(Targioni Tozzetti, 1886)
Diagnosis
Scale cover of adult female in life more or less circular, slightly convex, white or yellow-white, 1.5-2.8 mm in diameter, white with subcentral to submarginal yellow or reddish brown exuviae PSPENL2.jpg . Sometimes the scale cover is formed just under the cuticle of the host's stem, so the scale is partially concealed under plant tissue (Dekle, 1976). Scale cover of male smaller than that of female, narrow, with parallel sides, sometimes with a median longitudinal ridge; white with yellow terminal exuviae PSPENL8.jpg . Beneath the scale, the young adult female is pear shaped, orange-yellow; an older, egg-laying female appears almost circular. Exposed body of adult female cream to yellow PSPENL1.jpg and PSPENL5.jpg . Adult male winged (Ghauri, 1962) PSPENL7.jpg .
Body of slide-mounted female 0.9-1.1 mm long, turbinate (less than twice as long as wide), membranous; margins strongly lobed; dorsal ducts absent from median areas of thorax PSPENS.jpg . Pygidium with 3 pairs of well-developed lobes, each notched on the outer margin; median lobes zygotic, with pair of marginal setae between them; orifices of dorsal macroducts elongate oval, each marginal duct with long axis of orifice perpendicular to margin; single gland spine present on each side of pygidial segment VI; each pygidial gland spine containing several microducts and usually with a trifurcate tip PSPENP.jpg .
Host range
Pseudaulacaspis pentagona is polyphagous. However, it cannot complete development on some of the hosts that have been listed by various authors, which indicates that some may not be true host plants. Davidson and Miller, 1990, record it from hosts in 115 genera in 55 plant families. Hosts include species of: Abelmoschus esculentus, Acacia, Acer, Actinidia, Aesculus, Aleurites, Allamanda, Annona, Argyreia, Artocarpus altilis, Asparagus, Azadirachta, Bauhinia, Berberis, Bignonia, Bouvardia, Brachychiton, Broussonetia, Buddleja, Cajanus, Callicarpa, Calotropis, Camellia sinensis, Campsis, Capsicum, Carica papaya, Cassia, Casuarina, Catalpa, Cedrela, Cinnamomum, Celtis, Clematis, Cocos nucifera , Consolida, Cornus, Crotalaria, Cycas, Cydonia, Cytisus, Diospyros kaki, Ehretia, Erythrina, Euonymus, Euphorbia, Fagara, Ficus, Firmiana, Flacourtia spp., Fraxinus, Fuchsia, Genista, Geranium, Ginkgo, Gleditsia triacanthos, Gossypium, Guazuma, Gymnocladus, Heliotropium, Hevea brasiliensis, Hibiscus, Hypericum, Ipomoea, Jasminum, Juglans, Kalanchoe pinnata, Koelreuteria, Ligustrum, Macfadyena, Mallotus japonicus, Malus sylvestris, Mangifera indica, Manihot esculenta, Melia, Mikania, Morus spp., Nephelium, Nerium, Olea europaea, Ostrya, Paeonia, Passiflora, Paulownia, Pelargonium, Persea, Phaseolus, Phellodendron, Philadelphus, Phoenix, Photinia, Piper spp., Platanus, Plumeria, Polygala, Populus, Prunus spp., Psidium, Ptelea, Pterocarya, Pyrus, Quercus, Rhus, Ribes, Ricinus communis, Robinia, Rubus, Salix, Schinus, Sedum, Sida, Solanum, Sophora, Sorbus, Spartium, Sterculia urens, Strelitzia, Symphoricarpos, Tecoma, Theobroma, Triumfetta, Tylophora asthmatica, Ulmus, Urena, Veronica, Vincetoxicum, Vitis vinifera, Zamia, Zanthoxylum and Zelkova.
Affected plant stages: seedling, vegetative growing, flowering and fruiting stages
Affected plant parts: whole plant; stems, bark PSPENCOL.jpg and PSPENTWG.jpg , fruit PSPENDAM.jpg ; rarely on leaves or roots
Biology and ecology
Pseudaulacaspis pentagona reproduces sexually, with two to five generations per year depending on climate (Davidson and Peairs, 1966). Pheromones are emitted by females to attract males. The structure and composition of the P. pentagona pheromone is known (Heath et al., 1979).
