Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Popillia japonica
(Japanese beetle)

Klein M, 2008. Popillia japonica (Japanese beetle). Invasive Species Compendium. Wallingford, UK: CABI. DOI:10.1079/ISC.43599.20203373917



Popillia japonica (Japanese beetle)


  • Last modified
  • 11 December 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Popillia japonica
  • Preferred Common Name
  • Japanese beetle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • The Japanese beetle was first discovered in New Jersey, USA in 1916. It probably entered the USA as grubs with iris bulbs before 1912 when plant materials were first examined. Although not a pest in Japan, extensive, well-watered, turf, and a lack...

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Popillia japonica (Japanese beetle); Adult beetle. July 2016.
CaptionPopillia japonica (Japanese beetle); Adult beetle. July 2016.
Copyright©Theresa Cira, Minnesota Department of Agriculture
Popillia japonica (Japanese beetle); Adult beetle. July 2016.
AdultPopillia japonica (Japanese beetle); Adult beetle. July 2016.©Theresa Cira, Minnesota Department of Agriculture
Popillia japonica (Japanese beetle); Sexual dimorphism of Japanese beetle, male with tibial spurs (a) and female (b) can be differentiated by external leg features.
TitleSexual dimorphism between adults
CaptionPopillia japonica (Japanese beetle); Sexual dimorphism of Japanese beetle, male with tibial spurs (a) and female (b) can be differentiated by external leg features.
Copyright©Tom Hillyer
Popillia japonica (Japanese beetle); Sexual dimorphism of Japanese beetle, male with tibial spurs (a) and female (b) can be differentiated by external leg features.
Sexual dimorphism between adultsPopillia japonica (Japanese beetle); Sexual dimorphism of Japanese beetle, male with tibial spurs (a) and female (b) can be differentiated by external leg features.©Tom Hillyer
Adult P. japonica ('Japanese beetle') on flowerhead.
TitleAdult beetle
CaptionAdult P. japonica ('Japanese beetle') on flowerhead.
CopyrightAgriculture & Agri-Food Canada/Canadian Food Inspection Agency
Adult P. japonica ('Japanese beetle') on flowerhead.
Adult beetleAdult P. japonica ('Japanese beetle') on flowerhead.Agriculture & Agri-Food Canada/Canadian Food Inspection Agency
Popillia japonica (Japanese beetle); Grub raster, White grubs can be identified by the raster hair arrangement. August 2012.
CaptionPopillia japonica (Japanese beetle); Grub raster, White grubs can be identified by the raster hair arrangement. August 2012.
Copyright©Adam Sisson, Iowa State University
Popillia japonica (Japanese beetle); Grub raster, White grubs can be identified by the raster hair arrangement. August 2012.
LarvaPopillia japonica (Japanese beetle); Grub raster, White grubs can be identified by the raster hair arrangement. August 2012.©Adam Sisson, Iowa State University
Popillia japonica (Japanese beetle); Larva, third instar. May 2015.
CaptionPopillia japonica (Japanese beetle); Larva, third instar. May 2015.
Copyright©Erin Hodgson, Iowa State University
Popillia japonica (Japanese beetle); Larva, third instar. May 2015.
LarvaPopillia japonica (Japanese beetle); Larva, third instar. May 2015.©Erin Hodgson, Iowa State University


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Preferred Scientific Name

  • Popillia japonica Newman

Preferred Common Name

  • Japanese beetle

International Common Names

  • English: beetle, Japanese
  • Spanish: escarabajo japonés
  • French: hanneton japonais; scarabé japonais

Local Common Names

  • Denmark: Japanbille
  • Germany: Japankäfer; Kaefer, Japan-
  • Italy: scarabeo giapponese
  • Japan: mame-kogane
  • Sweden: Japanbagge

EPPO code

  • POPIJA (Popillia japonica)

Summary of Invasiveness

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The Japanese beetle was first discovered in New Jersey, USA in 1916. It probably entered the USA as grubs with iris bulbs before 1912 when plant materials were first examined. Although not a pest in Japan, extensive, well-watered, turf, and a lack of parasites, allowed populations to rapidly build up and spread steadily west to the Mississippi River. The loss of the chlorinated hydrocarbon insecticides, and the end of the Federal quarantine on nursery stock, has allowed beetles to move into western states at a rapid rate. Beetles are pests of quarantine concern in the western USA and Europe. P. japonica was found on Terceira Island, Azores, Portugal in the 1980s. Again, extensive turf allowed establishment of beetles, population explosions, the infestation of that island, and subsequently of three more of the Azorean Islands. Beetles have moved considerably outside of the climatic conditions in their native Japan.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Scarabaeidae
  •                             Genus: Popillia
  •                                 Species: Popillia japonica

Notes on Taxonomy and Nomenclature

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The Japanese beetle, Popillia japonica, is a member of the order Coleoptera, family Scarabaeidae, subfamily Rutelinae and tribe Anomlini.


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Newly-laid eggs are about 1.5 mm long, pearly white and oblong. Eggs absorb water from the soil, becoming spherical and nearly doubling in size within a week. The external surface of the protection chorion is marked with small hexagonal areas. The developing embryo can be seen within the eggs that are close to hatching (Fleming, 1972a).


P. japonica larvae are typical scarabaeid grubs (Fleming, 1972a). The head is yellowish-brown, with strong, dark-coloured mandibles. The body consists of three thoracic segments, each with a pair of jointed legs, and a 10-segmented abdomen. The grubs assume a typical, scarab, C-shaped position in the soil. The cuticle is transversely wrinkled and is covered with scattered brown hairs, which are interspersed with short, blunt, brown spines. The raster, located on the ventral side of the last abdominal segment, has many scattered, brown, hooked spines; medially, two conspicuous rows of 6-7 shorter straight spines are arranged in the form of a truncated V. This V-shaped arrangement on the raster distinguishes P. japonica from other larval scarabaeids in the USA. The last abdominal segment also bears many yellowish hairs at the sides and the end.

Newly-hatched grubs are about 1.5 mm long and translucent white. The abdominal area becomes dark once the larva has fed and the rectal sacs, or fermentation chambers, fill with soil. There are three instars. Just prior to moulting, first and second instars attain average middorsal lengths of 10.5 and 18.5 mm, respectively, whereas the mature, third instar, averages 32 mm. Head capsules of first, second and third instars average 1.2 mm wide and 0.7 mm long, 1.9 mm wide and 1.2 mm long, and 3.1 mm wide and 2.1 mm long, respectively.


