Signoret, 1869
Diagnosis
Scale cover of adult female in life broadly oval to circular, 1.5-2.0 mm across, fairly flat, very thin, pale and translucent, with subcentral yellow exuviae ASPDEL6.jpg . The pale yellow exuviae are more or less central on the scale (Williams and Watson, 1988). Scale cover of adult male similar to but smaller than that of adult female, oval with subcentral yellow exuviae ASPDEL3.jpg . Exposed body of living adult female yellow, pyriform, 0.6-1.1 mm long. Adult male winged (Ghauri, 1962).
Body of slide-mounted adult female 0.7-1.2 mm long, pyriform and membranous ASPDEO.jpg . Prepygidial macroducts absent. Pygidial macroducts each more than 16x as long as wide. Dorsal seta situated at the outer corner of each median lobe slender, normally more than 1.4 times as long as a median lobe ASPDEP3.jpg . Median lobes much larger than other lobes, sometimes slightly recessed into the apex of the the pygidium. Plates extending beyond lobe apices, finely fringed; 6-10 (usually 8) plates present lateral to each third lobe, the outer 2 or 3 slightly simpler than the others.
Considerable variation occurs in the relative sizes of distinguishing features of A. destructor, such as median and second lobes, pygidial macroducts and dorsal setae (Williams and Watson, 1988).
Host range
Aspidiotus destructor is a highly polyphagous species. Davidson and Miller, 1990, recorded it from hosts belonging to75 genera in 44 plant families, but its host range is wider than this. Its hosts are typically perennial plants and include many fruit tree species, such as avocado, breadfruit, mango, guava and papaya. Coconut is its favourite host; the undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. Older trees (over 4 years) or trees on well-drained soil are seldom seriously infested. Hosts include species of: Actinidia, Albizia, Aleurites, Allamanda, Alpinia, Anacardium, Annona spp., Annona muricata, Artocarpus altilis, Attalea, Averrhoa, Barringtonia, Brassica, Cajanus, Calophyllum, Camellia sinensis, Camellia, Canna, Capsicum, Carica papaya, Cassia, Ceiba pentandra, Celtis, Chamaerops, Cinnamomum verum, Citrus, Cocos nucifera, Colocasia esculenta, Copernicia, Crotalaria, Cucumis, Cycas, Dillenia, Dioscorea, Diplothemium, Elaeis guineensis, Eucalyptus deglupta, Eugenia, Euphorbia, Ficus carica, Ficus spp., Guarea, Heliconia, Hevea brasiliensis, Hibiscus, Indigophora, Jasminum, Lantana, Laportea, Laurus nobilis, Licania, Litsea, Lycopersicon esculentum, Macaranga, Mangifera indica, Mollinedia, Musa, Myristica fragrans, Osmanthus, Palmae, Pandanus, Passiflora, Persea americana, Phoenix dactylifera, Phoenix, Physalis, Phytolacca, Piper spp., Plumeria, Prunus persica, Psidium guajava, Randia, Raphanus, Rhizophora, Ricinus communis, Rollinia, Saccharum officinarum, Solanum spp., Spondias, Syzygium aromaticum, Tamarindus, Theobroma cacao, Trachycarpus, Vigna unguiculata, Vitis vinifera, Washingtonia, Xanthosoma and Zingiber.
Affected plant stages: seedling, vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: leaves ASPDEL8.jpg , stems and fruits/pods ASPDEL2.jpg
Biology and ecology
Aspidiotus destructor reproduces sexually. Males locate unmated females by following pheromones released by them. The life cycle of A. destructor typically lasts for 32-34 days. In one study the life cycle was found to be 32 days for females and 27 days for males (Tabibullah and Gabriel, 1973; Taylor, 1935, also did an in-depth study).
Each female deposits 20-50 eggs under her scale cover over a few days. In China on Actinidia, the average number of eggs laid by one female was 32-42 (Zhou et al., 1993). At room temperature (26-28°C), the egg stage lasted for 5 days, the larval stage lasted 17 days, the pre-oviposition stage in adult females lasted 25 days, the female generation lasted 44 days and the male generation lasted 38 days (Zhou et al., 1993). In the Philippines, on coconut, the egg stage lasted for 8 days in both sexes (Tabibullah and Gabriel, 1973). After hatching, the nymphs crawl under the scale edge out into the open ASPDEL.jpg and colonize the undersides of leaves and tender shoots. They drop off the leaves easily, so mortality is high during heavy rain.