Pseudaulacaspis pentagona is thermophilous, so it is found only under glass in colder countries. Females each lay about 100 eggs (Ball, 1980), which hatch 3-5 days after laying (Kosztarab, 1996). The species has between one and four generations per year depending on the climate; overwintering is as adult females and eggs in the USA (Davidson et al., 1983). Development under local conditions is described by Bobb et al., 1973; Shinano et al., 1976; Paloukis, 1979; Stimmel, 1982; Park and Kim, 1990; Hanks and Denno, 1993; and Erkiliç and Uygun, 1997. Pseudaulacaspis pentagona overwinters in cold countries as adult females. Crawlers appear 1-2 months after egg-laying (Ball, 1980). There are three generations per year on twigs and trunks of tea in Japan (Murakami, 1970).
Like other diaspidids, the main dispersal stage of P. pentagona is the mobile first instar, which has legs. Crawlers can walk up to perhaps 1 m, but can be distributed across much greater distances by wind, flying insects and birds. It is readily carried on consignments of plant material and fruit.
According to Nur, 1990, the females of this species each have 2n=16 chromosomes, but P. pentagona is unlikely to be tetraploid; males are 2n=8. During her life, a female initially produces offpring of one sex, then both sexes, and finally the other sex.
Symptoms
Heavy infestations of P. pentagona are often found as thick crusts of scales on tree trunks and older branches in temperate regions, and rarely on the roots. The large white colonies of females and males on the branches that make up a heavy infestation are easy to recognize. Leaves and fruits are not usually infested, but infestation of fruit can cause discolouration PSPENDAM.jpg .
Plant symptoms vary slightly between different hosts, but include: leaves with necrotic areas, yellowed or dead, showing early senescence and abnormal leaf fall; dieback or distortion of stems and discoloration of bark. The whole plant may be dwarfed, die back or even die. In the case of heavy infestations, branches or entire trees can die. Heavily infested plants may die some years after the onset of infestation; younger plants are more susceptible.
Economic impact
Pseudaulacaspis pentagona was described as a very destructive pest by Kosztarab, 1996, especially on flowering cherry, mulberry, peach and other deciduous fruit trees. It mainly damages deciduous fruits and nuts, including peach, currant, grape, kiwi, walnut; it also attacks some woody ornamental plants, including Morus, Sophora, Syringa, Catalpa, Euonymus and Paulownia (APPPC, 1987; Borchsenius, 1966; Konstantinova, 1976; Davidson and Miller, 1990; Kozár, 1990b; Kosztarab, 1996). In Argentina it is as common on garden plants as it is on fruit trees (Claps et al., 2001a). Crop losses caused by P. pentagona are difficult to assess, but Williams and Watson, 1988, describe it as a destructive species, and Danzig and Pellizzari, 1998, say it is a dangerous pest of fruit and ornamental plants. Foldi, 2001, lists this species as an economically important pest in France. It damages olives seriously in Greece (Argyriou and Kourmadas, 1981). Pseudaulacaspis pentagona can be a pest on cassava, attacking the stems and leaf bases of weak or old plants (Chua and Wood, 1990). Infested trees lose vigour and their lives are shortened, and young plants can die very quickly after infestation. Surrounding vegetation can be a source of re-infestation. When P. pentagona becomes newly established in a region, this may lead to the loss of whole trees and plantations. This species is a quarantine pest in many parts of the world; infested plants may be refused entry into many countries.
In recent years, outbreaks have been observed in different parts of the world on several species of fruit, especially kiwi and peach (Whitmore and Medina Gaud, 1974; Kozarzevskaja and Vlainic, 1981; Kozarzevskaja et al., 1986; Ram and Pathak, 1987; and Hickel et al., 1996). In Europe, heavy outbreaks have occurred on ornamental plants in Hungary (Kosztarab and Kozár, 1988) and Switzerland (Kozár et al., 1994; and Mani et al., 1997).
