Bouché, 1833
Taxonomic notes
When Signoret, 1868, attributed the authorship of Aspidiotus hederae (Vallot), Chermes hederae (Vallot), Aspidiotus capparis (Vallot) and Aspidiotus ? osmanthis (Vallot) to Vallot, 1829, he initiated several nomenclatural artifacts. Subsequent authors have perpetuated these errors. These names are not available according to the International Code of Zoological Nomenclature, and their usage should be excluded (Ben-Dov and Matile-Ferrero, 1999). The valid description of Aspidiotus hederae by Signoret, 1869b, which name he erroneously attributed to Vallot, 1829, refers to another species, not A. nerii Bouché (Ben-Dov and Matile-Ferrero, 1999).
Diagnosis
Scale cover of adult female 1.5-2.0 mm in diameter, nearly circular, flat to convex, semitranslucent yellowish white to light tan; with yellow or gold, central to subcentral exuviae ASPNEL3.jpg . Scale cover of male white, oval and translucent ASPNEL.jpg . Exposed body of adult female bright yellow in life (Gill, 1997). Adult males are winged (Ghauri, 1962; Kosztarab, 1996) ASPNEMAL.jpg .
Slide-mounted adult female pyriform and membranous ASPNES.jpg . Short, 1-barred submarginal and marginal macroducts (less than six times as long as they are wide) are present on both the pygidium and prepygidial segments. Pygidium lacking paraphyses at the bases of the lobes, although there is a basal sclerosis associated with each median lobe; the median lobes are never recessed, and the third lobes are sometimes pointed ASPNEP.jpg . Plates finely fringed, longer than lobes, especially near third lobe; lateral to the third lobes they become simpler and shorter with increasing distance from the lobe ASPNEMG.jpg .
The first instar nymph was described and illustrated by Howell and Tippins, 1977.
Host range
Aspidiotus nerii is a highly polyphagous species that has been recorded on hundreds of host species in over 100 plant families. Its many hosts include agricultural crops, palms, cut flowers and woody ornamentals (Williams and Watson, 1988; Flint, 1990; Zahradník, 1990; Dreistadt, 1994; Dymock and Holder, 1996; Heu, 2002). Hosts include species of: Acacia spp., Acer, Actinidia chinensis, Agave, Albizia, Aleurites, Aloe, Ananas comosus, Antirrhinum, Aralia, Arbutus, Areca, Artocarpus, Asparagus spp., Asplenium, Azalea, Bauhinia, Beilschmiedia, Brassica, Buxus, Callitris, Camellia, Capparis, Carica, Caryopteris, Cassia, Castela, Catalpa, Cedrela, Cedrus, Celtis, Ceratonia, Cercestis, Chrysanthemum, Citrus spp., Clerodendrum, Cocos nucifera, Cordyline, Crataegus, Cucurbita, Cupressus, Cyanotis, Cycas, Cyperus, Cytisus, Daphne, Datura, Dianthus, Diospyros, Dracaena, Elaeagnus, Elaeis, Elephantopus, Erica, Eucalyptus, Euonymus, Euphorbia, Fatsia, Ficus, Fraxinus, Gardenia, Genista, Grevillea, Hakea, Hedera, Helianthemum, Howea, Hoya, Hydrangea, Ilex, Ipomoea, Iris, Jasminum, Juniperus, Labiatae, Lapageria, Laurus nobilis, Leguminosae, Ligustrum, Lilaea, Lonicera, Lycopersicon, Macadamia ternifolia, Magnolia, Malus sylvestris, Mangifera indica, Medicago, Melia, Mentha, Morus, Muehlenbeckia, Musa, Myosporum, Myrsine, Nectandra, Nerium, Nicotiana, Oleaceae, Olea europaea, Opuntia, Orchidaceae, Palmae, Pandanus, Parkinsonia, Passiflora, Persea americana, Peumus, Phoenix dactylifera, Phormium, Physalis, Pinus, Piper, Pistacia vera, Pittosporum, Platanus, Plumeria, Poncirus, Populus, Prosopis, Prunus spp., Pteridophyta, Punica, Pyrus communis, Quercus, Rhododendron, Rhus, Ribes, Ricinus, Robinia, Rosa, Rosaceae, Rosmarinus, Rubia, Ruscus, Ruta, Salix, Salvia, Schinus, Sechium, Senna, Simmondsia, Smilax, Solanum, Sophora, Spartium, Strelitzia, Syringa, Tagetes, Taxus, Thuja, Tmesipteris, Tournefortia, Trachycarpus, Ulex, Ulmus, Veronica, Vinca, Vitis vinifera, Vitex, Vriesea, Washingtonia, Yucca, Zamia, Zanthoxylum and Ziziphus.