In China, A. destructor produced three generations annually, with the fertilized females overwintering on the stems of Actinidia trees (Zhou et al., 1993). In Japan on tea plants, A. destructor had only one generation per year (Murakami, 1970). However, in tropical conditions in Trinidad reproduction is continuous (Goberdhab, 1962).
The dispersal phase of A. destructor is the first instar, or crawler, 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 and transport of infested plant material by man.
Symptoms
On leaves, A. destructor causes yellow spots to develop beneath the insects ASPDEL7.jpg , due to the toxicity of saliva injected in to plant tissues while feeding ASPDEL5.jpg . Entire leaves may turn yellow to brown and fall, and fruits may be discoloured, stunted or fall prematurely. The bright yellow colour of affected coconut palms is clearly visible from a great distance. The undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. In extreme cases ASPDEL.jpg , the leaves dry out, entire fronds drop off and the crown dies.
Economic impact
Aspidiotus destructor is potentially the most destructive pest species on coconut, wherever it occurs in the world (Chua and Wood, 1990); the undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. In extreme cases, the leaves dry up, entire fronds drop off, the crown dies and the whole crop is lost. Neglected coconut plantations are particularly susceptible to damage by A. destructor. A. destructor is also an important economic pest of mango in Asia, Africa, the Philippines, India and Brazil; and of banana in Asia, the Pacific Islands, West Indies, Africa, Madagascar and South America. It attacks the leaves and fruits of oil palms, reducing the quality of the fruits (Chua and Wood, 1990). The species is also a pest of bananas worldwide (Chua and Wood, 1990). However, natural controls appear to keep A. destructor in check in most regions, and few major outbreaks have been recorded in recent years.
Before the introduction of successful biological control in 1955, copra production in Principe fell from 1400 to 500 tons per year owing to an invasion of A. destructor (Rosen, 1990a). After a heavy attack by A. destructor on coconuts in the Ivory Coast, yield was reduced by at least 25% over the next 2-3 years, although some heavily infested trees were able to catch up production in the 2 years after elimination of the infestation (Mariau and Julia, 1977).
Aspidiotus destructor is a cosmetic pest on a wide range of fruits, causing blemishes and other marks that reduce quality. On mango, A. destructor prefers grafted varieties; its economic impact is caused by feeding on tender shoots in nursery plants and because it adversely affects fruit setting in older plants. On oil palm, A. destructor is found feeding on leaves and fruit. It occasionally causes severe damage to guava in India (Hayes, 1970).
This species is highly polyphagous and therefore can easily be re-introduced, even if it is successfully controlled on the primary host crop.
Detection and inspection methods
Direct inspections can be made of the leaf undersides, aided by a x8 hand lens and a powerful torch if required, or by a dissection microscope in the laboratory. Look for yellow or brown patches on leaves, with scale insects on the underside. Look for pale buff, translucent, fairly flat, circular scales about 1.5-2.0 mm in diameter (the adult females).
Phytosanitary risk
Aspidiotus destructor is mentioned on quarantine lists (Burger and Ulenberg, 1990). Dharmaraju and Laird, 1984, described the transport of A. destructor around Oceania, mainly through human agency. They emphasized the importance of rigid quarantine procedures. The species is highly polyphagous and therefore can easily be re-introduced, even if it is successfully controlled on the primary host crop.
Natural enemies
Predators play a significant part in limiting A. destructor populations, and the most successful biological control of A. destructor has been achieved using predators rather than parasitoids (Rosen, 1990a). The commonest are the coccinellid beetles Chilocorus spp., Pseudoazya trinitatis, Cryptognatha nodiceps, Rhyzobius lophanthae and Pentilia castanea. The coccinellid Cryptognatha nodiceps is a particularly effective predator (Rosen, 1990a).