P. pentagona has caused major problems in areas where it was accidentally introduced in the absence of its natural regulators. The efficiency of natural enemies is reduced in urban areas by pollution; consequently, P. pentagona can cause severe damage to ornamental plants in towns and cities.
Detection and inspection methods
Infestations may be confirmed by laboratory examination with stereomicroscopic analyses of the branches, following the survey system described in detail by Kozár, 1990b; and Kozár, 1990c. Heavy infestations of P. pentagona on the bark of trees produce large, visible white colonies of female and male scalesPSPENCOL.jpg and PSPENTWG.jpg .
Pheromone traps are widely used for detection in newly infested regions, especially in Europe (Kozár et al., 1997). Colour and sticky traps were have also been developed to monitor the flight and dispersal of males and parasitoids (Kozár, 1990c; and Kozár et al., 1995).
Phytosanitary risk
Pseudaulacaspis pentagona is the second most frequently mentioned species on quarantine lists (after Diaspidiotus perniciosus) (Burger and Ulenberg, 1990), but not in EU countries. For details of quarantine regulations for P. pentagona see Anon, 1976; and Anon, 1993.
Natural enemies
Pseudaulacaspis pentagona is attacked by a large number of parasitoids and predators, which have been studied in detail especially in the Palaearctic region (Trjapitzin, 1978; Kosztarab and Kozár, 1988). Studies of natural enemies of P. pentagona include: Zaets, 1970; Rosen and DeBach, 1976; Gordon, 1978; Yasuda, 1981; Darling and Johnson, 1984; Noyes and Hui, 1987; Liebregts et al., 1989; Gordon and Hilburn, 1990; Habibian, 1991; Li, 1991, and Hanks and Denno, 1993. A comprehensive list of the natural enemies of P. pentagona is provided by Waterhouse and Norris, 1987.
These natural enemies are efficient regulators of P. pentagona; they keep pest density down in natural habitats. Where chemical control is used against P. pentagona in orchards its natural enemies can be killed, causing local outbreaks. Pseudaulacaspis pentagona has caused major problems in areas where it was accidentally introduced, in the absence of its natural regulators. Coccinellids and a number of parasitoids including Encarsia berlesei can be effective control agents. The efficiency of natural enemies is reduced in urban areas by pollution; consequently, P. pentagona can cause severe damage to ornamental plants in towns and cities.
There have been a number of successful biological control programmes against P. pentagona in different parts of the world, especially in the USA, Europe and Russia. Usually Encarsia (Prospaltella) berlesei is reared, released, and established, according to the protocols of Rosen, 1990a (see also Greathead, 1976). Encarsia berlesei was released in several European countries, and has spread into neighbouring countries. Encarsia diaspidicola has been successfully released in Western Samoa (Sands et al., 1990).
Many of the listed parasitoids of P. pentagona may also be hyperparasitoids. The pheromone released by P. pentagona females is not known to attract parasitoids.
