(Comstock, 1881)
Taxonomic note
Diaspidiotus perniciosus belongs to a group of species that are difficult to assign to distinct genera - see Genus Diaspidiotus, Background.
Diagnosis
In life, female scale cover more or less circular, flat to slightly convex, 1.0-2.2 mm diameter, grey or light brown with orange or yellow-brown, central or subcentral exuviae DIASPEL3.jpg and DIASPEL7.jpg .The first-[l][m]Glossary[/m][r]instar[/r]instar exuviae have a crater-like appearance. Male scale cover elongate oval, grey, smaller than that of female, with yellow subterminal exuviae (Davidson and Miller, 1990) DIASPEL9.jpg . First instar scale cover blue-black. Body of living female yellow to orange (Stoetzel, 1975; Gill, 1997); egg-laying females almost circular. Adult male alate, with orange body (Stoetzel, 1975).
Body of slide-mounted adult female membranous and pyriform, 0.8-1.5 mm long; several prepygidial abdominal segments each with a few submarginal macroducts on either side, but prepygidial marginal macroducts absent DIASPES.jpg . Anus smaller than a median lobe, situated at least 2.4 times its own length from base of median lobes; perivulvar pores absent. Pygidium with median and second lobes well developed, slightly convergent, each notched on outer margin; second lobes usually as long or longer than associated marginal seta; third lobes usually represented by points, occasionally absent; fourth lobes absent DIASPEP1.jpg . Paraphyses present between positions of third lobes only, shorter than the lobes, usually thickened towards their inner ends. Plates between lobes pointed or slightly fimbriate; median and first interlobular spaces each containing two plates DIASPEMG.jpg .
The adult male of D. perniciosus was described by Bustshik, 1958.
Host range
Diaspidiotus perniciosus is highly polyphagous, especially on trees and shrubs, and has been recorded from hosts belonging to 240 genera in 81 plant families, but favours Rosacae, e.g. Malus, Prunus, Pyracantha and Pyrus (Davidson and Miller, 1990). It will probably feed on a wider range of plants than indicated below, but on some of these plants the insect cannot finish its development. The main host plants are different in different parts of the world (Kozár, 1990b), so for each region only the local literature sources can give a true picture. A range of conifers can be attacked (Zahradník, 1990). Different varieties of fruit species show different degrees of susceptibility, and many fruit species have been studied in this respect (Bichina and Gatina, 1976; Shalamberidze, 1978; Verma and Srivastava, 1990). Host records include species of: Acacia, Acer, Actinidia deliciosa, Aesculus, Ailanthus, Akebia, Albizia, Aleurites, Alnus, Aloe, Althaea, Amelanchier, Ampelopsis, Antirrhinum, Aralia, Arbutus, Aristolochia, Asclepias, Asparagus officinalis, Aster, Aucuba, Baccharis, Bambusa, Berberis, Betula, Bignonia, Buddleja, Buxus, Callistemon, Camellia, Canna, Caragana, Carpinus, Carya, Caryopteris, Castanea, Catalpa, Ceanothus, Cedrus, Celtis, Cephalanthus, Cercis, Chaenomeles, Chamaecyparis, Chrysanthemum, Cinnamomum, Cistus, Citrus spp., Clematis, Clerodendrum, Convolvulus, Cornus, Corylopsis, Corylus, Cotoneaster, Crataegus, Cupressus, Cydonia, Cytisus, Dahlia, Daphne, Daucus, Deutzia, Diervilla, Diospyros spp., Elaeagnus, Erica, Erigeron, Eriobotrya, Eucalyptus, Euonymus, Euphorbia, Fagus, Ficus, Forsythia, Fragaria, Fraxinus, Gaylussacia, Genista, Ginkgo, Gymnocladus, Hamamelis, Hibiscus, Hydrangea, Hypericum, Ilex, Inula, Jasminum, Juglans regia, Juniperus, Kalmia, Kerria, Laburnum, Lagerstroemia, Larix, Laurus, Lespedeza, Ligustrum, Lilium, Lindera, Liquidambar, Liriodendron, Lonicera, Lycium, Maclura, Magnolia, Mahonia, Malus spp., Malva, Melia, Mesembryanthemum, Mespilus, Morus, Myrica, Myrtus, Nerium, Nyssa, Olea, Ostrya, Paeonia, Panicum, Paulownia, Persea, Petroselinum, Phellodendron, Philadelphus, Phlox, Photinia, Physocarpus, Picea, Pinus, Piper, Pittosporum, Platanus, Polygonum, Poncirus, Populus, Potentilla, Prunus spp., Ptelea, Punica, Pyracantha, Pyrus, Quercus, Rhamnus, Rhododendron, Rhus, Ribes, Ricinus, Robinia, Rosa, Rosmarinus, Rubus, Rudbeckia, Ruscus, Salix, Sambucus, Sassafras, Schinus, Sciadopitys, Smilax, Solanum, Sophora, Sorbaria, Sorbus, Spartium, Spiraea, Stephanandra, Symphoricarpos, Tamarix, Taxodium, Taxus, Tecoma, Ternstroemia, Thuja, Tilia, Trachelospermum, Tsuga, Ulmus, Urtica, Vaccinium, Viburnum, Vinca, Vitex, Vitis vinifera, Wisteria and Yucca.