Parasites of local significance include species of Comperiella, Aphytis and Encarsia. A number of parasitoids and predators of A. destructor are described by Rosen, 1990a. The parasitoids of A. destructor in India are described by Tandon and Srivastava, 1980, and those in Pakistan by Rafiq Ahmad and Ghani, 1972. Natural enemies of A. destructor have been described from Sri Lanka (Sinnathamby, 1980), China (Zhou et al., 1993) and Taiwan (Wu and Tao, 1976).
Natural enemies of other scale insects may adapt to feeding on A. destructor as it colonizes new areas. However, continuous unchecked attacks of A. destructor were recorded on coconuts in Fiji, Mauritius and New Hebrides until natural enemies were introduced.
Narendra and Rao, 1974, describe an entomogenous fungus attacking A. destructor in India. Evans and Prior, 1990, list fungal pathogens of A. destructor.
The presence of ants on crop plants can deter natural enemies from attacking A. destructor. Rai, 1977, reported this problem on palms in Guyana; ants attending colonies of honeydew-producing mealybugs deterred natural enemies to such an extent that the mealybug and A. destructor populations reached damaging levels.
Parasitoids:
- Aphytis chionaspis, attacking: nymphs, adults, in Philippines, Taiwan, Malaysia, Sri Lanka, Indonesia. Introduced to: Fiji, Hawaii
- Aphytis lingnanensis, attacking: nymphs, in China, Taiwan and Japan. Introduced: USA (California), South Africa, Mexico, Argentina, Chile, Cyprus, Israel, Morocco
- Aphytis melinus, attacking: nymphs, in India, Pakistan. Introduced: USA (California), South Africa, Australia, Argentina, Chile, Cyprus, Israel, Italy, Morocco
- Comperiella bifasciata, attacking: nymphs, in India, China and Japan. Introduced to: USA (California), South Africa, Swaziland, Australia, Mexico, Agrentina, Israel.
- Comperiella unifasciata, attacking: adults, in Japan
- Encarsia citrina, attacking: nymphs, adults, in Japan. Introduced: Indonesia (Bali), Tahiti, Fiji, Cook Islands, Africa
- Signiphora fax
- Zaomma lambinus, attacking: adults, in Java
Predators:
- Aleurodothrips fasciapennis, attacking: eggs, nymphs, adults, in Indonesia; introduced to: Fiji
- Chilocorus circumdatus, attacking: nymphs, adults
- Chilocorus dohrni, attacking: eggs, nymphs, adults, in Ivory Coast
- Chilocorus kuwanae, attacking: nymphs, adults in China, Russian far east, Korea and Japan. Introduced to: Caucasus, Italy, India and USA
- Chilocorus nigrita, attacking: eggs, nymphs, adults, in Oriental region, Indonesia. Introduced to: southern Africa, Oman, Seychelles, American Samoa, Hawaii, Pohnpei and Mauritius and has spread naturally
- Chilocorus politus, attacking: nymphs, adults
- Chilocorus schioedtii, attacking: eggs, nymphs, adults, in Ivory Coast
- Cryptognatha nodiceps, attacking: eggs, nymphs, adults, in Trinidad. Introduced to: Fiji, Jamaica, St Kitts, Angola, Principe
- Cybocephalus gibbulus, attacking: eggs, larvae, adults, in Taiwan
- Pentilia castanea
- Pharoscymnus c-luteum, attacking: nymphs, adults
- Pseudoazya trinitatis, attacking: eggs, nymphs, adults, in Puerto Rico, Trinidad. Introduced to: Saipan, USA (Florida), Jamaica, St Lucia
- Pseudoscymnus anomalus, attacking: eggs, nymphs, adults, in Guam, Palau, Truk, Vanuatu. Introduced to: Hawaii
- Pseudoscymnus cadapani, attacking: eggs, larvae, adults, in Philippines
- Rhyzobius lophanthae, attacking: eggs, nymphs, adults, in Australia. Introduced to: Hawaii, Republic of Georgia
- Rhyzobius pulchellus, attacking: eggs, nymphs, adults, in New Caledonia, Vanuatu. Introduced to: Fiji
- Rhyzobius satelles, attacking: eggs, nymphs, adults, in Australia. Introduced to: Vanuatu, Wallis
- Rodolia rubea, attacking: nymphs, adults
- Scymnus luteus, attacking: eggs, larvae, adults, in Indonesia, Pacific Islands
- Scymnus luteus, attacking: eggs, larvae, adults, in Indonesia, Pacific Islands
- Scymnus severini, attacking: nymphs, adults
- Telsimia nitida, attacking: eggs, larvae, adults, in Taiwan, Hawaii, Indonesia
- Telsimia sanchezi, attacking: eggs, larvae, adults, in Philippines
Distribution
See Aspidiotus destructor distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species.