Parasitoids:
- Ablerus atomon, attacking: nymphs, adults
- Ablerus lepidus, attacking: nymphs, adults
- Ablerus pentagona, attacking: nymphs, adults
- Ablerus perspeciosus, attacking: nymphs, adults, in Japan
- Ablerus platensis, attacking: nymphs, adults
- Aphytis chionaspis, attacking: nymphs, adults
- Aphytis diaspidis, attacking: nymphs, adults, in China; Japan; Introduced: Bermuda; Jamaica
- Aphytis lingnanensis, attacking: nymphs, adults
- Aphytis proclia, attacking: nymphs, adults, in Southeast Asia; Introduced: Italy
- Aprostocetus purpureus
- Arrhenophagus albitibiae
- Arrhenophagus chionaspidis in Japan
- Coccophagoides kuwanai, attacking: nymphs, adults
- Comperiella bifasciata
- Encarsia berlesei, attacking: nymphs, adults, in China; Japan; Introduced: Austria; France; Italy; Spain; Switzerland; Russia; Argentina; Brazil; Peru; Puerto Rico; Uraguay; Samoa
- Encarsia citrina, attacking: nymphs, adults
- Encarsia diaspidicola, attacking: nymphs, adults, in Southern Europe, Italy; Introduced: Japan; Samoa
- Encarsia perniciosi, attacking: nymphs, adults
- Epitetracnemus comis
- Marietta leopardina, attacking: nymphs, adults
- Pteroptrix bicolor, in Japan
- Pteroptrix orientalis, attacking: nymphs, adults
- Thomsonica amathus, in Japan
- Signiphora aspidioti
- Zaomma lambinus
Predators:
- Adalia bipunctata, attacking: nymphs, adults
- Chilocorus cacti, attacking: nymphs, adults
- Chilocorus circumdatus, attacking: nymphs, adults
- Chilocorus hupehanus, attacking: nymphs, adults
- Chilocorus kuwanae, attacking: nymphs, adults, in China, Japan
- Chilocorus nigrita, attacking: nymphs, adults
- Chilocorus politus, attacking: nymphs, adults
- Chilocorus renipustulatus, attacking: nymphs, adults
- Coccidophilus cariba, attacking: nymphs, adults
- Coccidophilus citricola, attacking: nymphs, adults
- Cryptognatha nodiceps, attacking: nymphs, adults
- Cryptognatha simillima, attacking: nymphs, adults
- Cybocephalus gibbulus, attacking: nymphs, adults
- Decadiomus hughesi, attacking: nymphs, adults
- Dentifibula viburni
- Exochomus quadripustulatus, attacking: nymphs, adults
- Pentilia insidiosa, attacking: nymphs, adults
- Pharoscymnus horni, attacking: nymphs, adults
- Pharoscymnus tomeensis, attacking: nymphs, adults
- Prodilis sp. nr. gorhami, attacking: nymphs, adults
- Rhyzobius lophanthae, attacking: nymphs, adults, in Australia; Introduced: Southern Europe
- Rhyzobius pulchellus, attacking: nymphs, adults, in Vanuatu, New Caledonia, Peru
- Sticholotis quadrisignata, attacking: nymphs, adults
- Sukunahikona prapawan, attacking: nymphs, adults
Pathogens:
- Nectria flammea in Japan
Distribution
See Pseudaulacaspis pentagona distribution.
Microscopic examination of slide-mounted adult females is required to identify P. pentagona authoritatively.
Borchsenius, 1966 treated P. prunicola (Maskell) as a synonym of P. pentagona. However, Davidson and Miller, 1990; Davidson et al., 1983; Danzig, 1993; and Kosztarab, 1996, treated these species separately and they are treated as separate species here. Pseudaulacaspis pentagona can be differentiated from P. prunicola by the number and shape of gland spines on the pygidial margin. In each of the spaces between the second and third, and third and fourth lobes, P. pentagona has a single gland spine with usually a trifurcate tip PSPENP.jpg; P. prunicola usually has two gland spines in this position, each with a bifid or pointed tip PSPRUMG1.jpg. Also, P. pentagona has 28-106 large macroducts on one side of the body, while P. prunicola has 24-86 large macroducts in this area (Davidson et al., 1983). However, these characters show great variability depending on temperature, geographic locality and host plant (Danzig, 1993). Pseudaulacaspis pentagona is a tropical/subtropical species, whereas P. prunicola prefers cool climates (Kozár, 1990b; Davidson and Miller, 1990). Pseudaulacaspis prunicola (white prunicola scale) PSPRUNL4.jpg occurs in USA (Alabama, Connecticut, District of Colombia, Florida, Louisiana, Maryland, Massachusetts, Mississippi, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, Virginia, Western Virginia; possibly also in Oregon and Hawaii), China, Taiwan, Korea, and Japan (Amami, Honshu, Okinawa) (1977 and 1990 data from S. Kawai photographs; Kawai, 1980; Davidson et al., 1983). It is commonly found on the bark/stems and fruit of its hosts. It has been recorded on hosts belonging to 21 genera in 16 plant families (Davidson and Miller, 1990) including species of Jasminum, Ligustrum, Prunus (especially flowering cherries) and Syringa (Davidson and Miller, 1990; Danzig and Pellizzari, 1998); in contrast, P. pentagona is not usually found on Syringa (Davidson et al., 1983). Pseudaulacaspis prunicola causes aesthetic damage to ornamental plants because the white male scale covers form concentrations on the undersides of branches, making the plant unsightly PSPRUNL1.jpg ; heavy infestations can cause dieback or even death of flowering cherries (Davidson and Miller, 1990). In the USA there are two (Pennsylvania) or three (Maryland) generations per year and the species overwinters as gravid females. Scale cover of adult female more or less circular, slightly convex, white or yellow-white, with subcentral to submarginal yellow exuviae PSPRUNL3.jpg . Scale cover of male smaller than that of female, narrow, with parallel sides, sometimes with a median longitudinal ridge; white with yellow terminal exuviae (Davidson and Miller, 1990). Male scale covers PSPRUNL2.jpg .
