineata is attracted to blends of specific chemicals emitted by their host plants. In further investigations, Dickens (2002) reported a synthetic blend of three compounds ((Z)-3 hexenyl actetate. ±-linalool and methyl salicylate) which attracted second and fourth larval instars. This blend attracted newly emerged and 5-day-old adults in greenhouse trials (Martel et al., 2005a), and initial field tests confirmed the effect of the attractant blend and its usefulness for semiochemical-based control (Martel et al., 2005b). Pitfall traps baited with the attractant captured a greater number of L. decemlineata in potato fields. Moreover, fewer egg masses and larvae were observed in neighbouring untreated crops. The use of trap crops with the attractive blend allowed for a 44% pesticide reduction for similar yields compared to conventional methods (Martel et al., 2005b). In another study (Martel et al., 2007), this blend was coupled with pyrethroid insecticide at different concentrations. The use of this attracticide showed the same control efficiency as the commercial insecticide whilst using 92% less insecticide. Since many L. decemlineata populations have developed the resistance to permethrin, the authors suggested imidacloprid as a replacement.

A male-produced aggregation pheromone for L. decemlineata was identified as (S)-3, 7-dimethyl-2-oxo-oct-6-ene-1, 3-diol [(S)- L. decemlineata I] by Dickens et al. (2002). The pheromone has been shown to be attractive to L. decemlineata larvae (Hammock et al., 2007) as well as male and female adults (Dickens et al., 2002). Kuhar et al. (2006) demonstrated the attractiveness of (S)- L. decemlineata I in the field as well as its potential for integrated pest management of L. decemlineata. However, the synthetic routes for (S)- L. decemlineata I of high purity have been complicated and expensive. Therefore Kuhar et al. (2012) conducted pitfall trap studies to assess the relative attraction of L. decemlineata adults to synthetic mixtures of the (S) and (R) enentiomers of the pheromone. The results indicated that any further research as well as IPM strategies that incorporate L. decemlineata I as an aggregation pheromone should utilize blends containing more than 87% optical purity of the (S)-enentiomer of the pheromone. The main difficulty with semiochemicals is ensuring a controlled release during a long period (Sablon et al., 2013). The choice of dispenser is very important to control the release and to prevent molecule degradation (Heuskin et al., 2011).

(3) The basic concept of the use of antifeedants is to spray them on potatoes to deter L. decemlineata feeding and reduce damage of potato foliage. Many of antifeedants are constituent of plants from sagebrush community, hydroxides which also act as fungicides, alchocol extracts of the leaves and bark of Quercus alba L., limonin, sesquiterpenes (Sablon et al., 2013), α-mangostin (Kim and Lan, 2011), terpenoid lactones (Szczepanik et al., 2005) and extracts from various plants (Gokce et al., 2006a, Pavela et al., 2004, 2009) including potatoes (Szafranek et al., 2008). Neem extract, a well-recognized botanical insecticide, also shows antifeeding activity against L. decemlineata, but the magnitude of the effects depends on the dominant L. decemlineata life stage present when application was made and on attack intensity (Zehnder and Warthen, 1988; Bezjak et al., 2006; Igrc Barcic et al., 2006). Such molecules and blends of chemicals not only reduce feeding but also may deter oviposition by females as shown for saponins (Waligora-Rosada 2010) and citrus limonoids (Murray et al., 1995). Very often antifeedants are considered as synthetic insecticides whereas they originate from plants and should therefore be considered as botanical insecticides. According to Sablon et al. (2013a) this is one of the reasons why the commercialization of antifeedants has been generally unsuccessful.

Biological Control

Edovum puttleri has been released in several areas in the eastern USA and is being investigated for possible release in Europe (Schauff, 1991). Research on E. puttleri in relation to its potential for biological control of the Colorado potato beetle in the USA is reviewed by Schroder and Athanas (1989a, b) and Schroder et al. (1985).