Affected plant stages: vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: usually on bark of twigs DIASPEL4.jpg , sometimes on fruit and leaves DIASPEL6.jpg ; can occur on all plant parts including the roots
Biology and ecology
Reproduction is sexual. There are up to four or five generations per year, depending on climate; overwintering in cold climates is as diapausing larvae (Davidson and Miller, 1990; Kozár, 1990b; Gill, 1997). Stoetzel, 1975, recorded the adult female laying a batch of 10-20 eggs and then 4-5 eggs per day until death. In central Europe the adults appear at the end of April, and in northern Europe 1 or 2 months later. The crawlers continue to appear for 1-2 months: each female produces about 100 crawlers. The development of one generation requires about 45-80 days (Kosztarab and Kozár, 1988). In Iran, D. perniciosus has 4 generations per year (Abivardi, 2001). This pest is tolerant of cold winter temperatures, and can develop at 7°C or above. It can establish in areas with temperature averages of -24.2°C in January and 31.5°C in July, and an annual precipitation of 348-2465 mm: an effective temperature sum of 404.6-577.1°C is necessary for the larvae to emerge (Kozár, 1990).
The first instar crawler is the dispersal phase, and can be distributed by wind, birds or flying insects. The most important means of transport, however, is infested nursery material. This pest has been accidentally introduced to many countries by this method.
The pheromone of this species is known (Gieselmann et al., 1979a); it attracts the males and parasitoids. Colour and sticky traps have been developed to follow the flight of the males and parasitoids (Kozár, 1990c).
According to Nur, 1990, males of D. perniciosus are haploid, with n=4, and females are diploid, with 2n=8.
Symptoms
Diaspidiotus perniciosus injects toxic saliva into the host plant as it feeds; this severely disrupts normal tissue growth. Dwarfing and discolouration near the feeding site is common, especially on first year growth DIASPEL8.jpg and on fruit DIASPEL2.jpg , which develop red patches (Gill, 1997). Infested leaves develop necrotic patches at the feeding sites DIASPEL6.jpg . Stems may become distorted (Davidson and Miller, 1990). In cases of heavy infestation, early senescence occurs and branches or even whole plants may die. Red spotting on the fruits and/or necrotic spots on the leaves do not cause great damage; however, these symptoms can cause problems at plant quarantine inspection, because the species is on the quarantine schedules of many countries.