Rutherfordia major (Cockerell) (lychee bark scale, white litchi scale, large snow scale) could be misidentified as P. pentagona, but differs by having gland spines on the pygidium thick and blunt, and dorsal macroducts present in median areas of the thorax. In contrast, P. pentagona has gland spines on the pygidium branched or bifid distally, and dorsal macroducts absent from median areas of the thorax (Williams and Watson, 1988) PSPENP.jpg. Rutherfordia major is a polyphagous species known from Tanzania, Madagascar, Mauritius, Seychelles, Sri Lanka, India (Assam, Kerala), Nepal, Sri Lanka, Indonesia (Java), Taiwan, Tonga, USA (Hawaii), Central America, Brazil (Sao Paulo), Colombia, Venezuela, Antigua and Cuba on the stems and bark of species of Alphitonia, Bridelia, Casearia, Cordia, Cupania, Elattostachys, Euphorbia, Ficus, Flacourtia, Heliotropium, Ipomoea, Litchi, Nephelium, Salix, Santalum and Scolopia (Nakahara, 1982; Williams and Watson, 1988; Williams and Williams, 1988; Takagi et al., 1989; Danzig and Pellizzari, 1998; Tao, 1999; Kondo, 2001; Heu, 2002). The status, affinities and morphology of R. major are discussed by Takagi et al., 1989. See also genus Rutherfordia, Background.
Comments
Pseudaulacaspis pentagona originated in eastern Asia, and is a tropical/subtropical species (Davidson et al., 1983; Kozár, 1990b; Longo et al., 1995) that is now cosmopolitan in distribution. It is found in colder countries only under glass. There are many publications concerning its detailed distribution and importance in different parts of the world (Konstantinova, 1976; Kozár and Konstantinova, 1981; Davidson and Miller, 1990; Kozár et al., 1994; and IIE, 1996a).
Many of the records listed below were published before P. pentagona and P. prunicola were distinguished by Davidson et al., 1983, so the distribution below probably reflects the combined distribution of these species. As currently understood, P. pentagona is tropical/subtropical, whereas P. prunicola is a temperate species (Kozár, 1990b).
In spite of records in the past literature (Nakahara, 1982; EPPO, 1999), P. pentagona is not present in New Zealand now (Charles and Henderson, submitted). It has also not been collected in California since before 1920, so is probably not present there now (Gill, 1997).
This species is probably intercepted at plant quarantine inspection in most countries; however, interceptions go largely unreported. In the past 20 years it has started to spread northwards in Europe in field conditions, possibly as a result of climate change. Where 'eradication' has been reported, P. pentagona may have died out locally because of unsuitable conditions, or it may persist at low densities.