The ability of augmentative releases of E. puttleri to control Colorado potato beetle was assessed on aubergine in New Jersey, USA (Lashomb et al., 1987) and Italy (Pucci and Dominici, 1988) and on potato in Maryland, USA (Schroder and Athanas, 1989a, b) and Ontario and New Brunswick, Canada (Sears and Boiteau, 1989). The most favourable results were obtained for aubergine in New Jersey (Lashomb et al., 1987) and for potato in Maryland (Schroder et al., 1985; Schroder and Athanas, 1989a, b). However, detailed comparisons between these studies were limited by the variation in numbers of wasps released and the methods used to evaluate the impact on the host populations (Van Driesche et al., 1991). Results reported by Van Driesche et al. (1991) for augmentative releases of E. puttleri against L. decemlineata on potatoes in Massachusetts, USA, confirmed the findings of Sears and Boiteau (1989) that releases of E. puttleri are ineffective against first-generation eggs. Parasitoid releases had greater impact on second generation eggs. The release of 47,000 wasps in 1988 against the second generation, without pesticide, resulted in 34.4% parasitism and 16.1% host feeding; a total reduction of 50.5%.

Releases of Podisus maculiventris, at 76,000/ha, reduced egg numbers of Colorado potato beetle in the second and third generations in field model experiments in Moldova, Russia (Novozhilov et al., 1991). The complex of natural enemies, including 26 carabids, 11 coccinellids and three chrysopids, added to the beneficial effect of the pentatomid. When L. decemlineata eggs were exposed to P. maculiventris and twelve-spotted ladybeetle (Coleomegilla maculata), predation on L. decemlineata eggs by both predators together did not increase significantly over levels inflicted by either predator alone (Mallampalli et al., 2002). Aldrich and Cantelo (1999) demonstrated that pheromones of P. maculiventris could be used to attract females and thereby enhance populations of the predator within a potato planting.

Augmentative releases of Perillus bioculatus controlled first-generation eggs and larvae of L. decemlineata in field-plot tests conducted during two seasons under short-season conditions in Quebec, Canada (Cloutier and Bauduin, 1995).

Sablon et al. (2011) reported on the results of laboratory assay in which the use of lacewing larva Chysoperla carnea allowed an efficient reduction of L. decemlineata eggs and larvae. Sablon et al. (2013b) have shown that the first and second instar of C. carnea only consumed eggs, whereas the third instar of consumed all L. decemlineata immature stages. The third instar killed four times as many L. decemlineata larvae than other larval stages. However, field assays are needed to confirm the efficiency of this natural enemy under field conditions.

The nematode Pristionchus uniformis reduced populations of L. decemlineata in greenhouse and field experiments in Poland (Fedorko and Stanuszek, 1971). Variable suppression of L. decemlineata populations was achieved in New Brunswick and Prince Edward Island, Canada, following application of the entomopathogenic nematode Steinernema carpocapsae (Stewart et al., 1998). Among the four investigated nematode species (Steinernema feltiae, S. carpocapsae, Heterorhabditis bacteriophora and H. megidis), Trdan et al. (2009) found the application of higher concentrations of S. feltiae to be the best for the control of overwintering adults for the purpose of preventing mass appearance. Adel and Hussein (2010) reported a significant reduction of the number of damaged leaves and lower index of damage achieved with Steinernema feltiae and Heterorhabditis bacteriophora foliage application. Ebrahimi et al. (2011) investigated the effect of the sublethal concentrations of the same nematode species on L. decemlineata. They found that sublethal nematode concentrations caused wing deformation and delayed metamorphosis which may affect L. decemlineata adult fitness and have long term effect on population pressure.

Most work on fungi for the biological control of chrysomelid hosts has focused on the use of Beauveria bassiana against the Colorado potato beetle. Attempts at controlling L. decemlineata with Beauveria in Russia and Eastern Europe began with large-scale research programmes in Russia that led to the mass production and relatively widespread use of Beauveria-based products, generally bearing the name 'boverin' (Ayleshina, 1978; Lipa, 1985). This Beauveria-based approach to the control of the Colorado potato beetle was most effective on low and moderate populations of the beetles. The extensive use of B. bassiana against L. decemlineata in biological control programmes in Russia during the 1970s and 1980s is summarized by Feng et al. (1994).