When mature, the grub stops feeding, voids the gut so that the rectal sacs lose their dark appearance, and become cream-coloured and a pale, somewhat shrunken prepupa. The body straightens out, except for a slight crook at the caudal end. Eventually, the developing appendages are everted from their sacs and lie outside the newly developed pupal cuticula, beneath the old larval cuticula. The transformation to prepupa and pupa, both of which are very delicate, occurs in an earthen cell formed by the mature larva (Fleming, 1972a).


The young pupa forms within the old larval and prepupal exuviae, which changes in appearance to a fine, light tan, mesh-like tissue. This shroud-like covering splits along the middorsal line as the pupa develops. The pupa, which averages 14 mm long and 7 mm wide, resembles the adult beetle, except the wings and other appendages are closely folded to the body. It is a pale-cream colour at first, gradually becoming tan and finally taking on the metallic green of the adult (Fleming, 1972a).


Fleming (1972a) published an illustrated description of the beetle, including characters on the fore tibia and tarsi by which the sexes can be distinguished (the male tibial spur is hook-like, not spatulate, and the tarsi attach at the end of the tibia, not upto a few mm). The adult is an attractive, broadly oval beetle, 8-11 mm long, and about 5-7 mm wide. Females are usually slightly bigger than males. The head and body are dark, metallic green, with darker copper-green legs. The coppery-brown elytra, which do not quite reach the tip of the abdomen, expose a row of five lateral patches of white hairs on each side of the abdomen, and a pair of these patches on the dorsal surface of the last abdominal segment. These white patches on the green abdomen distinguish P. japonica from all other beetles that resemble it in the USA. However, there are several similar Popillia species in the far-east that require an expert to distinguish them from P. japonica (see Ping, 1988).


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P. japonica originates from north-eastern Asia where it is native in northern Japan and in the far east of Russia (Fleming, 1972a). Fleming’s (1972a) report of P. japonica in China and Korea, probably referred to closely-related species, but not the Japanese beetle (Ping, 1988; Reed et al., 1990).

In Japan, the beetle is most abundant in northern Honshu and all of Hokkaido where grasslands occur, but it does not reach the high population densities that occur in the USA. It is common, but not abundant in Kyushu, Shikoku and southern Honshu. It was not considered to be a pest in Japan until the increase in golf courses, and is still not the major scarab pest species.

In the USA, P. japonica, is established in all states boarding, or east of the Mississippi River, with the possible exception of Florida. Several western states have isolated established populations of the beetle. Pest survey data are submitted to NAPIS by participating USA states in the Cooperative Agricultural Pest Survey (CAPS) with USDA, APHIS, PPQ and the resulting distribution of P. japonica in the USA is mapped from 2001 to the present (2010), and can be found here: See also Smith et al. (1997).

In Russia, the last report of P. japonica restricted it to the South Kirile region of Sakhalin, on the island of Kunashir (Chebanov, 1977). It is absent in the EPPO region except for Terceira Island, Azores (Portugal), where the pest probably spread from the USA air base (Simoes, 1984). An infestation on the island of Faial was found in 1996. More recently Pico and San Miguel both become infested in 2006. Both Terceira and Faial are generally infested, but eradication programmes are being considered for the isolated infestations on Pico and San Miguel.

Distribution Table

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

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


ChinaAbsent, Invalid presence record(s)
-HeilongjiangAbsent, Invalid presence record(s)
-HunanAbsent, Invalid presence record(s)
-JilinAbsent, Invalid presence record(s)
Hong KongAbsent, Invalid presence record(s)
IndiaAbsent, Formerly present
JapanPresent, Widespread
-HokkaidoPresentWhole island
-KyushuPresentLight infestation
North KoreaAbsent, Unconfirmed presence record(s)
South KoreaAbsent, Invalid presence record(s)
TaiwanAbsent, Formerly present


ItalyPresent, Few occurrences
LithuaniaAbsent, Confirmed absent by survey
NetherlandsAbsent, Intercepted only
PortugalPresent, Localized
-AzoresPresent, LocalizedOn four of nine islands
RussiaPresent, Localized
-Russian Far EastPresent, Localized
SloveniaAbsent, Confirmed absent by survey
SwitzerlandPresent, Transient under eradication

North America

CanadaPresent, Localized
-British ColumbiaPresent, Few occurrences
-New BrunswickPresent, Localized
-Nova ScotiaPresent, Localized
-Prince Edward IslandPresent, Localized
United StatesPresent, Localized1916
-AlabamaPresent, Few occurrences
-CaliforniaAbsent, Eradicated
-ColoradoPresent, Few occurrences
-ConnecticutPresent, Localized
-DelawarePresent, Localized
-District of ColumbiaPresent, Localized
-GeorgiaPresent, Localized
-IdahoAbsent, Formerly present
-IllinoisPresent, Localized
-IndianaPresent, Localized
-IowaPresent, Localized
-KansasPresent, Few occurrences
-KentuckyPresent, Localized
-MainePresent, Localized
-MarylandPresent, Localized
-MassachusettsPresent, Localized
-MichiganPresent, Localized
-MinnesotaPresent, Localized
-MissouriPresent, Localized
-NebraskaPresent, Few occurrences
-NevadaAbsent, Eradicated
-New HampshirePresent, Localized
-New JerseyPresent, Localized
-New MexicoPresent
-New YorkPresent, Localized
-North CarolinaPresent, Localized
-OhioPresent, Localized
-OklahomaPresent, Few occurrences
-OregonAbsent, Eradicated
-PennsylvaniaPresent, Localized
-Rhode IslandPresent, Localized
-South CarolinaPresent, Localized
-South DakotaPresent, Few occurrences
-TennesseePresent, Localized
-VermontPresent, Localized
-VirginiaPresent, Localized
-West VirginiaPresent, Localized
-WisconsinPresent, Localized

Risk of Introduction

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P. japonica is an A1 quarantine organism for EPPO (OEPP/EPPO, 1980), and is also of quarantine significance for CPPC, JUNAC, NAPPO, and OIRSA. Within the USA, P. japonica is the object of an USDA/APHIS quarantine, restricting interstate movement of aircraft from regulated airports (USDA-APHIS, 2004). The interstate shipment of plant material is covered by the US Domestic Japanese Beetle Harmonization Plan (National Plant Board, 2004). Very few of the P. japonica infestations in states west of the Mississippi River have been associated with movement of beetles by aircraft.