Europe
Bulgaria: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Croatia: widespread (Kosztarab and Kozár, 1988)
Former USSR
Abkhazia: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Adjaria: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Azerbaijan: present, no further details (IIE, 1996a)
Georgia, Republic of: present, no further details (IIE, 1996a)
Russia (Europe): present, no further details (EPPO, 1999)
Russian Far East: present, no further details (EPPO, 1999)
Sakhalin: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Southern Russia: present, no further details (EPPO, 1999)
Transcaucasus: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Ukraine: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Former Yugoslavia: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
France: restricted distribution (Kosztarab and Kozár, 1988; Foldi, 2001)
Germany: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Greece: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Hungary: present, no further details (Danzig and Pellizzari, 1998; Ripka, 1999)
Italy: present, no further details (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)
Macedonia: present, no further details (Kosztarab and Kozár, 1988)
Malta: present, no further details (Danzig and Pellizzari, 1998)
Montenegro: northeast (EPPO, 1999)
Netherlands: present, no further details (Jansen, 1995)
Portugal
Madeira: present, no further details (IIE, 1996a)
Slovakia: present, no further details (Kosztarab and Kozár, 1988)
Slovenia: present, no further details (Kosztarab and Kozár, 1988)
Spain: present in Madrid and Tarragona (Amparo Blay Golcoechea, 1993)
Canary Islands: present, no further details (IIE, 1996a; Amparo Blay Golcoechea, 1993)
Switzerland: present, no further details (Kozár et al., 1994; Kosztarab and Kozár, 1988)
United Kingdom: restricted distribution, currently under eradication (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
Asia
Azerbaijan: present, no further details (EPPO, 1999)
Brunei Darussalam: present, no further details (IIE, 1996a)
China: widespread (Borchsenius, 1966)
Anhui: present, no further details (IIE, 1996a)
Fujian: present, no further details (Tao, 1999)
Gansu: present, no further details (Tao, 1999)
Guangdong: present, no further details (Tao, 1999)
Guangxi: present, no further details (Tao, 1999)
Guizhou: present, no further details (EPPO, 1999)
Hebei: present, no further details (Tao, 1999)
Hennan: present, no further details (Tao, 1999)
Hong Kong: present, no further details (Tao, 1999)
Hubei: present, no further details (IIE, 1996a)
Hunan: present, no further details (Tao, 1999)
Inner Mongolia: present, no further details (Tao, 1999)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Jilin: present, no further details (Tao, 1999)
Liaoning: present, no further details (Tao, 1999)
Nei Menggu: present, no further details (IIE, 1996a)
Ningxia: present, no further details (Tao, 1999)
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)
Xinjiang: present, no further details (Tao, 1999)
Xizang: present, no further details (Tao, 1999)
Yunnan: present, no further details (Tao, 1999)
Zhejiang: present, no further details (Tao, 1999)
India: present, no further details (Nakahara, 1982)
Assam: present, no further details (IIE, 1996a)
Himachal Pradesh: present, no further details (IIE, 1996a)
Jammu and Kashmir: present, no further details (IIE, 1996a)
Karnataka: The Natural History Museum collection, London, UK
Manipur: present, no further details (IIE, 1996a)
Punjab: present, no further details (IIE, 1996a)
Sikkim: present, no further details (IIE, 1996a)
Uttar Pradesh: present, no further details (IIE, 1996a)
West Bengal: present, no further details (IIE, 1996a)
Indonesia: present, no further details (Nakahara, 1982)
Irian Jaya: present, no further details (IIE, 1996a)
Java: present, no further details (IIE, 1996a)
Iran: present, no further details (Seghatoleslami, 1977; Abivardi, 2001)
Iraq: present, no further details (IIE, 1996a)
Israel: present, no further details (Danzig and Pellizzari, 1998)
Japan: present, no further details (Kawai, 1980; Ozawa, 1994; Tao, 1999)