Aspidiotus cryptomeriae Kuwana ASPCRYS.jpg is similar to A. destructor; however, the marginal setae by the outer corners of the median lobes are shorter than the lobes in A. cryptomeriae ASPCRYP.jpg, whereas they are longer than the lobes in A. destructor ASPDEP2.jpg. Aspidiotus cryptomeriae ASPCRYL1.jpg is known from USA (Connecticut, Delaware, Indiana, Maryland, New York and Pennsylvania), former USSR (Far East), China (Fujian, Shandong, Yunnan, Zhejiang), Japan (Honshu), Korea, Saghalien and Taiwan on the needles/leaf undersides of conifers such as species of Abies, Cedrus, Cephalotaxus, Chamaecyparis, Cryptomeria, Juniperus, Keteleeria, Picea, Pinus, Pseudotsuga, Taxus and Torreya (Takagi, 1969; 1972 data from S. Kawai photograph; Kawai, 1980; Nakahara, 1982; Zahradník, 1990; Kosztarab, 1996; Tao, 1999). The species ASPCRYL2.jpg is univoltine and overwinters as second instars; it has been recorded damaging hosts in Sakhalin (Zahradník, 1990), and is an important pest of conifers in Japan (Tachikawa, 1982). Gardosik, 2001, discusses its life history, damage and control. In Maryland it has two generations per year and is sufficiently damaging to shade and roadside trees to warrant control measures (Smith and Raupp, 1986; Kosztarab, 1996). The immature stages and natural enemies were studied in detail by Stoetzel and Davidson, 1974. ASPCRYL.jpg
Aspidiotus destructor subspecies rigidus Reyne was described from Indonesia (Sulawesi), damaging Cocos nucifera (Reyne, 1948). This subspecies can only be separated reliably from A. destructor sensu stricto by details of its biology, but the prosomal cuticle is rather thicker than that in A. destructor sensu stricto.
In life, the appearance of A. destructor and A. excisus Green is very similar. In A. excisus ASPEXS.jpg, the median lobes are strongly recessed into the pygidial margin; short marginal prepygidial macroducts are present on either side of the abdomen; the dorsal seta situated at the outer corner of each median lobe is shorter than the median lobe, and the dorsal pygidial macrocucts are no more than 8x as long as wide ASPEXP.jpg. In A. destructor, the median lobes are never strongly recessed; dorsal pygidial macrocucts are at least 16x as long as wide; the dorsal seta situated at the outer corner of each median lobe is normally more than 1.4 times as long as a median lobe, and short marginal prepygidial macroducts are absent ASPDEP2.jpg. Williams and Watson, 1988, provide a key and give detailed descriptions and illustrations of the adult females of both species. Aspidiotus excisus has been recorded from Kenya, Pakistan, India (Lakshaweep Is, Tamil Nadu), Bangladesh, Sri Lanka, Maldive Is, West Malaysia, Singapore, North Borneo, Thailand, Indonesia, China (Hong Kong), Taiwan, Japan, Philippines, Papua New Guinea, Mariana Is, Palau, Caroline Is, Fiji, Costa Rica, El Salvador, Honduras, Mexico, Panama, Colombia, Ecuador, Guatemala, Guyana, Suriname, Venezuela, Antigua, Dominican Republic, Grenada, Jamaica, Martinique, Puerto Rico, St Lucia, Trinidad and USA (Florida), on hosts in 11 plant families: Araceae, Caricaceae, Commelinaceae, Compositae, Convolvulaceae, Labiatae, Musaceae, Oleaceae, Orchidaceae, Rutaceae and Verbenaceae including species of Carica, Citrus, Cyanotis, Euphorbia and Thespesia (Beardsley, 1966; Velasquez, 1971; Kawai, 1980; Nakahara, 1982; Williams and Watson, 1988; Miller, 1996; Kondo, 2001; The Natural History Museum collection, London, UK). ASPEXL1.jpg The adult male is winged (Ghauri, 1962) ASPEXL2.jpg and ASPEXL3.jpg
Comments
Aspidiotus destructor apparently originated in the Pacific islands (Burger and Ulenberg, 1990) but is now recorded in tropical and subtropical regions worldwide. It is present in nearly all countries where coconuts are grown. In the northern parts of its range, it is found only under glass (Danzig and Pellizzari, 1998).