Affected plant stages: seedling, vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: on aerial parts - leaves ASPNEL2.jpg , bark and fruit
Biology and ecology
Reproduction is either sexual or pathenogenetic; in California, both parthenogenetic and bisexual races of A. nerii occur. Studies by Gerson and Hazan, 1979, indicate that there may be six different parthenogenetic populations in various parts of the world. They found that the parthenogenetic strains reproduced more slowly that the sexual population, and were less fecund. Females lay eggs for 1-2 weeks, averaging a total of ca 100-150 eggs from each female. Development time is about 5 weeks, with 2-3 generations produced each year depending on climatic conditions. In olives, infestations are often heavier on the lower branches, which may indicate that the insects prefer cooler, shady conditions.
Crawlers are the primary dispersal stage and move to new areas of the plant or are dispersed by wind or animal contact. Mortality due to abiotic factors is high in this stage. Dispersal of sessile adults and eggs occurs through human transport of infested plant material.
Symptoms
Symptoms are generally related to the severity of attack. Infestations on the leaves and stems may cause wilting and reduce the photosynthetic area of the plants, leading to lower yield. Damage to fruit occurs in heavy infestations, where spotting and often deformity of fruits affects market value. In olives this is seen as green spots on purple fruits. Damage to foliage is particularly important in ornamentals. Heavy infestation can lead to yellowing, leaf and shoot malformation or shriveling and plant death (Beardsley and Gonzalez, 1975; Debach and Rosen, 1991; Dreistadt et al., 1994).
Economic impact
Aspidiotus nerii is particularly important where aesthetic value of the crop is high, for example in cut flowers, palms and ornamentals (VanDriesche et al., 1998; Zahradník, 1990). In olive crops, the presence of a single scale makes a fruit unmarketable. Economic loss on table olives due to damage to fruits and reduced oil yield can be up to 70% (Alexandrakis and Bénassy, 1981; Flint, 1990). Large infestations can develop on olives, reducing oil content and discolouring and deforming fruit, which has to be culled; A. nerii can also be a pest on Citrus and carob (Gill, 1997). It can be an economically important pest of forest trees (Zahradník, 1990a). The species is a pest of olive in Peru (Canales Canales and Valdivieso, 1999), Argentina, Chile, and all Mediterranean countries (Argyriou, 1990; Claps et al., 2001a). Danzig and Pellizzari, 1998, describe A. nerii as a very dangerous pest in the Palaearctic region, and Foldi, 2001, lists it as an occasional pest in France. Quiroga et al., 1991, reported that A. nerii was the most severe pest of jojoba (Simmondsia chinensis) fruits in northern and central Chile. Heavy infestation on a single host caused distortion, discoloration, wilting and eventually host death. Aspidiotus nerii is a pest of bananas in the Canary Is (Chua and Wood, 1990). Gill, 1997, described A. nerii as a troublesome pest of ornamental plants in gardens, nurseries and under glass/indoors in California. Charles and Henderson, submitted, record it as a pest of economically important plants in New Zealand, and that it occurs on native plants also.