Temperature and particularly soil moisture are the main factors limiting potential spread of the beetle into new areas. According to Fleming (1972a), the Japanese beetle is adapted to regions were the mean soil temperature is between 17.5 and 27.5oC during the summer, and above -9.4oC in the winter. It should be noted that the winter temperature is 0.5-1.0 m below ground surface where the larvae overwinter. In addition, precipitation must be adequate and rather uniformly distributed throughout the year, averaging at least 25 cm during the summer. However, these parameters were established before irrigation was prevalent in much of the midwest and western USA. The Japanese beetle distribution in the USA is far south of the beetle distribution in its native Japan. The distribution in Japan may be influenced by other species of Popillia, or other scarabs, competing for limited resources.

Allsopp (1996) used a computer-generated modified Match Index to analyze climatic suitability and predict the potential worldwide distribution of P. japonica. According to the model, in North America the beetle has the potential to spread west to the middle of Nebraska, Kansas, Oklahoma, and Texas, south to the middle of South Carolina and Georgia, and most of Alabama and Mississippi. The Japanese beetle has already met or exceeded all of these parameters in the past 12 years. The southern parts of the Canadian Maritimes and eastern British Columbia, and parts of Washington and Oregon are also suitable. Indeed, Oregon has had several P. japonica infestations, as has California, and now Colorado and Utah. In addition Ontario, Quebec and Nova Scotia all have infestations in eastern Canada. Most of continental Europe, except most of Scandinavia and the Mediterranean areas, are suitable, as are the UK and Ireland. Suitable areas in Asia include the Caucasus and areas to the north, eastern central China, and the Korean Peninsula. In Africa, the mountains of Morocco and coastal south-eastern South Africa seem suitable. In the Southern Hemisphere, the south-eastern highlands of Australia, Tasmania, New Zealand, and the Rio de la Plata area between Argentina and Uruguay, and parts of costal Chile are suitable. Once established in these areas, P. japonica could cause enormous damage and significant economic loss. It is interesting to note that unlike the weevils, no scarab has established on the opposite side of the equator from its native land (Jackson and Klein, 2006).

Habitat List

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Hosts/Species Affected

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In the USA, adult P. japonica have been observed feeding on at least 295 species of plants in 79 plant families (Fleming, 1972a). These include small fruits, tree fruits, vegetable and garden crops, field crops, woody and herbaceous ornamentals, shade trees, various weeds, and many non-economic species. Economic damage has been recorded on more than 100 species. The beetles are particularly attracted to certain species of Aceraceae, Anacardiaceae, Betulaceae, Clethraceae, Ericaceae, Fagaceae, Gramineae, Hippocastanaceae, Juglandaceae, Lauraceae, Leguminosae, Liliaceae, Lythraceae, Malvaceae, Onagraceae, Platanaceae, Polygonaceae, Rosaceae, Salicaceae, Tiliaceae, Ulmaceae and Vitaceae. The grubs feed on roots of a wide range of vegetable crops, ornamental plants and tender grasses. In Japan, the host range appears to be smaller than in North America.

Within the EPPO region, the host range of P. japonica would be similar. Malus, Prunus, Rubus and Vitis, with their wide distribution and intensive cultivation, would be especially favourable food sources for the adults. Lush pasture and turfgrasses provide a favourable habitat for the grubs.

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Acer (maples)AceraceaeMain
    Acer campestre (field maple)AceraceaeUnknown
    Acer palmatum (Japanese maple)AceraceaeUnknown
    Acer platanoides (Norway maple)AceraceaeUnknown
    Aesculus (buckeye)HippocastanaceaeOther
      Alnus japonica (Japanese alder)BetulaceaeUnknown
      Althaea (hollyhocks)MalvaceaeOther
        Asparagus officinalis (asparagus)LiliaceaeMain
        Berchemia racemosaUnknown
        Betula (birches)BetulaceaeOther
          Betula nigra (river birch)BetulaceaeUnknown
          Betula utilis (Himalayan silver birch)BetulaceaeUnknown
          Castanea (chestnuts)FagaceaeOther
          Castanea crenata (Japanese chestnut)FagaceaeUnknown
          Cayratia japonica (Sorrel vine)VitaceaeUnknown
          Cercis canadensis (eastern redbud)FabaceaeUnknown
          Cotinus coggygria (fustet)AnacardiaceaeUnknown
          Cynodon dactylon (Bermuda grass)PoaceaeUnknown
          Dioscorea japonica (Japanese yam)DioscoreaceaeUnknown
          Fagus sylvatica (common beech)FagaceaeUnknown
          Fallopia convolvulus (black bindweed)PolygonaceaeUnknown
          Fallopia japonica (Japanese knotweed)PolygonaceaeUnknown
          Festuca arundinacea (tall fescue)PoaceaeUnknown
          Glycine max (soyabean)FabaceaeMain
            Glycine sojaFabaceaeUnknown
            Hibiscus (rosemallows)MalvaceaeOther
              Hibiscus syriacus (shrubby althaea)MalvaceaeUnknown
              Hypericum (st Johnsworts)ClusiaceaeUnknown
              Juglans nigra (black walnut)JuglandaceaeOther
                Lagerstroemia indica (Indian crape myrtle)LythraceaeWild host
                  Lolium perenne (perennial ryegrass)PoaceaeUnknown
                  Malus (ornamental species apple)RosaceaeMain
                  Malus floribundaRosaceaeUnknown
                  Malus toringo (toringo crab-apple)RosaceaeUnknown
                  Malus zumiRosaceaeUnknown
                  Melia azedarach (Chinaberry)MeliaceaeUnknown
                  Oenothera biennis (common evening primrose)OnagraceaeUnknown
                  Parthenocissus quinquefolia (Virginia creeper)VitaceaeWild host
                    Phaseolus vulgaris (common bean)FabaceaeUnknown
                    Platanus (planes)PlatanaceaeOther
                      Platanus orientalis (plane)PlatanaceaeUnknown
                      Poa pratensis (smooth meadow-grass)PoaceaeUnknown
                      Polygonum (knotweed)PolygonaceaeWild host
                        Polygonum lapathifolium (pale persicaria)PolygonaceaeUnknown
                        Polygonum thunbergiiUnknown
                        Populus (poplars)SalicaceaeOther
                          Populus maximowiczii (Japanese poplar)SalicaceaeUnknown
                          Populus nigra (black poplar)SalicaceaeUnknown
                          Prunus (stone fruit)RosaceaeMain
                            Prunus cerasifera (myrobalan plum)RosaceaeUnknown
                            Prunus cistenaRosaceaeUnknown
                            Prunus domestica (plum)RosaceaeOther
                              Prunus japonica (Japanese bush cherry tree)RosaceaeUnknown
                              Prunus sargentii (sargent's cherry)RosaceaeUnknown
                              Prunus serrulata (Japanese flowering cherry)RosaceaeUnknown
                              Prunus subhirtella (weeping Japanese cherry)RosaceaeUnknown
                              Prunus virginiana (common chokecherrytree)RosaceaeUnknown
                              Prunus yedoensisRosaceaeUnknown
                              Pteridium aquilinum (bracken)DennstaedtiaceaeUnknown
                              Quercus serrata (glandbearing oak)FagaceaeUnknown
                              Quercus variabilis (oriental cork oak)FagaceaeUnknown
                              Rheum hybridum (rhubarb)PolygonaceaeMain
                                Rosa (roses)RosaceaeMain
                                Rosa multiflora (multiflora rose)RosaceaeUnknown
                                Rubus (blackberry, raspberry)RosaceaeMain
                                  Rubus crataegifoliusRosaceaeUnknown
                                  Rumex (Dock)PolygonaceaeUnknown
                                  Salix (willows)SalicaceaeOther
                                    Salix purpurea (purple willow)SalicaceaeUnknown
                                    Sassafras albidum (common sassafras)LauraceaeOther
                                      Smilax china (Chinaroot)SmilacaceaeUnknown
                                      Sorbus americana (American mountainash)RosaceaeOther
                                        Tilia (limes)TiliaceaeMain
                                          Tilia americana (basswood)TiliaceaeUnknown
                                          Tilia cordata (small-leaf lime)TiliaceaeUnknown
                                          Tilia euchlora (crimean lime)TiliaceaeUnknown
                                          Tilia japonicaTiliaceaeUnknown
                                          Tilia miquelianaTiliaceaeUnknown
                                          Tilia tomentosa (silver lime)TiliaceaeUnknown
                                          Trifolium pratense (red clover)FabaceaeUnknown
                                            Ulmus (elms)UlmaceaeMain
                                              Ulmus parvifolia (lacebark elm)UlmaceaeUnknown
                                              Vaccinium corymbosum (blueberry)EricaceaeUnknown
                                              Vitis (grape)VitaceaeMain
                                              Vitis riparia (riverbank grape (USA))VitaceaeUnknown
                                              Vitis thunbergiiUnknown
                                              Vitis vinifera (grapevine)VitaceaeUnknown
                                              Wisteria floribunda (Japanese wisteria)FabaceaeUnknown
                                              Zea mays (maize)PoaceaeMain
                                                Zelkova serrata (Japanese selkova)UlmaceaeUnknown