Europe
Former USSR:
Azerbaijan: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Central European territory: present under glass, no further details (Danzig and Pellizzari, 1998)
Georgia, Republic of: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Russian Federation: present, no further details (CIE, 1966)
South European territory: present under glass, no further details (Danzig and Pellizzari, 1998)
Transcaucasus: present, no further details (CIE, 1966)
Germany: present under glass, no further details (Danzig and Pellizzari, 1998)
Italy: under glass (Longo et al., 1995)
Portugal
Madeira: present, no further details (CIE, 1966)
Spain
Canary Islands: present, no further details (CIE, 1966)
United Kingdom
England: under glass at a few botanic gardens (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
Asia
Bangladesh: present, no further details (APPPC, 1987)
Bhutan: The Natural History Museum collection, London, UK
Brunei Darussalam: present, no further details (Waterhouse, 1993)
Cambodia: present, no further details (CIE, 1966; Waterhouse, 1993)
Chagos Archipelago: present, no further details (CIE, 1966)
China
Fujian: present, no further details (Tao, 1999)
Guangdong: present, no further details (Tao, 1999)
Guangxi: present, no further details (Tao, 1999)
Hainan: present, no further details (CIE, 1966)
Hubei: present, no further details (Tao, 1999)
Hunnan: present, no further details (Tao, 1999)
Hong Kong: present, no further details (CIE, 1966)
Hunan: present, no further details (Zhou et al., 1993)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Quichou: present, no further details (Tao, 1999)
Shandong: present, no further details (Tao, 1999)
Sichuan: present, no further details (Tao, 1999)
Zhejiang: present, no further details (Tao, 1999)
India
Andaman and Nicobar Islands: The Natural History Museum collection, London, UK
Andhra Pradesh: present, no further details (CIE, 1966)
Assam: present, no further details (CIE, 1966)
Bihar: present, no further details (CIE, 1966)
Gujarat: present, no further details (CIE, 1966)
Jammu and Kashmir: present, no further details (CIE, 1966)
Karnataka: present, no further details (CIE, 1966)
Kerala: present, no further details (CIE, 1966)
Lakshadweep Is: present, no further details (CIE, 1966)
Madhya Pradesh: present, no further details (CIE, 1966)
Maharashtra: present, no further details (CIE, 1966)
Orissa: present, no further details (CIE, 1966)
Punjab: present, no further details (CIE, 1966)
Sikkim: present, no further details (CIE, 1966)
Tamil Nadu: present, no further details (CIE, 1966)
Uttar Pradesh: present, no further details (CIE, 1966)
West Bengal: present, no further details (CIE, 1966)
Indonesia: present, no further details (Waterhouse, 1993)
Irian Jaya: present, no further details (Williams and Watson, 1988)
Bali: present, no further details (CIE, 1966)
Java: present, no further details (CIE, 1966)
Sumatra: present, no further details (CIE, 1966)
Sulawesi: present, no further details (CIE, 1966)
Iran: present, no further details (Danzig and Pellizzari, 1998)
Japan: present, cannot read any further details (Kawai, 1980)
Honshu: present, no further details (CIE, 1966)
Korea: present, no further details (Danzig and Pellizzari, 1998)
Malaysia: present, no further details (Waterhouse, 1993)
West Malaysia: present, no further details (CIE, 1966)
Sabah: present, no further details (CIE, 1966)
Sarawak: present, no further details (CIE, 1966)
Maldive Is: present, no further details (Watson et al., 1995)
Myanmar: present, no further details (CIE, 1966; Waterhouse, 1993)
Nepal: The Natural History Museum collection, London, UK
Oman: widespread (Kinawy, 1991)
Pakistan: present, no further details (CIE, 1966)