During the mid-1980s, a pilot test programme studying the efficacy of B. bassiana for the biocontrol of Colorado potato beetle on potatoes in the USA showed mixed results, but demonstrated that, at its best, the fungus could provide foliar protection and result in yields approaching those of pesticide-treated control plots (Watt and LeBrun, 1984; Hajek et al., 1987). In America, the use of B. bassiana to control L. decemlineata never achieved either the scale or the reported success of the Soviet effort and was eventually abandoned by the commercial firm that provided the formulated fungus for the pilot test programme.

Kryukov et al. (2014) established for the first time that the peroral effect of the fungal culture of Cordyceps militaris resulted in a dose-dependent decrease in survival, and delayed developmental time and moulting, as well as increased sensitivity of larvae to the fungus Beauveria bassiana.

Following the invasion of the potato-growing areas of Continental Europe, attempts were made to introduce North American natural enemies, known to be important mortality factors, to France, from 1929-1940. Two tachinids, Myiopharus doryphorae and M. aberrans; two pentatomids, Perillus bioculatus and Podisus maculiventris; and a carabid, Lebia grandis, were cultured and released, but none became established. Work resumed in 1957 under a co-operative programme involving 12 countries. Efforts concentrated on attempting to establish P. bioculatus, again without success; it was concluded that P. bioculatus was not adapted to conditions in Europe. Further introductions of M. doryphorae were also made in France but the European strain of L. decemlineata was resistant to parasitism by this tachinid (Greathead, 1976). Attempts at introducing natural enemies were then abandoned until the discovery of Endovum puttleri, which was imported into several countries and released but did not persist for long periods. Recent efforts in the Czech Republic to use Beauveria to control L. decemlineata have shown some success (Dirlbekova and Dirlbek, 1987; Dirlbek and Dirlbekova, 1987; Dirlbek, 1988; Dirlbekova et al., 1992).

Effective control of L. decemlineata was achieved in potato fields in New Jersey, USA, in 1989-1990, using foliar applications of Bacillus thuringiensis subsp. tenebrionis. The bacillus was sprayed when an average 1-30% of marked egg masses had hatched. Significantly greater defoliation, greater numbers of third- and fourth-instar larvae, and lower yields occurred in plots when the initial application was delayed until 6 days after 30% egg hatch, compared with plots treated at 30% egg hatch (Ghidiu and Zehnder, 1993). When applied properly, foliar applications of commercial formulations of B. thuringiensis subsp. tenebrionis provided effective control of L. decemlineata and protected the potato crop from defoliation. Because B. thuringiensis subsp. tenebrionis must be ingested to be effective, is most effective against small larvae, and has only limited residual activity once applied to foliage, proper timing of applications and thorough spray coverage of the crop are critical to effective control (Zehnder and Gelernter, 1989; Ferro and Lyon, 1991; Zehnder et al., 1992; Dubois and Jossi, 1993; Korol' et al., 1994; Ferro, 2000). Ghassemi-Kahrizeh and Aramideh (2014) have proven in laboratory assays that there is a remarkable synergistic effect of Henna powder on Bacillus thuringiensis efficiency.

In organic agriculture pest management, the mainstay for insect control, including L. decemlineata control, has been pyrethrum Chrysanthemum cinerariaefolium. This has a knock-down effect, but a short residual activity (Zehnder et al., 2007). Piperonil butoxide (PBO) is also used as a synergist for pyrethrins and the synthetic pyrethroid insecticides. Because pyrethrins are harmful to beneficial animals, such as predatory mites, parasitoids, and honey bees, the use of the synergist reduces the concentration of pyrethrins necessary for pest control. Owing to the concern over PBO, dillapiol is considered as a potential replacement. Dillapiol, which was first found in the Indian dill, Anethum sowa Roxb. ex Fleming (Apiaceae), is nearly as effective as PBO as a pyrethrin synergist (Joffe et al. 2012). In laboratory trials that pyrethrum efficacy was increased 2.2 times with the SS strain and 9.1 times with the RS strains of L. decemlineata by using pyrethrum + diallipol (Liu et al., 2014). In field trials with the pyrethrum+ diallipol formulation demonstrated efficacy ≥10 times than pyrethrum alone.