                                                Growth Stages

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                                                Flowering stage, Fruiting stage, Pre-emergence, Seedling stage, Vegetative growing stage


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                                                Skeletonized foliage is the most common symptom of feeding by the adult. The beetles generally feed from the upper surface of leaves, chewing out the tissue between the veins and leaving a lacelike skeleton. Severely damaged leaves soon turn brown and drop. The adults are gregarious, usually beginning to feed on foliage at the top of a plant and working downward. On some plants with thin leaves and fine venation, and on petals of flowers, the beetles consume irregularly-shaped sections in the same manner as many Lepidoptera. Plants with thick, tough leaves are usually not attacked, but when such leaves are eaten (Concord grapes), the feeding is often restricted to the palisade mesophyll and does not penetrate to the lower leaf surface.

                                                Maize is one field crop seriously damaged in North America. The beetles feed on the maturing silk, preventing pollination; this results in malformed kernels and reduced yield. This appears to be more of a back-yard maize growing situation because the light-loving beetles rarely venture more than 1-2 rows into a maize field. However, beetles can feed over an entire soyabean field and cause their damage. They also defoliate asparagus, nearly all varieties of grapes, and many fruit-bearing trees, especially apple, cherry, plum, and peach. Beetles can aggregate and feed in large numbers on the fruit of early-ripening varieties of apple, peach, nectarine, plum, raspberries, and quince. This feeding renders fruit unmarketable, unless they have been protected by pesticides.

                                                The larvae are most abundant in well-kept lawns and golf courses, and less often in pastures. As the grub feeds just below the surface, it cuts off and consumes the grass roots. Early symptoms include thinning, yellowing, and wilting, culminating in large patches of dead, brown grass that appears in late summer or early autumn because of water stress, and less often in the following spring when more moisture is normally available. When grubs are numerous (400/m2+) the root system is completely severed and the sod can be lifted or rolled back like a carpet. Secondary damage from skunks, raccoons, crows, or other predators often causes more disruption to the sward than the grubs themselves. Feeding by grubs on roots of maize, beans, tomatoes, strawberries, nursery seedlings, or other crops reduces their vitality and yield and sometimes kills the plants. Damage is often most severe when these crops are planted into areas which were previously turf.

                                                List of Symptoms/Signs

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                                                SignLife StagesType
                                                Fruit / abnormal shape
                                                Fruit / external feeding
                                                Inflorescence / external feeding
                                                Leaves / external feeding
                                                Roots / external feeding
                                                Roots / reduced root system
                                                Whole plant / dwarfing
                                                Whole plant / external feeding
                                                Whole plant / plant dead; dieback

                                                Biology and Ecology

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                                                Recent general reviews of P. japonica should be consulted for additional information on this pest (Potter, 1998; Vittum et al., 1999; Potter and Held, 2002; Jackson and Klein, 2006). Fleming (1972a) provided a detailed account of the biology of P. japonica. Larvae (mainly third-instar) overwinter in an earthen cell about 15-20 cm deep in the soil. In early spring, when the soil temperatures increase to about 10oC, the grubs move closer to the surface and resume feeding on plant roots at 2.5 to 5.0 cm depth. Pupation usually occurs after 4-6 weeks of feeding, and the adults emerge from mid-May to mid-July, depending on latitude.