Economic impact
This species is very damaging to deciduous fruit trees including apple, pear, peach, plum, currants and some woody ornamental plants (Konstantinova, 1976; APPPC, 1987; Davidson and Miller, 1990; Kozár, 1990b; Kosztarab, 1996). Diaspidiotus perniciosus is the most serious scale pest of deciduous tree crops, especially pome and stone fruit, throughout the world; it causes serious damage to almost all kinds of deciduous fruit trees, especially to apple, pear, plum and currant (Kozár, 1990b); it is also a pest of ornamental plants (Kosztarab and Kozár, 1988). It has a noticeable effect on the host within 24 hours of infestation, and the trees lose their vigour and have a shorter lifespan. If left unchecked, D. perniciosus can kill mature trees in 2-3 years. Shortly after introduction to a new country, this pest can infest and kill whole orchards. Discoloured fruit are reduced in grade and overall value (Gill, 1997). Local outbreaks have been observed in different parts of the world on fruit trees and ornamental plants, as in Hungary (Kozár and Drozdjak, 1988), Switzerland (Kozár et al., 1994; Mani et al., 1995), the European part of Russia (Kozár and Konstantinova, 1981), Australia (Baker, 1977), Pakistan (Baluchistan) (Simmonds, 1960a) and Canada (Ker and Sears, 1986). It was regarded as by far the most important pest of apple in the southern USSR, western Transcaucasia and Central Asia (Konstantinova, 1976). In India (Kashmir) it threatened to wipe out the apple-growing industry when it was first accidentally introduced around 1910, and it subsequently became a serious pest of apple in Kulu Valley and south India (Simmonds, 1960a). Charles and Henderson, submitted, record D. perniciosus as a pest on economically important plants in New Zealand. In Chile it is a serious pest of pome and other fruit trees, requiring quarantine inspection of fruits before export (Claps et al., 2001a). Danzig and Pellizzari, 1998, say D. perniciosus is a dangerous pest all over the world. Foldi, 2001, lists this species as an economically important pest in France. It has been mentioned as a pest of almond (Prunus dulcis) (Chua and Wood, 1990).
Crop loss caused by D. perniciosus on different tree species is difficult to assess. Other important economic effects of D. perniciosus are the government regulations regarding movement of infested fruit between various areas and particularly between countries. Quarantine regulation of the movement of fruit has often resulted in considerable loss of revenue (Gill, 1997).
Detection and inspection methods
In cases of heavy infestation, greyish scales can be found on the bark of the trees. Scales feeding on fruit cause red spots on the skin. Lighter infestations can be found by stereomicroscopic examination of the branches, following the survey system described in detail by Kozár, 1990, and Kozár, 1990c.
Phytosanitary risk
Diaspidiotus perniciosus is the species most frequently mentioned on quarantine lists (usually as Quadraspidiotus perniciosus) (Burger and Ulenberg, 1990). In recent years, however, the European Union has deleted D. perniciosus from its quarantine list because it is present in most of the member countries of the European Union.
Immature stages of D. ancylus can be mistaken for D. perniciosus; this sometimes causes problems at pre-export inspection of pome fruits in Chile (Claps et al., 2001a).
Natural enemies
Diaspidiotus perniciosus has numerous parasitoids and predators that have been studied in detail, especially in the Palearctic region (Trjapitzin, 1978; Kosztarab and Kozár, 1988). There are many papers on natural enemies of this pest in different countries (Ahmad and Ghani, 1971; Rosen and DeBach, 1976; Fol'kina, 1978; Myartseva, 1978; Trjapitzin, 1978; Popova, 1979; Darling and Johnson, 1984; Katsoyannos, 1984; Yasnosh, 1985; Bhagat et al., 1988; Titayavan and Davis, 1988; Thakur et al., 1989; Sharma et al., 1990; Bull et al., 1993). These natural enemies are efficient regulators of D. perniciosus, and keep down the population density in natural habitats. Where chemical control is inefficient against D. perniciosus, but kills the natural enemies, local outbreaks can occur in orchards. The efficiency of natural enemies is also reduced in urban regions by pollution, where subsequently the pest can cause damage on ornamental plants.
Parasitoids:
- Ablerus dozieri, in USA
- Aphytis aonidiae, attacking: nymphs, adults; cosmopolitan
- Aphytis diaspidis, attacking: nymphs, adults
- Aphytis mytilaspidis, attacking: nymphs, adults; cosmopolitan
- Aphytis proclia, attacking: nymphs, adults; cosmopolitan
- Coccophagoides murtfeldtii, attacking: nymphs, adults
- Encarsia citrina, attacking: nymphs, adults
- Encarsia perniciosi, attacking: nymphs, adults, in China, Korea, Japan, Taiwan, Caucasus, Soviet Far East; introduced to: India, Pakistan, Australia, USA (California), Chile, Germany, Italy, Europe. Also present in North America, New Zealand
Predators:
- Aphytis sp., in USA
- Chrysopa spp.
- Chilocorus spp.
- Chilocorus bipustulatus, in Greece
- Chilocorus kuwanae, in Sakhalin I. (former USSR)
- Coccidencyrtus sp., in USA
- Coccidencyrtus ensifer, in USA
- Cybocephalus fodori, in Greece
- Hemisarcoptes malus, in Italy
- Leptothrips mali, in USA (California)
- Prospaltella perniciosi, in USA
Pathogens:
- Alternaria sp.