Besides pyrethrum extracts, many plant extracts and essential oils have been tested against L. decemlineata. Among the tested plants, Humulus lupulus extract has been proven by Gokce et al. (2006b) and by Cam et al. (2012) to be toxic to L. decemlineata larvae. The essential oils of Mentha longifolia (Rafiee-Dastjerdi et al., 2014), Letharia vulpina and Peltigera rufescens (Emsen et al., 2013) and Piper nigrum (Scott et al., 2003) can also control L. decemlineata.

Terpenes are a large and diverse class of organic compounds, produced by a variety of plants, particularly conifers. Several studies have shown that some of the monoterpens have potential for L. decemlineata control. Monoterpene hydrocarbons exhibited high toxicity as compared with oxygenated monoterpenes; 1,8-cineole, fenchone, β-pinene and γ-terpinene can be used as potential control agents against both L. decemlineata larvae and adults (Kordali et al., 2007). Khorram et al. (2011) tested 6 pure monoterpene hydrocarbons as a single compounds or mixtures; all six tested compounds (monoterpene hydrocarbons) can be used as potential control agents against both larvae and adults of L. decemlineata, either as single compounds or in mixtures. Mahdi et al. (2011) tested 12 pure oxygenated monoterpenes as two different doses for their toxicity against second and third instar of larvae and adults of L. decemlineata; out of 12 tested, oxygenated monoterpenes, fenchone, linalool, citronella and menthone showed a strong toxicity against the tested developmental stages.

Transgenic Plants

Transgenic potatoes expressing the gene for B. thuringiensis subsp. tenebrionis Cry3A delta -endotoxin were approved for commercial use in the USA in 1995. In 1998, transgenic potato varieties expressing the Cry3A toxin were planted on approximately 20,000 hectares in the USA. Although these transgenic potato varieties are highly toxic to Colorado potato beetle (Perlak et al., 1993; Wierenga et al., 1996) and provide excellent control of the beetle, planting of Cry3A toxin-expressing transgenic potato varieties declined dramatically by 2000 due to concern over consumer resistance to purchasing transgenic potatoes and products made from them. An additional factor contributing to this decline was competition from new, conventional insecticides that controlled a broader spectrum of potato pests. The future role of transgenic potato varieties in the Colorado potato beetle management is currently uncertain, despite their effectiveness and considerable evidence that they have no significant effect on populations of natural enemies (Hoy et al., 1998; Riddick et al., 1998). Transgenic lines of aubergine expressing the Cry3A endotoxin (Hamilton et al., 1997b) and lines expressing the Cry3B endotoxin (Arpaia et al., 1997; Mennella et al., 1998), although not commercially available, have been shown to provide control of Colorado potato beetle.

Because of the potential that Colorado potato beetle will develop resistance to the Cry3A endotoxin if these transgenic potato varieties are widely planted, significant effort has been directed towards the development of resistance management strategies for transgenic potatoes. Proposed resistance management strategies for potato focus on using varieties that express a high dose of the toxin in conjunction with plantings of refuge areas of non-transgenic potato or other hosts in which beetles are not controlled. The level of toxin expressed should be high enough to kill any individuals that are heterozygous for resistance alleles. The size and location of refuge plantings must be such that any homozygous resistant beetles selected on the transgenic crop mate with homozygous susceptible beetles produced in the refuge. The progeny of such matings would be heterozygous for the resistance allele and would be killed if they fed on the transgenic crop. Strategies for managing resistance to transgenic potatoes are discussed by Gould et al. (1994) and Hoy (1999). Ochoa-Campuzano et al. (2013) identified prohibitin, an essential protein for L. decemlineata larval viability. They explored the possibility for prohiobitin-1 silencing in L. decemlineata larvae in order to reach higher efficacy of Cry3Aa toxin. Cooper et al. (2006) assessed the effectiveness of the protein avidin against the Colorado potato beetle neonates in a no-choice detached leaf bioassay at 0, 17, 34, 51, 102, and 204 μg avidin/ml over 12 d. The combined effects of avidin (136 μg avidin/ml) with Bt-Cry3A or leptines were evaluated with neonates and third instars over 12 and 6 days, respectively. Survival of third instars on the Bt-Cry3A with avidin was significantly reduced after 3 days compared with survival on the Bt-Cry3A, suggesting the addition of avidin may increase susceptibility to Bt-Cry3A.