                                                Mating begins at emergence, and egg-laying soon follows. Virgin females produce a volatile sex pheromone (Ladd, 1970), identified and called Japonilure (Tumlinson et al., 1977). Early in the seasonal flight period, aggregations containing up to many dozens of males form on the ground around a single, emerging female. Females also remate on food plants between bouts of oviposition, with the last male’s sperm being used. The beetles normally feed in groups, usually starting near the top of a plant and working downward (Fleming, 1972a; Rowe and Potter, 1996). The adults are attracted to feeding-induced plant volatiles, resulting in aggregation on damaged plants (Fleming, 1972a; Loughrin et al., 1996). Females may leave host plants during the day and fly to suitable sites for oviposition. Areas with moist, loamy soil covered with turf or pasture grasses are preferred (Fleming, 1972a; Allsopp et al., 1992). Eggs are laid singly or in small clusters (2-4 eggs) in the upper 7.5 cm of soil. The cycle of feeding, mating and oviposition is repeated every few days. The average lifespan of a female is 30-45 days, during which she may lay 40-60 eggs.

                                                Eggs hatch in about 2 weeks, and the young larvae begin feeding on fine roots and organic matter. They moult and become second-instars after 2-3 weeks, and third-instars after 3-4 weeks more. Feeding continues until late autumn, when the grubs move deeper into the soil in response to declining soil temperatures. At the latitude of Virginia and Maryland, the populations consist of mainly adults and eggs in July, first- and second-instars by mid-August, second- and third-instars by early September, and third-instars from late September to late April, and prepupae and pupae in May and early June. Normally, there is one generation per year, even in the most southern areas, but at the northern edge of its range, a few individuals may need 2 years to complete the life cycle (Fleming, 1972a; Vittum, 1986; Vittum et al., 1999).

                                                Natural enemies

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                                                Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                Adapsilia flaviseta USA ornamental plants
                                                Bacillus popilliae lentimorbus Pathogen
                                                Bacillus thuringiensis galleriae Pathogen
                                                Bacillus thuringiensis thuringiensis Pathogen Larvae USA, Japan turf
                                                Beauveria bassiana Pathogen Adults/Larvae USA ornamentals, turf
                                                Campsomeris annulata Parasite Larvae Fleming (1968)
                                                Campsomeris marginella modesta Parasite Larvae Fleming (1968)
                                                Carabidae Predator Eggs/Larvae USA turf
                                                Craspedonotus tibialis Predator Larvae Fleming (1968)
                                                Dexia ventralis Parasite Larvae Fleming (1968)
                                                Entoderma colletosporium Pathogen Larvae Fleming (1968)
                                                Erythrocera genalis Parasite Adults Fleming (1968)
                                                Eutrixopsis javana Parasite Adults Fleming (1968)
                                                Formicidae Eggs/Larvae USA turf
                                                Hamaxia incongrua Parasite Adults Fleming (1968)
                                                Heterorhabditis bacteriophora Parasite Larvae Azores, USA turf
                                                Heterorhabditis heliothidis Parasite
                                                Heterorhabditis marelatus Parasite Larvae USA turf
                                                Heterorhabditis megidis Parasite Larvae USA turf
                                                Istocheta aldrichi Parasite Adults USA ornamental plants, turf
                                                Istocheta ussuriensis Parasite Adults Fleming (1968)
                                                Metarhizium anisopliae Pathogen Adults/Larvae Azores, USA ornamentals, turf
                                                Nomuraea rileyi Pathogen
                                                Ovavesicula popilliae Pathogen Larvae Connecticut, Michigan, USA turf
                                                Paenibacillus popilliae Pathogen Larvae USA turf
                                                Pexopsis clauseni Parasite
                                                Prosena siberita Parasite Larvae New Jersey, USA turf
                                                Rickettsiella popilliae Pathogen Larvae Fleming (1968)
                                                Scolia japonica Parasite Larvae Fleming (1968)
                                                Serratia Pathogen Larvae Fleming (1968)
                                                Steinernema carpocapsae Parasite Larvae USA turf
                                                Steinernema feltiae Parasite Larvae Fleming (1968)
                                                Steinernema glaseri Parasite Larvae Azores, New Jersey, USA turf
                                                Steinernema kushidai Parasite Larvae Japan turf
                                                Steinernema riobravis Parasite Larvae Fleming (1968)
                                                Steinernema scapterisci Parasite Larvae Fleming (1968)
                                                Steinernema scarabaei Parasite Larvae New Jersey, USA turf
                                                Tiphia asericae Parasite Larvae Fleming (1968)
                                                Tiphia biseculata Parasite Larvae Fleming (1968)
                                                Tiphia brevilineata Parasite Larvae Fleming (1968)
                                                Tiphia burrelli Parasite USA ornamental plants
                                                Tiphia communis Parasite Larvae Fleming (1968)
                                                Tiphia koreana Parasite Larvae Fleming (1968)
                                                Tiphia matura Parasite Larvae Fleming (1968)
                                                Tiphia notopolita Parasite Larvae Fleming (1968)
                                                Tiphia phyllophagae Parasite Larvae Fleming (1968)
                                                Tiphia popilliavora Parasite Larvae Virginia, USA turf
                                                Tiphia pullivora Parasite Larvae Fleming (1968)
                                                Tiphia tegitiplaga Parasite Larvae Fleming (1968)
                                                Tiphia vernalis Parasite Larvae eastern USA turf
                                                Trophops clauseni Parasite Adults Fleming (1968)

                                                Notes on Natural Enemies

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                                                Several indigenous, generalist, predators, especially ants and ground beetles, help to suppress Japanese beetle populations by feeding on eggs and grubs (Terry et al., 1993; Zenger and Gibb, 2001). Both adults and larvae are fed upon by various birds including starlings, crows, grackles, and gulls. Moles, skunks, raccoons and armadillos, feed on the grubs, but cause considerable damage to the turf or pastures. No native parasitic insect attacks P. japonica in North America. However, at least three species of imported parasitic insects have become established in eastern and midwest states (see 'Prevention and Control'). Grubs are susceptible to several naturally-occurring fungal pathogens including Metarrhizium anisopliae and Beauveria bassiana, entomopathogenic nematodes including Steinernema and Heterorhabditis species, bacterial pathogens such as Paneibacillus popilliae, the protozoan, Ovavesicula popilliae, and the rickettsia Rickettsiella popilliae. Unfortunately, humans are also susceptible to the rickettsia.

                                                For further information see reviews by Fleming (1968), Potter and Held (2002), and Jackson and Klein (2006), and the text section on 'Prevention and Control'. Natural enemy records from older literature are presented in the 'Natural Enemy' table, and referenced as Fleming (1968).