- Coniotherium sp.
- Fusarium sp.
- Sphaerostilbe sp.
Distribution
See Diaspidiotus perniciosus distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species. This species is often found together with other species of Diaspidiotus. It is easy to distinguish because D. perniciosus lacks the perivulvar pores that are present in other Diaspidiotus species infesting fruit trees. Diaspidiotus perniciosus overwinters as first-instar larvae, whereas other species overwinter as second instars or females. Diaspidiotus perniciosus is ovoviviparous, whereas the other species lay eggs.
Diaspidiotus africanus (Marlatt) (grey scale) (Aspidiotus pectinatus Lindinger is a synonym) DIASAFRP.jpg could be confused with D. perniciosus but in slide-mounted adult females, D. africanus differs in lacking prepygidial submarginal macroducts; second lobe usually shorter than the associated marginal seta DIASAFMG.jpg ; and third lobe may or may not be represented by a marginal point. In contrast, D. perniciosus has a few prepygidial submarginal macroducts present on each side of several of the prepygidial segments DIASPEP1.jpg ; second lobe usually moderately developed, as long or longer than associated marginal seta; and third lobe usually represented by a marginal point DIASPEMG.jpg , but occasionally absent. Diaspidiotus africanus is known only from South Africa and South West Africa (Balachowsky, 1956; Munting, 1971a) on twigs of species of Acacia, Albizia, Clerodendrum, Combretum, Cydonia, Ehretia, Ficus carica, Gleditsia, Grewia, Laurus, Ligustrum, Nerium, Olea europaea, Phoenix dactylis, Prunus spp., Psoralea, Pyrus spp., Robinia and Schinus (Balachowsky, 1956; Borchsenius, 1966). This species is a declared pest of deciduous fruit trees in the Western Cape, South Africa, where its control is required by law for export quarantine reasons, as it cannot be distinguished from D. perniciosus in life; hovever, D. africanus does not cause serious damage to fruit trees (Munting, 1971a). Scale cover of adult female circular, 1.5-1.8 mm across, slightly convex, mostly light brown, with central exuviae; scale cover of male similar to that of female but oval and smaller, 1.0 mm long. Munting, 1971a, redescribed D. africanus and discussed its synonyms, morphological variation and distinction from other, similar species.
In life, immature stages of D. ancylus (Putnam) (convex scale, Putnam scale, cochenille de Putnam) DIASANS.jpg can be mistaken for D. perniciosus (Claps et al., 2001a). However, in slide-mounted adult females, D. ancylus possesses perivulvar pores DIASANP.jpg whereas D. perniciosus lacks them DIASPEP1.jpg. Diaspidiotus ancylus is dimorphic. The leaf form of this scale has 3 pairs of well-developed pygidial lobes and has been called Abgrallaspis [Hemiberlesia] howardi (Cockerell) in the past (Gill, 1997); it would key out as Hemiberlesia lataniae in the Picture Key. Diaspidiotus ancylus is a polyphagous species known from Canada (Ontario), USA (continental including Arizona, California, Indiana, Iowa, New Mexico, New York, Tennessee), Mexico, Central America, Brazil (Sao Paulo), Chile (first to eighth regions), Spain (Huelva), Portugal, Germany, South Africa and Australia; it is often found on the bark of branches and trunk, and sometimes on leaves, of basswood, elm and maple. Hosts include species of Acer, Aesculus, Betula, Carya, Catalpa, Celtis, Cephalanthus, Crataegus, Cydonia, Fagus, Fraxinus, Hydrangea, Juglans, Liriodendron, Maclura, Malus sylvestris, Muehlenbeckia, Olea europaea, Oxalis, Persea americana, Populus spp., Prunus spp., Pyrus, Quercus, Ribes, Robinia spp., Salix, Sorbus, Tilia, Ulmus, Viburnum and Vitis vinifera (Munting, 1971a; Konstantinova, 1976; Nakahara, 1982; Zahradník, 1990a; Amparo Blay Golcoechea, 1993; Kosztarab, 1996; Miller, 1996; Gill, 1997; Danzig and Pellizzari, 1998; Claps et al., 2001a; CSIRO, 2001; The Natural History Museum collection, London, UK). Immature stages of D. ancylus can be mistaken for D. perniciosus (Claps et al., 2001a). This species has been a pest on walnuts, cranberries, blueberries and on elms and other ornamentals in the USA in the past; it is probably native to North America (Gill, 1997). Diaspidiotus ancylus was listed as a pest of deciduous fruit trees of regional importance by Kozár, 1990b; heavy infestations can kill twigs and branches (Kosztarab, 1996). One or two generations per year have been recorded in the USA, and overwintering is as adult females (second instar females in Chile) (Kosztarab, 1996). This species does not occur in the former USSR, but featured on the list of quarantine pests for this region (Konstantinova, 1976).