The Colorado potato beetle spiroplasma (CPBS) appears to be host (genus)-specific and is transmitted among larvae and adults during regurgitation and defecation. However, researchers have adopted a strategy to engineer spiroplasma with an insect-lethal gene because the spiroplasma appears to be a commensal (Hackett et al., 1988; Gasparich et al., 1993a, b). The delta-endotoxin gene for B. thuringiensis subsp. tenebrionis (Btt) may be an appropriate gene for this purpose because the Colorado potato beetle is susceptible to the toxin and the CPBS adheres to the midgut microvilli (Hackett and Clark, 1989), which is the site of action of the delta-endotoxin. This system would allow multiplication and spread of the genetically-engineered microorganism throughout the crop in contrast to direct treatment of beetles with the Btt beta-endotoxin.

RNAi interference

Zhu et al. (2011) reported the results of study in which the potential of feeding dsRNA expressed in bacteria or synthesized in vitro to manage populations of L. decemlineata was investigated. Feeding RNA interference (RNAi) successfully triggered the silencing of all five target genes tested and caused significant mortality and reduced body weight gain in the treated beetles. This study provides the first example of an effective RNAi response in insects after feeding dsRNA produced in bacteria. The obtained results suggest that the efficient induction of RNAi using bacteria to deliver dsRNA is a possible method for management of L. decemlineata. This method is still in the experimental stage.

Host-Plant Resistance

The incorporation of varietal resistance to Colorado potato beetle has emphasised the transfer of resistance traits to S. tuberosum from other Solanum species using a variety of techniques to obtain successful interspecific crosses (Shapiro et al., 1991; Tingey and Yencho, 1994). Emphasis has been placed on resistance derived from Solanum berthaultii, which is mediated in large part by glandular trichomes on the foliage (Yencho et al., 1996) and on resistance derived from Solanum chacoense, which is mediated by high concentrations of leptine glycoalkaloids in the foliage (Sinden et al., 1986; Sanford et al., 1997; Yencho et al., 2000). Potato breeding lines with resistance to potato beetle have been released (Plaisted et al., 1992; Lorenzen and Balbyshev, 1997). Other potentially valuable mechanisms of resistance have been identified as well (for example, Balbyshev and Lorenzen, 1997).

Chemical Control

Chemical insecticides have constituted the primary method of control for Colorado potato beetle for much of its history as a pest of potato. However, the extraordinary ability of the beetle to develop resistance to virtually all insecticides used to control it has led repeated control failures in many areas (Casagrande, 1987; Georghiou and Lagunes-Tejeda, 1991; Bishop and Grafius, 1996). The commercialisation of several new insecticides with novel modes of action, which are effective in controlling Colorado potato beetle populations resistant to organophosphate, carbamate, chlorinated hydrocarbon and pyrethroid insecticides has restored the ability to control the beetle with insecticides in areas most severely affected by resistance. In the past 60 years L. decemlineata has developed resistance to 54 different insecticides, including imidacloprid and eight other neonicotinoids (Whalon et al., 2013). Up to 155-fold resistance to imidacloprod was reported in adult L. decemlineata from selected fields on Long Island, New York state, USA, after three seasons of use (Zhao et al., 2000), followed by thiamethoxam and clothianidin in other regions of the USA (Mota-Sanchez et al., 2006; Alyokhin et al., 2007; 2008).

Because of the importance of insecticides in managing Colorado potato beetle in commercial potato production, great emphasis has been placed on the development of procedures for managing resistance. Key processes involved in adaptation of Colorado potato beetle populations to insecticides have been explored using simulation models (Follett et al., 1993, 1995).

Resistance management efforts for Colorado potato beetle focus primarily on using pest management procedures, including crop rotation, scouting and action thresholds to minimize the use of insecticides in conjunction with resistance monitoring and rotations of insecticides having different modes of action (Kennedy and French, 1994; Grafius, 1997; Midgarden et al., 1997; Dively et al., 1998). A computerized approach to insecticide management for the Colorado potato beetle has been used in commercial potato fields on the eastern shore of Virginia, USA (Vencill et al., 1995). The Potato-Insect Expert System (PIES) uses insect life stages, potato growth stage, percentage defoliation and other factors to determine whether the application of insecticide is necessary to prevent tuber yield loss. Recommendations from PIES were compared with commercial spray thresholds based on the number of L. decemlineata per stem. Tuber yields were not significantly different between the two methods, although on average PIES recommended fewer insecticide applications than the conventional method. Visual defoliation ratings for PIES also required less time than conventional sampling for L. decemlineata on potato stems.