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                                                P. japonica is the single most destructive insect pest on golf courses, lawns and pastures, and on herbaceous and woody landscape plants in the eastern USA (Tashiro, 1987; Potter, 1998; Vittum et al., 1999). A decade ago it was estimated that more than $460 million is spent each year to control the grubs and adults, and about $156 million in renovating or replacing damaged turf or ornamental plants (USDA/APHIS, 2000). Damage to tree fruits, small fruits, maize, and soybeans is also significant. In addition, many millions of US dollars, and considerable quantities of pesticides, are also lost trying to limit the beetle’s spread by nursery stock and aeroplanes in North America. The Japanese beetle has never been a major pest in Japan, and has not caused extensive damage up to this point in the Azores. Costs connected with quarantine concerns are likely to increase greatly with the discovery of the beetle on San Miguel Island, USA.

                                                Risk and Impact Factors

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                                                • Proved invasive outside its native range
                                                • Highly adaptable to different environments
                                                • Capable of securing and ingesting a wide range of food
                                                • Highly mobile locally
                                                • Benefits from human association (i.e. it is a human commensal)
                                                • Gregarious
                                                Impact outcomes
                                                • Altered trophic level
                                                • Damaged ecosystem services
                                                • Ecosystem change/ habitat alteration
                                                • Host damage
                                                • Negatively impacts agriculture
                                                • Negatively impacts cultural/traditional practices
                                                • Negatively impacts livelihoods
                                                • Reduced amenity values
                                                • Reduced native biodiversity
                                                • Soil accretion
                                                • Transportation disruption
                                                Impact mechanisms
                                                • Herbivory/grazing/browsing
                                                Likelihood of entry/control
                                                • Highly likely to be transported internationally accidentally
                                                • Difficult/costly to control


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                                                A diagnostic protocol for P. japonica is given in OEPP/EPPO (2006).

                                                Detection and Inspection

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                                                Adult P. japonica are easily detected by inspecting the most vulnerable plants for aggregations of beetles on foliage, flowers, or fruits (not blueberries), or for skeletonized leaves during the beetles flight period in early- to mid-summer. Adults are most active on warm days between 10 am and 3 pm. Traps containing the three part food-type lure (phenethyl propionate + eugenol + geraniol) and the sex attractant (Japonilure) (Ladd et al., 1981) are widely used in the USA and Azores for monitoring and survey purposes, and to delineate infestations. Grubs can be detected by cutting sections of sod with a spade or golf cup cutter in late summer, autumn, or in early spring and examining the soil and roots to a depth of about 8 cm. For grubs in nursery trees, removal and examination of soil down to 30 cm may be required to get an accurate sample.

                                                Prevention and Control

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                                                Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.


                                                The destructive potential and economic importance of this pest have led to intensive studies of various means for control. Note that the adults and grubs cause very different types of damage, above ground and below ground. Because the adults can fly considerable distances, controlling one life stage will not necessarily preclude problems with the other.


                                                Host-plant resistance

                                                Fleming (1972a) provided a ranking of the extent of feeding by adult P. japonica on 435 plant species in 95 families. Within some generally susceptible genera such as Betula, Malus and Tilia, less susceptible cultivars have been found (Ranney and Walgenbach, 1992; Spicer et al. 1995; Potter et al., 1998). Use of resistant, or less susceptible species and cultivars when planning a landscape, or replacing damaged plant material, is a key to managing adults. Highly susceptible trees such as Sassafras, Prunus cerasifera, Ancer ploatanoides, and Tillia spp., or certain wild plants, such as species of Malva, Parthenocissus, Polygonum, and Vitis, will attract numerous beetles.

                                                Although tolerance varies, all species of cool-season turfgrasses are susceptible to the grubs (Potter et al., 1992). Infection of tall fescue, Festuca arundinacae or perennial ryegrass, Lolium perenne, with fungal endophytes (Neotyphodium spp.) does not provide resistance to this pest.

                                                Physical removal and exclusion

                                                Hand removal may provide some control for small plantings. Beetles on plants are sluggish in the morning, before 9 am, or when the temperature is <21oC, and can be killed by picking them, or shaking them, into a bucket of soapy water (Ladd and Klein, 1982). This is most effective when done before damage to the plants. High-value plants such as roses can be protected with fine netting or Reemay fabric around each blossom during the period of beetle activity.


                                                Although mass trapping has held isolated populations in check, and reduced the regulatory situation at some airports, it has not been effective in reducing established P. japonica infestations. However, Japanese beetle traps are an important tool in the identification and delimitation of new P. japonica infestations. California (Potter and Held, 2002) and Oregon monitor 10,000 and 5,000 traps per year, respectively, and have both eradicated isolated infestations in their states. Other western states utilize traps to a lesser extent. Small-scale trapping may aggravate defoliation damage in landscapes because the traps may attract more beetles than actually enter the traps (Gordon and Potter, 1985). Although beetles can fly up to 5 miles, they rarely do, and are not attracted to traps more than 50-100 m away (Lacey et al., 1994).

                                                Cultural control

                                                Female beetles seek out sites that are most optimal for egg laying and survival (Allsopp et al., 1992). Withholding irrigation during peak beetle flight can help to reduce subsequent grub populations in time of drought, and in naturally dry areas (Potter et al., 1996). In contrast, rainfall or irrigation in summer and early autumn, during early instar feeding, promotes tolerance and recovery of grub-damaged turfgrass. Vigorous, well-watered turf can withstand two to three times the normal threshold of grubs (ca. 100/m2) that would destroy a weak, wilted, or starved sward. Raising cutting height, and maintaining a balanced fertility regime to promote growth of roots also enhances tolerance of root-feeding by grubs (Crutchfield et al., 1995).

                                                Biological control

                                                During 1920-1933, the USDA imported about 49 species of parasites of P. japonica and related scarabs from the orient and Australia and released them into Japanese beetle-infested areas in the USA (Fleming, 1968). Only a few of these became established, the most widely distributed are Tiphia vernalis, a wasp that parasitizes overwintered grubs in the spring, and Istocheta aldrichi, a tachinid fly that parasitizes adults ( The spring Tiphia seems to be well-established throughout the beetle-inhabiting areas. Istocheta had been restricted to the New England states, but has recently been established in North Carolina, Michigan, Minnesota, and Missouri, USA (Jackson and Klein, 2006). A third established parasitoid, Tiphia popilliavora, the fall tiphia, has not been recovered since 1969, although isolated populations may still be present. These parasitoids provide some suppression, particularly I. aldrichi, in areas with restricted turf, but do not usually provide adequate beetle control.