Melanaspis bromiliae (Leonardi) MELBRS.jpg could be misidentified in the key as D. perniciosus, but differs in its host preference and in having the anus located near the centre of the pygidium MELBRP.jpg; in contrast, D. perniciosus has the anus located near the posterior margin of the pygidium DIASPEP1.jpg. Melanaspis bromiliae is known from Cameroon, Côte d'Ivoire, Guinea, South Africa, Seychelles, Togo, Portugal including the Azores, Belgium (under glass), Hungary, Italy (under glass), Netherlands, Spain (Canary Is), Taiwan, Japan, USA (Florida, and under glass in Pennsylvania), Brazil, Bahamas, Bermuda, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, Guatemala, Haiti, Honduras, Jamaica, Martinique, Mexico, Panama, Puerto Rico, USA (District of Colombia, Florida, Hawaii), India, Federated Malay States, Philippines, Singapore, South Mariana Is (Guam), Palau Is, Yap Is, and Pohnpei on pineapple fruit (Ananas spp.) (on which it is a pest), and leaves of dryland taro (Colocasia esculenta), Cocos nucifera, and species of Neoglaziovia and Pandanus and other hosts (Beardsley, 1966; Kawai, 1980; Nakahara, 1982; Deitz and Davidson, 1986; Longo et al., 1995; Miller, 1996; Gill, 1997; Danzig and Pellizzari, 1998; Tao, 1999; Heu, 2002). Colonies MELBRL1.jpg , MELBRIL1.jpg
Comments
Diaspidiotus perniciosus is a temperate species, apparently native to northern China (Burger and Ulenberg, 1990). It was accidentally introduced to the USA, and subsequently to other parts of the world. It is widely distributed in the Palearctic and Nearctic regions (Kozár, 1990a) and continues to spread in Central Asia (Konstantinova et al., 1984). There are many publications about its detailed distribution and importance in different parts of the world (Konstantinova, 1976; Baker, 1977; Chowdhuri, 1977; Kozár and Konstantinova, 1981; CABIIE, 1986; APPPC, 1987; Kozár and Drozdjak, 1988; Davidson and Miller, 1990; Kozár et al., 1994; Mani et al., 1995; CABI/EPPO, 1998c). Diaspidiotus perniciosus has been reported from South Africa but its presence there has not been confirmed (Danzig, 1993; EPPO, 1999). Diaspidiotus perniciosus has not been recorded from most of the Pacific islands.