Philippines: present, no further details (Velasquez, 1971; Waterhouse, 1993)
Saudi Arabia: present, no further details (Danzig and Pellizzari, 1998)
Singapore: present, no further details (APPPC, 1987; Waterhouse, 1993)
Sri Lanka: present, no further details (CIE, 1966)
Taiwan: present, no further details (Takagi, 1969; Wong et al., 1999)
Thailand: present, no further details (APPPC, 1987; Waterhouse, 1993)
Vietnam: present, no further details (CIE, 1966; Waterhouse, 1993)
Yemen: The Natural History Museum collection, London, UK
Africa
Angola: present, no further details (CIE, 1966)
Benin: present, no further details (CIE, 1966)
Burundi: present, no further details (CIE, 1966)
Cameroon: present, no further details (CIE, 1966)
Cape Verde: present, no further details (CIE, 1966)
Congo Democratic Republic: present, no further details (Buyckx, 1962; CIE, 1966)
Côte d'Ivoire: present, no further details (CIE, 1966)
Egypt: present, no further details (Danzig and Pellizzari, 1998)
Eritrea: present, no further details (CIE, 1966)
Ethiopia: present, no further details (CIE, 1966)
Ghana: present, no further details (CIE, 1966)
Guinea-Bissau: present, no further details (CIE, 1966)
Guinea: present, no further details (CIE, 1966)
Kenya: present, no further details (CIE, 1966)
Madagascar: present, no further details (CIE, 1966)
Mauritania: present, no further details (CIE, 1966)
Mauritius: present, no further details (Williams and Williams, 1988)
Mozambique: present, no further details (CIE, 1966)
Nigeria: present, no further details (CIE, 1966)
Principe: present, no further details (CIE, 1966)
Rwanda: present, no further details (CIE, 1966)
Réunion: present, no further details (Williams and Williams, 1988)
Sao Tomé: present, no further details (CIE, 1966; Fernandes, 1974)
Senegal: present, no further details (CIE, 1966)
Seychelles: present, no further details (CIE, 1966)
Sierra Leone: present, no further details (CIE, 1966)
Somalia: present, no further details (CIE, 1966)
South Africa: present, no further details (CIE, 1966)
Sudan: present, no further details (CIE, 1966)
Tanzania: present, no further details (CIE, 1966)
Zanzibar: present, no further details (CIE, 1966)
Togo: present, no further details (CIE, 1966)
Uganda: present, no further details (CIE, 1966)
Zambia: present, no further details (CIE, 1966)
Zimbabwe: The Natural History Museum collection, London, UK
Western Hemisphere
Antigua: present, no further details (CIE, 1966)
Bahamas: The Natural History Museum collection, London, UK
Barbados: present, no further details (Bennett and Alam, 1985)
Belize: present, no further details (CIE, 1966)
Brazil:
Alagôas: present, no further details (Foldi, 1988)
Amazonas: present, no further details (Foldi, 1988; Claps et al., 2001a)
Bahia: present, pest on cocoa (Silva, 1950; Claps et al., 2001a)
Ceará: present, no further details (CIE, 1966; Claps et al., 2001a)
Fernando Noronha: present, no further details (Foldi, 1988; Claps et al., 2001a)
Guanabara: present, no further details (Foldi, 1988)
Maranhao: present, no further details (Foldi, 1988; Claps et al., 2001a)
Pará: present, no further details (Foldi, 1988; Claps et al., 2001a)
Paraíba: present, no further details (Foldi, 1988; Claps et al., 2001a)
Pernambuco: present, no further details (Foldi, 1988; Claps et al., 2001a)
Piaui: present, no further details (Foldi, 1988; Claps et al., 2001a)
Rio Grande do Norte: present, no further details (Foldi, 1988; Claps et al., 2001a)
Rio de Janeiro: present, no further details (Foldi, 1988; Claps et al., 2001a)
Santa Caterina: present, no further details (Foldi, 1988; Claps et al., 2001a)
Sao Paulo: present, no further details (Foldi, 1988; Claps et al., 2001a)
Sergipe: present, no further details (CIE, 1966)