                                                Spores of Paenibacillus (=Bacillus) popilliae, the primary causal agent of milky disease in P. japonica were widely distributed in colonization programmes around the middle of the last century in eastern USA (Fleming, 1968). Although milky disease is one of the primary natural biological agents reducing Japanese beetle populations, the value of augmenting this natural incidence with commercial spore powder has come under question (Redmond and Potter, 1995; Potter and Held, 2002; Jackson and Klein, 2006). Another bacterium, Bacillus thuringiensis – serovar japonensis, strain Buibui, has shown strong larvacidal activity against P. japonica and other grubs (Ohba et al., 1992; Alm et al., 1997), but lacks a commercial product in the USA.

                                                Entomopathogenic nematodes in the genera Steinernema and Heterorhabditis are the most commonly used pathogens against P. japonica. Nematodes such as Steinernema glaseri and Heterorhabditis bacteriophora are better-adapted to locate and parasitize the grubs in the soil (Gaugler et al., 1997). Wright et al. (1988) showed that nematodes could be used to control P. japonica grubs in container-grown nursery plants. Techniques for using these nematodes can be found at, or in the training video “Entomopathogenic Nematodes: Tools for pest management” (Gaugler and Klein, 1998). Autodissemination of the fungus Metarrhizium anisopliae has been used to suppress Japanese beetle populations in the Azores and the USA (Klein and Lacey, 1999; Vega et al., 2007).

                                                Chemical control

                                                Adult beetles have been controlled by treating susceptible plants with carbamates, organophosphates, or more recently, pyrethroid insecticides (Potter, 1998; Potter and Held, 2002). After the banning of long-residual insecticides grubs were treated with short-residual organophosphates and carbamates. More recently, neonicotinyl (imidacloprid) and moult accelerators (halofenozide) have been used for preventive larval control in turf (Potter and Held, 2002), and may have an effect for more than 1 year (George et al., 2007).

                                                Phytosanitary Measures

                                                Control of larvae in nursery stock is a far greater problem with quarantine concerns. Approved procedures are limited to chlorpyrifos dips of balled and burlapped nursery stock, and application of Marathon (imidacloprid) and Discus (imidacloprid + cyfouthrin) to field stock (Mannion et al., 2000; 2001; National Plant Board, 2004; Oliver et al., 2007). None of these materials provide 100% mortality of grubs, resulting in the three west-coast states (CA, OR, WA) prohibiting plants with any soil. Adult beetles are usually eliminated from fresh produce by commercial grading. However, grading has failed to remove P. japonica from blueberries, where adult beetles eat inside the berry and can not be seen. Movement of beetles by aircraft from infested to protected States is regulated by USDA/APHIS (2004). Compliance agreements require aircraft at regulated airports to be sprayed with an insecticide. In addition, express package services have expended millions of dollars to use physical excluders and teams of beetle spotters to reduce the chance of beetles getting on a plane. Aircraft are examined upon arrival in protected USA states, and retreated if live beetles are found.


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                                                Spicer PG, Potter DA, McNiel RE, 1995. Resistance of flowering crabapple cultivars to defoliation by the Japanese beetle (Coleoptera: Scarabaeidae). Journal of Economic Entomology, 88(4):979-985

                                                Switzer, P. V., Cumming, R. M., 2014. Effectiveness of hand removal for small-scale management of Japanese beetles (Coleoptera: Scarabaeidae). Journal of Economic Entomology, 107(1), 293-298. doi: 10.1603/EC12303

                                                Szendrei, Z., Mallampalli, N., Isaacs, R., 2005. Effect of tillage on abundance of Japanese beetle, Popillia japonica Newman (Col., Scarabaeidae), larvae and adults in highbush blueberry fields. Journal of Applied Entomology, 129(5), 258-264. doi: 10.1111/j.1439-0418.2005.00961.x

                                                Tashiro H, 1987. Turfgrass insects of the United States and Canada. Ithaca, New York, USA; Cornell University Press, xiv + 391 pp

                                                Terry LA, Potter DA, Spicer PG, 1993. Insecticides affect predatory arthropods and predation on Japanese beetle (Coleoptera: Scarabaeidae) eggs and fall armyworm (Lepidoptera: Noctuidae) pupae in turfgrass. Journal of Economic Entomology, 86(3):871-878

                                                Tumlinson JH, Klein MG, Doolittle RE, Ladd TL, Proveaux AT, 1977. Identification of the female Japanese beetle sex pheromone: inhibition of male response by an enantiomer. Science, l97:789-792

                                                USDA, 1975. Japanese beetle quarantines (map). Cooperative Economic Insect Report, 25(9):110

                                                USDA/APHIS, 2000. Managing the Japanese Beetle. A Homeowner's Handbook.,cic_text/housing/japanese-beetle/jbeetle.html

                                                USDA/APHIS, 2004. Japanese beetle program manual for airports. Marketing and Regulatory Programs.

                                                Vega, FE, Dowd PF, Lacey LA, Pell JK, Jackson DM, Klein MG, 2007. Dissemination of beneficial microbial agents by insects. In: Field Manual of Techniques in Invertebrate Pathology [ed. by Lacey LA, Kaya HK] Dordrecht, The Netherlands: Springer, 127-148

                                                Villani MG, Wright RJ, 1988. Entomogenous nematodes as biological control agents of European chafer and Japanese beetle (Coleoptera: Scarabpidae) larvae infesting turfgrass. Journal of Economic Entomology, 81(2):484-487

                                                Villani MG, Wright RJ, Baker PB, 1988. Differential susceptibility of Japanese beetle, Oriental beetle, and European chafer (Coleoptera: Scarabaeidae) larvae to five soil insecticides. Journal of Economic Entomology, 81(3):785-788

                                                Vittum PJ, 1986. Biology of the Japanese beetle (Coleoptera: Scarabpidae) in eastern Massachusetts. Journal of Economic Entomology, 79(2):387-391

                                                Vittum PJ, Villani MG, Tashiro H, 1999. Turfgrass Insects of the United States and Canada. Ithaca, USA: Cornell University Press, 422 pp

                                                Wickizer SL, Gergerich RC, 2007. First report of Japanese beetle (Popillia japonica) as a vector of Southern bena mosaic virus and Bean pod mottle virus. Plant Disease, 91(5):637. HTTP://

                                                Wright RJ, Villani MG, Agudelo-Silva F, 1988. Steinernematid and heterorhabditid nematodes for control of larval European chafers and Japanese beetles (Coleoptera: Scarabpidae) in potted yew. Journal of Economic Entomology, 81(1):152-157

                                                Zenger JT, Gibb TJ, 2001. Identification and impact of egg predators of Cyclocephala lurida Bland and Popillia japonica Newman (Coleoptera: Scaraaeidae) in turfgrass. Environmental Entomology, 30:425-430

                                                Distribution References

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

                                                CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

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                                                Commonwealth Institute of Entomology, 1978. [Nos. 379-384 and 16, 64 and 215 (revised).]. In: Distribution Maps of Pests, Series A (Agricultural),

                                                EPPO, 2014. EPPO Global database (available online). Paris, France: EPPO.