Europe
Albania: present, no further details (EPPO, 1999)
Austria: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Bulgaria: restricted distribution (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Croatia: widespread (Kosztarab and Kozár, 1988)
Czech Republic: restricted distribution (EPPO, 1999)
Denmark: present, no further details (Gertsson, 2001)
Former Czechoslovakia: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Former USSR
Azerbaijan: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
East Siberia: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Far East: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Georgia, Republic of: present, no further details (Danzig, 1993; Aleksidze, 1995; Danzig and Pellizzari, 1998)
Kazakhstan: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Middle Asia: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Moldova: restricted distribution (CABIIE, 1986)
South European Territory: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Tadjikistan: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Transcacasus: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Uzbekistan: present, no further details (Borchsenius, 1966; Danzig, 1993; Danzig and Pellizzari, 1998)
Ukraine: restricted distribution (CABIIE, 1986; Kosztarab and Kozár, 1988)
Former Yugoslavia: widespread (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
France: restricted distribution (Danzig and Pellizzari, 1998; Foldi, 2001)
Germany: restricted distribution (Danzig and Pellizzari, 1998)
Greece: restricted distribution (Kosztarab and Kozár, 1988)
Hungary: widespread (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Italy: widespread (Longo et al., 1995; Danzig and Pellizzari, 1998)
Sardinia: present, no further details (Longo et al., 1995)
Sicily: present, no further details (Longo et al., 1995)
Netherlands: present (Jansen, 2001)
Poland: restricted distribution (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Portugal: restricted distribution (Kosztarab and Kozár, 1988)
Madeira: present, no further details (Kosztarab and Kozár, 1988)
Romania: restricted distribution (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Slovakia: restricted distribution (Kosztarab and Kozár, 1988)
Slovenia: restricted distribution (Kosztarab and Kozár, 1988)
Spain: fairly widespread (Amparo Blay Golcoechea, 1993)
Canary Islands: present, no further details (Amparo Blay Golcoechea, 1993)
Switzerland: restricted distribution (Kosztarab and Kozár, 1988; Kozár et al., 1994; Danzig and Pellizzari, 1998)
Asia
Afghanistan: present, no further details (Ramaseshiah, 1985; Mohammad Ullah, 1988)
Armenia: present, no further details (Danzig, 1993)
Bangladesh: present, no further details (Kosztarab, 1996)
Bhutan: present, no further details (Nakahara, 1982; The Natural History Museum collection, London, UK)
Brunei Darussalam: present, no further details (Kosztarab and Kozár, 1988)
China
Anhui: present, no further details (Tao, 1999)
Guangdong: present, no further details (Tao, 1999)
Hebei: present, no further details (Tao, 1999)
Heilongjiang: present, no further details (Tao, 1999)
Henan: present, no further details (Tao, 1999)
Hong Kong: present, few occurrences (EPPO, 1999)
Hubei: present, no further details (CABIIE, 1986)
Hunan: present, no further details (Tao, 1999)
Inner Mongolia: present, no further details (CABIIE, 1986)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Jilin: present, no further details (CABIIE, 1986)
Liaoning: present, no further details (Tao, 1999)
Manchuria: present, no further details (CABIIE, 1986)
Shaanxi: present, no further details (Tao, 1999)
Shandong: present, no further details (Tao, 1999)
Shanxi: present, no further details (Xie, 1982)
Sichuan: present, no further details (Tao, 1999)
Yunnan: present, no further details (Tao, 1999)
Xinjiang: present, no further details (CABIIE, 1986)
Zhejiang: present, no further details (Tao, 1999)
India: present, no further details (Nakahara, 1982)
Andhra Pradesh: present, no further details (CABIIE, 1986)
Assam: present, no further details (CABIIE, 1986)
Bihar: The Natural History Museum collection, London, UK
Delhi: present, no further details (CABIIE, 1986)
Himachal Pradesh: present, no further details (CABIIE, 1986)
Jammu and Kashmir: present, no further details (Thakur et al., 1989)
Karnataka: present, no further details (CABIIE, 1986)
Maharashtra: present, no further details (CABIIE, 1986)
Orissa: present, no further details (CABIIE, 1986)
Punjab: present, no further details (CABIIE, 1986)
Tamil Nadu: present, no further details (CABIIE, 1986)
Uttar Pradesh: present, no further details (CABIIE, 1986)
West Bengal: present, no further details (CABIIE, 1986)
Iran: restricted distribution (Seghatoleslami, 1977; Abivardi, 2001)
Iraq: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Japan: present (Kawai, 1980)
Hokkaido: widespread (CABIIE, 1986)
Honshu: widespread (CABIIE, 1986)