In order to establish an integrated pest management system, a temperature-driven decision support system (SIMLEP DSS) was designed for Europe, consisting of two modules (Jorg et al., 2007). SIMLEP 1 is a regional forecasting model for the first occurrence of hibernating beetles and the start of egg laying. SIMLEP 3 is a field-specific model which forecasts the occurrence of the developmental stages of L. decemlineata. From 1999 to 2004 SIMLEP 3 was validated in Germany, Austria, Italy and Poland. In about 90% of cases SIMLEP 3 correctly predicted the periods of maximum egg laying and young larval occurrence, which are the optimal periods for field assessments and treatments with conventional and biological insecticides. SIMLEP 3 has since been validated in Slovenia (Kos et al., 2009). SIMLEP 3 is used in practice in Germany and Austria on a large scale and in the western part of Poland.

Integrated Pest Management

IPM programs for potato emphasize reducing insect pest populations by using a number of measures against the pest, including crop rotation, altered planting dates (to avoid peak pest populations), and use of B. thuringiensis toxins and other pesticides only when necessary to prevent damaging populations from developing. IPM systems seek to maximise the impact of naturally occurring biological control in suppressing potato beetle populations (Hazzard et al., 1991; Hilbeck et al., 1997, 1998). One such management system is described by Ferro (2000).

The suggestion of adding sub lethal dosages of chemical insecticides to biological insecticides in order to improve their efficacy was raised by Kovacevic (1960) and was later discussed by Benz (1971). Some investigations showed that the combinations of the subnormal dosages of some insecticides could result in the improved efficacy of B. thuringiensis insecticides and independent synergism against L. decemlineata (Dobrincic 1996; Dobrincic and Igrc Barcic, 1998). If the combinations of insecticides at lower dosages are used, various benefits could be expected (Igrc Barcic et al., 2006): (1) ecological, for lower dosage means less pollution; (2) biological, the combinations could slow up resistance development, and the joint action of the insecticides used could result in a synergistic effect; and (3) economic, as there would be lower costs per treatment. In field trials one application of a combination of reduced spinosad and B.t.t. doses, plus pyrethrin, resulted in good protection from L. decemlineata larval attack, which resulted in satisfactory protection of the leaves and in significant yield increase of potato yield. Adding the low dose of spinosad to B.t.t. insecticide in laboratory trials resulted in a low increase in efficacy of B.t.t., and adding a low dose of B.t.t. to spinosad did not result in any change of the efficacy of spinosad (Bažok et al., 2008). No synergistic effect between the insecticides was noted; the joint actions of combinations were mainly described as independent synergism. Even with the independent synergism the combinations could be recommended for use if they had some advantages over other possibilities for the control of L. decemlineata.

Considerable effort has been directed toward enhancing naturally occurring biological control through augmentative releases of predators and parasitoids. In research plots releases of nymphs of Perillus bioculatus, either alone or in combination with weekly or less frequent applications of B. thuringiensis subsp. tenebrionis, were investigated in Delaware, USA (Hough-Goldstein and Whalen, 1993). In Quebec, Canada, inundative releases of P. bioculatus showed potential for suppressing Colorado potato beetle populations when used within an IPM programme (Cloutier and Bauduin, 1995).

Releases of the parasitoid Edovum puttleri were a critical component in the management of insecticide resistant potato beetles in aubergine in New Jersey, USA. However, the programme was expensive and has been largely discontinued following the introduction of new insecticides highly effective against potato beetle populations resistant to traditional insecticides.

Phytosanitary Measures

Countries at risk should require that consignments of any plants or plant products be found free from L. decemlineata after having been subjected to sorting and packaging techniques in suitable premises. In addition, potatoes and certain vegetable crops should be grown in fields which have been inspected during the growing season and found to be free from the pest, and/or in areas where either the pest does not occur or is under intensive official control.

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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.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

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