Hokkaido: present, no further details (EPPO, 1999)
Honshu: present, no further details (IIE, 1996a)
Kyushu: present, no further details (IIE, 1996a)
Korea, DPR: present, no further details (Danzig, 1993; IIE, 1996a)
Korea, Republic of: present, no further details (Danzig, 1993; IIE, 1996a)
Malaysia
West Malaysia: present, no further details (IIE, 1996a)
Sabah: present, no further details (IIE, 1996a)
Sarawak: The Natural History Museum collection, London, UK
Maldive Is: present, no further details (Watson et al., 1995)
Nepal: present, no further details (IIE, 1996a)
Philippines: present, no further details (IIE, 1996a)
Ryukyu Archipelago: present, no further details (EPPO, 1999)
Singapore: present, no further details (Waterhouse, 1993; Tao, 1999)
Sri Lanka: present, no further details (IIE, 1996a)
Syria: present, no further details (Danzig and Pellizzari, 1998)
Taiwan: present, no further details (Wong et al., 1999)
Turkey: present, no further details (Kozár et al., 1979; Danzig and Pellizzari, 1998)
Vietnam: present, no further details (Danzig, 1993; IIE, 1996a)
Africa
Cape Verde: present, no further details (IIE, 1996a)
Egypt: present, no further details (IIE, 1996a)
Ghana: present, no further details (IIE, 1996a)
Madagascar: present, no further details (IIE, 1996a)
Malawi: present, no further details (IIE, 1996a)
Mauritius: present, no further details (Williams and Williams, 1988)
Principe: The Natural History Museum collection, London, UK
Réunion: present, no further details (Williams and Williams, 1988)
Sao Tomé: present, no further details (IIE, 1996a)
Seychelles: present, no further details (IIE, 1996a)
South Africa: present, no further details (IIE, 1996a)
St Helena: present, no further details (IIE, 1996a)
Tanzania: present, no further details (IIE, 1996a)
Zanzibar: present, no further details (Nakahara, 1982; IIE, 1996a)
Zimbabwe: present, no further details (IIE, 1996a)
Western Hemisphere
Antigua: present, no further details (IIE, 1996a)
Argentina: very common (Claps et al., 2001a)
Buenos Aires: present, no further details (Claps et al., 2001a)
Coórdoba: present, no further details (Claps et al., 2001a)
Mendoza: present, no further details (Claps et al., 2001a)
Paraná Delta: present, no further details (IIE, 1996a)
San Juan: present, no further details (Claps et al., 2001a)
San Luís: present, no further details (IIE, 1996a)
Santa Fe: present, no further details (Claps et al., 2001a)
Satiago del Estero: present, no further details (Claps et al., 2001a)
Tucumán: present, no further details (Claps et al., 2001a)
Bahamas: present, no further details (IIE, 1996a)
Barbados: present, no further details (Bennett and Alam, 1985)
Bermuda: widespread and common (Hodgson and Hilburn, 1991)
Bolivia: present, no further details (Nakahara, 1982; IIE, 1996a)
Brazil: present, no further details (Tao, 1999)
Amazonas: present, no further details (IIE, 1996a)
Bahia: present, no further details (Claps et al., 2001a)
Ceará: present, no further details (Claps et al., 2001a)
Guanabara: present, no further details (Silva et al., 1968)
Minas Gerais: present, no further details (Claps et al., 2001a)
Pará: present, no further details (Claps et al., 2001a)
Paraíba: present, no further details (Claps et al., 2001a)
Paraná: present, no further details (IIE, 1996a)
Pernambuco: present, no further details (Claps et al., 2001a)
Rio Grande do Norte: present, no further details (Claps et al., 2001a)
Rio Grande do Sul: present, no further details (Claps et al., 2001a)
Rio de Janeiro: present, no further details (Claps et al., 2001a)
Santa Catarina: present, no further details (IIE, 1996a)
Sao Paulo: present, no further details (Claps et al., 2001a)
Canada: absent, not established (Borchsenius, 1966; EPPO, 1999)
Ontario: absent, not established (EPPO, 1999)
Caribbean Islands: present, no further details (Davidson et al., 1983)
Central America: present, no further details (Davidson et al., 1983)
Colombia: present, no further details (IIE, 1996a; Kondo, 2001)
Costa Rica: present, no further details (IIE, 1996a)
Cuba: present, no further details (IIE, 1996a)
Curaçao: present, no further details (IIE, 1996a)
Dominica: present, no further details (IIE, 1996a)