Kyushu: widespread (CABIIE, 1986)
Shikoku: widespread (CABIIE, 1986)
Korea: present, no further details (CABIIE, 1986; Danzig and Pellizzari, 1998)
Korea, Democratic People's Republic: present, no further details (Danzig, 1993)
Korea, Republic of: present, no further details (Danzig, 1993)
Nepal: present, no further details (CABIIE, 1986)
Pakistan: present, no further details (CABIIE, 1986; Danzig, 1993)
South-East Asia: present, no further details (Danzig and Pellizzari, 1998)
Taiwan: present, no further details (Wong et al., 1999)
Thailand: present, no further details (Danzig, 1993)
Turkey: restricted distribution (Kozár et al., 1979; Danzig and Pellizzari, 1998)
Vietnam: present, no further details (Danzig, 1993)
Africa
Algeria: widespread (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Angola: present, no further details (Nakahara, 1982)
Congo Democratic Republic: present, no further details (CABIIE, 1986; Danzig, 1993)
Eritrea: The Natural History Museum collection, London, UK
Morocco: restricted distribution (EPPO, 1999)
South Africa: present, no further details (Munting, 1971a)
Tunisia: restricted distribution (EPPO, 1999)
Zimbabwe: restricted distribution (Munting, 1971a; CABIIE, 1986)
Western Hemisphere
Argentina: widespread, commoner in the South (Borchsenius, 1966; Claps et al., 2001a)
Buenos Aires: present, no further details (CABIIE, 1986; Claps et al., 2001a)
La Pampa: present, no further details (Claps et al., 2001a)
Mendoza: present, no further details (Claps et al., 2001a)
Santa Fe: present, no further details (Claps et al., 2001a)
Tucumán: present, no further details (Claps et al., 2001a)
Bolivia: widespread (CABIIE, 1986)
Brazil: widely distributed (Claps et al., 2001a)
Minas Gerais: present, no further details (Claps et al., 2001a)
Paraná: present, no further details (Claps et al., 2001a)
Rio de Janeiro: present, no further details (Claps et al., 2001a)
Rio Grande do Sul: present, no further details (Claps et al., 2001a)
Santa Caterina: present, no further details (Claps et al., 2001a)
Sao Paulo: present, no further details (Claps et al., 2001a)
Canada: restricted distribution (Borchsenius, 1966; CABIIE, 1986)
British Columbia: restricted distribution (CABIIE, 1986)
Nova Scotia: present, no further details (CABIIE, 1986)
Ontario: restricted distribution (Kosztarab, 1996)
Québec: restricted distribution (CABIIE, 1986)
Central America: present, no further details (Danzig and Pellizzari, 1998)
Chile: widespread (CABIIE, 1986)
Antofagasta: present, no further details (Claps et al., 2001a)
Atacama: present, no further details (Claps et al., 2001a)
Biobío: present, no further details (Claps et al., 2001a)
Coquimbo: present, no further details (Claps et al., 2001a)
La Araucanía: present, no further details (Claps et al., 2001a)
Los Lagos: present, no further details (Claps et al., 2001a)
Maule: present, no further details (Claps et al., 2001a)
O'Higgins: present, no further details (Claps et al., 2001a)
Santiago: present, no further details (Claps et al., 2001a)
Valparaiso: present, no further details (Claps et al., 2001a)
Cuba: present, no further details (CABIIE, 1986)
Ecuador: restricted distribution (EPPO, 1999)
Mexico: present (Miller, 1996)
Paraguay: restricted distribution (Nakahara, 1982)
Peru: widespread (CABIIE, 1986)
St. Martin: restricted distribution (EPPO, 1999)
Uruguay: present, no further details (CABIIE, 1986)
USA: generally distributed in continental USA (Nakahara, 1982) but not present in Maine, North and South Dakota, Wyoming (CABIIE, 1986) or California (Gill, 1997)
District of Colombia: present, no further details (CABIIE, 1986)
Hawaii: present, no further details (Borchsenius, 1966)
Illinois: present, no further details (CABIIE, 1986)
Massachusetts: present, no further details (CABIIE, 1986)
Michigan: present, no further details (CABIIE, 1986)
New Jersey: present, no further details (CABIIE, 1986)
New York: present, no further details (CABIIE, 1986)
Ohio: present, no further details (Kosztarab, 1996)
Oregon: present, no further details (CABIIE, 1986)
Tennessee: present, no further details (EPPO, 1999)
Utah: The Natural History Museum collection, London, UK
Virginia: The Natural History Museum collection, London, UK
Washington: present, no further details (CABIIE, 1986)
West Virginia: present, no further details (Kozár et al., 1994)
Venezuela: widespread (CABIIE, 1986; EPPO, 1999)
Oceania
Australia: restricted distribution (Kosztarab and Kozár, 1988)
New South Wales: present, no further details (CABIIE, 1986)
Queensland: widespread (CABIIE, 1986)
South Australia: present, no further details (CABIIE, 1986)
Victoria: restricted distribution (CABIIE, 1986)
Tasmania: present, no further details (CABIIE, 1986)
Western Australia: present, no further details (CABIIE, 1986)
Caroline Islands: restricted distribution (EPPO, 1999)
New Zealand: widespread (CABIIE, 1986; Charles and Henderson, submitted)