                                                EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO.

                                                Fleming WE, 1972. Biology of the Japanese beetle. In: USDA Technical Bulletin 1449, Washington, DC,

                                                Imura O, 2003. Herbivorous arthropod community of an alien weed Solanum carolinense L. Applied Entomology and Zoology. 38 (3), 293-300. DOI:10.1303/aez.2003.293

                                                Keathley C P, Potter D A, 2008. Quantitative resistance traits and suitability of woody plant species for a polyphagous scarab, Popillia japonica Newman. Environmental Entomology. 37 (6), 1548-1557. DOI:10.1603/0046-225X-37.6.1548

                                                Kim J Y, Leal W S, 2000. Ultrastructure of pheromone-detecting sensillum placodeum of the Japanese beetle, Popillia japonica Newmann (Coleoptera: Scarabaeidae). Arthropod Structure & Development. 29 (2), 121-128. DOI:10.1016/S1467-8039(00)00022-0

                                                Klein M G, Lacey L A, 1999. An attractant trap for the autodissemination of entomopathogenic fungi into populations of the Japanese beetle, Popillia japonica (Coleoptera: Scarabaeidae). Biocontrol Science and Technology. 151-158.

                                                Lacey L A, Amaral J J, Coupland J, Klein M G, Simões A M, 1995. Flight activity of Popillia japonica (Coleoptera: Scarabaeidae) after treatment with Metarhizium anisopliae. Biological Control. 5 (2), 167-172. DOI:10.1006/bcon.1995.1020

                                                Lacey L A, Martins A, Ribeiro C, 1994. The pathogenicity of Metarhizium anisopliae and Beauveria bassiana for adults of the Japanese beetle, Popillia japonica (Coleoptera: Scarabaeidae). European Journal of Entomology. 91 (3), 313-319.

                                                Loughrin J H, Potter D A, Hamilton-Kemp T R, 1995. Volatile compounds induced by herbivory act as aggregation kairomones for the Japanese beetle (Popillia japonica Newman). Journal of Chemical Ecology. 21 (10), 1457-1467. DOI:10.1007/BF02035145

                                                Mabry T R, Hobbs H A, Steinlage T A, Johnson B B, Pedersen W L, Spencer J L, Levine E, Isard S A, Domier L L, Hartman G L, 2003. Distribution of leaf-feeding beetles and bean pod mottle virus (BPMV) in Illinois and transmission of BPMV in soybean. Plant Disease. 87 (10), 1221-1225. DOI:10.1094/PDIS.2003.87.10.1221

                                                Morales-Rodriguez A, Peck D C, 2009. Synergies between biological and neonicotinoid insecticides for the curative control of the white grubs Amphimallon majale and Popillia japonica. Biological Control. 51 (1), 169-180. DOI:10.1016/j.biocontrol.2009.06.008

                                                NAPIS, 2009. Reported Status of Japanese Beetle - Popillia japonica. In: Reported Status of Japanese Beetle - Popillia japonica, Washington DC, USA: USDA Animal and Plant Health Inspection Service.

                                                Ping L, 1988. The Popillia fauna of China., Pianze Eldonejo. 71 pp.

                                                Rahemi A, Dale A, Fisher H, Kelly J, Taghavi T, Singleton C, Bonnycastle A, 2015. Distribution of pests on Vitis riparia in sandy soils of the south-western Ontario. Journal of Plant Studies. 4 (1), 21-26.

                                                Ranney T G, Walgenbach J F, 1992. Feeding preference of Japanese beetles for taxa of birch, cherry and crabapple. Journal of Environmental Horticulture. 10 (3), 177-180.

                                                Reed D K, Lee M H, Kim S H, Klein M G, 1991. Attraction of scarab beetle populations (Coleoptera: Scarabaeidae) to Japanese beetle lures in the Republic of Korea. Agriculture, Ecosystems & Environment. 36 (3-4), 163-174. DOI:10.1016/0167-8809(91)90013-N

                                                Seagraves B L, Redmond C T, Potter D A, 2013. Relative resistance or susceptibility of maple (Acer) species, hybrids and cultivars to six arthropod pests of production nurseries. Pest Management Science. 69 (1), 112-119. DOI:10.1002/ps.3375

                                                Switzer P V, Escajeda K, Kruse K C, 2001. Pairing patterns in Japanese beetles (Popillia japonica Newman): effects of sex ratio and time of day. Journal of Insect Behavior. 14 (6), 713-724. DOI:10.1023/A:1013075915697

                                                Szendrei Z, Mallampalli N, Isaacs R, 2005. Effect of tillage on abundance of Japanese beetle, Popillia japonica Newman (Col., Scarabaeidae), larvae and adults in highbush blueberry fields. Journal of Applied Entomology. 129 (5), 258-264. DOI:10.1111/j.1439-0418.2005.00961.x

                                                Tashiro H, 1987. Turfgrass insects of the United States and Canada. Ithaca, New York, USA: Cornell University Press. xiv + 391pp.

                                                Wickizer S L, Gergerich R C, 2007. First report of Japanese beetle (Popillia japonica) as a vector of Southern bena mosaic virus and Bean pod mottle virus. Plant Disease. 91 (5), 637. HTTP:// DOI:10.1094/PDIS-91-5-0637C

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                                                GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway source for updated system data added to species habitat list.


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                                                27/03/2008 Updated by:

                                                Michael Klein, Ohio Agricultural Research and Development Center, Adjunct Associate Professor, 1680 Madison Avenue, Wooster, OH 44691, USA

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