Dominican Republic: present, no further details (IIE, 1996a)
Guadeloupe: present, no further details (IIE, 1996a)
Haiti: present, no further details (IIE, 1996a)
Honduras: present, no further details (IIE, 1996a)
Jamaica: present, no further details (IIE, 1996a)
Netherlands Antilles: present, no further details (EPPO, 1999)
Nevis: present, no further details (IIE, 1996a)
Panama: present, no further details (IIE, 1996a)
Peru: present, no further details (Nakahara, 1982; IIE, 1996a)
Puerto Rico: present, no further details (IIE, 1996a)
St Kitts: present, no further details (EPPO, 1999)
St Vincent: present, no further details (IIE, 1996a)
Suriname: present, no further details (IIE, 1996a)
Trinidad: present, no further details (IIE, 1996a)
Uruguay: present, no further details (IIE, 1996a)
USA: widespread (Borchsenius, 1966)
Alabama: present, no further details (Davidson et al., 1983)
Connecticut: present, no further details (Nakahara, 1982)
Delaware: present, no further details (Nakahara, 1982)
District of Colombia: present, no further details (Davidson et al., 1983)
Florida: present, no further details (Davidson et al., 1983)
Georgia (USA): present, no further details (Davidson et al., 1983)
Hawaii: present on Hawaii island only (Heu, 2002)
Louisiana: present, no further details (Davidson et al., 1983)
Maine: present, no further details (Nakahara, 1982)
Maryland: present, no further details (Davidson et al., 1983)
Mississippi: present, no further details (Davidson et al., 1983)
Missouri: present, no further details (IIE, 1996a)
Montana: present, no further details (Nakahara, 1982)
New Jersey: present, no further details (Nakahara, 1982)
New York: present, no further details (Nakahara, 1982)
North Carolina: present, no further details (Davidson et al., 1983)
Ohio: present, no further details (Nakahara, 1982)
Oregon: present, no further details (Nakahara, 1982)
Pennsylvania: present, no further details (Nakahara, 1982)
Rhode Island: present, no further details (Nakahara, 1982)
South Carolina: present, no further details (Davidson et al., 1983)
Tennessee: present, no further details (Davidson et al., 1983)
Texas: present, no further details (Davidson et al., 1983)
Virginia: present, no further details (Davidson et al., 1983; Kosztarab, 1996)
Washington: present, no further details (Kosztarab, 1996)
West Virginia: present, no further details (Kosztarab, 1996)
United States Virgin Islands: present, no further details (EPPO, 1999)
Venezuela: present, no further details (IIE, 1996a)
Virgin Is: present, no further details (IIE, 1996a)
Oceania
Australia: present, no further details (Davidson et al., 1983; Davidson and Miller, 1990)
New South Wales: present, no further details (IIE, 1996a)
Queensland: present, no further details (IIE, 1996a)
Belau: present, no further details (EPPO, 1999)
Bonin Is: present, no further details (Nakahara, 1982)
Caroline Islands: present, no further details (Beardsley, 1966)
Fiji: present (Williams and Watson, 1988; IIE, 1996a)
Guam: present, no further details (IIE, 1996a)
Micronesia, Federated States of: present, no further details (EPPO, 1999)
New Caledonia: present, no further details (Williams and Watson, 1988; IIE, 1996a)
New Guinea: present, no further details (Nakahara, 1982)
Norfolk Island: present, no further details (Williams and Watson, 1988; IIE, 1996a)
Northern Mariana Islands: present, no further details (IIE, 1996a)
Ogasawara-shoto: present, no further details (IIE, 1996a)
Palau: present (Nakahara, 1982; Beardsley, 1966)
Papua New Guinea: present (Williams and Watson, 1988; IIE, 1996a)
Pohnpei: present, no further details (Nakahara, 1982)
Saipan: present, no further details (Nakahara, 1982)
Samoa: present, no further details (Williams and Watson, 1988)
Solomon Islands: present (Williams and Watson, 1988; IIE, 1996a)
South Mariana Is: present (Beardsley, 1966)
Tonga: present (Williams and Watson, 1988; IIE, 1996a)
Truk Is: present (Nakahara, 1982; Beardsley, 1966)
Vanuatu: present (Williams and Watson, 1988; IIE, 1996a)
Wallis I.: present, no further details (Williams and Watson, 1988; IIE, 1996a)
Western Samoa: present (Williams and Watson, 1988)