Cayman Islands: present (CIE, 1966)
Chile
Easter Island: present, no further details (Charlín, 1973; Claps et al., 2001a)
Colombia: present, no further details (CIE, 1966; Kondo, 2001)
Costa Rica: present, no further details (CIE, 1966)
Cuba: present, no further details (CIE, 1966)
Dominica: present, no further details (CIE, 1966)
Dominican Republic: present, no further details (CIE, 1966)
Ecuador: present, no further details (CIE, 1966; Kondo, 2001)
Galapagos Is: present, no further details (CIE, 1966)
Grenada: present, no further details (CIE, 1966)
Guadeloupe: present, no further details (CIE, 1966)
Guatemala: present, no further details (CIE, 1966)
Guyana: present, no further details (CIE, 1966)
Haiti: present, no further details (CIE, 1966)
Honduras: present, no further details (CIE, 1966)
Jamaica: The Natural History Museum collection, London, UK
Lesser Antilles: present, no further details (Kondo, 2001)
Martinique: present, no further details (CIE, 1966)
Mexico: present, no further details (CIE, 1966; Miller, 1996)
Montserrat: present, no further details (CIE, 1966)
Netherlands Antilles: present, no further details (CIE, 1966)
Nevis: The Natural History Museum collection, London, UK
Nicaragua: present, no further details (CIE, 1966)
Panama: present, no further details (CIE, 1966)
Peru: present, no further details (CIE, 1966)
Puerto Rico: present, no further details (CIE, 1966)
St Kitts and Nevis: present, no further details (CIE, 1966)
St Lucia: present, no further details (CIE, 1966)
St Vincent: present, no further details (CIE, 1966)
Suriname: present, no further details (CIE, 1966)
Trinidad: present, no further details (CIE, 1966)
USA
Connecticut: under glass (Nakahara, 1982)
Florida: present, no further details (CIE, 1966)
Georgia: present, no further details (Nakahara, 1982)
Hawaii: present on Oahu, Hawaii, Maui and Kauai (Heu, 2002)
Pennsylvania: present, no further details (Nakahara, 1982)
United States Virgin Islands: present, no further details (CIE, 1966)
Venezuela: present, no further details (CIE, 1966)
Virgin Is: present, no further details (CIE, 1966; Williams and Greathead, 1990)
Oceania
American Samoa: present (Williams and Watson, 1988)
Australia
Northern Territory: present, no further details (CIE, 1966; CSIRO, 2001)
Queensland: present, no further details (CSIRO, 2001)
Belau: present, no further details (CIE, 1966)
Bonin Is: present, no further details (Nakahara, 1982)
Caroline Islands: present, no further details (CIE, 1966)
Fiji: present, widespread (Williams and Watson, 1988)
French Polynesia: present, widespread (Williams and Watson, 1988)
Guam: present, no further details (CIE, 1966)
Marquesas Is: present, no further details (CIE, 1966)
Mariana Is: present (Nakahara, 1982)
Marshall Islands: present, no further details (CIE, 1966)
New Britain: present, no further details (CIE, 1966)
New Caledonia: present, no further details (CIE, 1966)
Palau: present, no further details (Beardsley, 1966)
Papua New Guinea: present (Williams and Watson, 1988)
Pohnpei: present, no further details (Beardsley, 1966)
Samoa: present, no further details (Nakahara, 1982)
Society Is: present, no further details (Nakahara, 1982)
Solomon Islands: present, widespread (Williams and Watson, 1988)
South Mariana Is: present (Beardsley, 1966)
Tahiti: present, no further details (CIE, 1966)
Truk Is: present, no further details (Beardsley, 1966)
Tuamotu Is: present, no further details (Nakahara, 1982)
Tuvalu: present, no further details (CABI, 2000)
Vanuatu: present, widespread (Williams and Watson, 1988)
Wallis Is: present, no further details (CIE, 1966)
Western Samoa: present, widespread (Williams and Watson, 1988)
Yap Is: present, no further details (Beardsley, 1966)