Detection and inspection methods
Closely examine leaves, bark and fruit o fthe host-plants listed above, for almost circular, flat to convex, semitranslucent yellowish white to light tan scale covers, each with yellow or gold, central to subcentral exuviae. In olives, infestations are often heavier on the lower branches.
Phytosanitary risk
Infested plant material is mainly responsible for passive transport of this insect.
Natural enemies
Parasitoids:
- Aphytis spp.
- Aphytis chilensis, attacking: nymphs, adults, in Europe; Middle East; North and South Africa; North and South America; USA, California; Australia; New Zealand
- Aphytis chionaspis, attacking: nymphs, adults
- Aphytis coheni, attacking: nymphs, adults, in Greece
- Aphytis dubius ssp. intermedius in Argentina
- Aphytis lingnanensis, attacking: nymphs, adults, in China; introduced: Greece; Mexico; USA (California, Texas)
- Aphytis melinus, attacking: nymphs, adults, in India, Pakistan. Introduced: USA (California), South Africa, Australia, Argentina, Chile, Spain, Greece, Cyprus, Turkey, Israel, Italy, Morocco
- Encarsia aurantii in South America
- Encarsia citrina, attacking: nymphs, adults, in Tahiti; Fiji; Indonesia, Bali, Chile
- Encarsia ectophaga in Argentina
- Habrolepis rouxi
- Mytilaspidis sp. in Chile
- Signiphora merceti in Argentina
- Signiphora xanthographa in Argentina
Predators:
- Aleurodothrips fasciapennis, attacking: eggs, nymphs, adults, in Indonesia; introduced to: Fiji
- Chilocorus circumdatus, attacking: nymphs, adults
- Chilocorus infernalis, attacking: nymphs, adults
- Chilocorus kuwanae, attacking: nymphs, adults, in China, Korea
- Chilocorus nigrita, attacking: nymphs, adults, in India; Pakistan; Sri Lanka; Introduced: Africa (accidental); Oman
- Chilocorus renipustulatus, attacking: nymphs, adults
- Hemisarcoptes coccophagus, attacking all stages apart from eggs, widespread including Israel; Introduced: New Zealand
- Karnyothrips flavipes
- Hemisarcoptes malus, attacking all stages apart from eggs
- Rhyzobius lophanthae, attacking: nymphs, adults
- Rhyzobius pulchellus, attacking: nymphs, adults, in Vanuatu, New Caledonia
- Signiphora spp.
- Signiphora flavopalliata
Distribution
See Aspidiotus nerii distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species.
Aspidiotus elaeidis Marchal resembles A. nerii in possessing short prepygidial macroducts; however, in A. elaeidis, the third lobes are rounded, the anus is wider and longer than a median lobe, and the dorsal pygidial macroducts are 14x as long as wide ASPELP.jpg. In A. nerii, the third lobes are sometimes pointed; the anus is usually the same size as, or smaller than, a median lobe; and the dorsal pygidial macroducts are up to 6x as long as wide ASPNEP.jpg. Aspidiotus elaeidis has been recorded from Denmark (under glass) and Central and southern Africa, e.g. Angola, Benin, Cameroon, Congo, Côte d'Ivoire, Eritrea, Ethiopia, Ghana, Nigeria, Senegal, Somalia, Sierra Leone and Tanzania; on the leaves of hosts in 9 plant families: Apocynaceae, Ebenaceae, Euphorbiaceae, Loranthaceae, Meliaceae, Ochnaceae, Oleaceae, Palmae and Rosaceae, such as species of Bridelia, Carissa, Chaetachme, Chrysobalanus, Cocos, Diospyros, Elaeis, Funtumia, Lophira, Manihot, Musa, Olea, Persea, Viscum and Xylocarpus (Balachowsky, 1956; Munting, 1971; Danzig and Pellizzari, 1998; The Natural History Museum collection, London, UK). Aspidiotus elaeidis attacks the leaves and fruits of oil palm in Congo (Chua and Wood, 1990) and Benin (Munting, 1971).
Aspidiotus excisus Green ASPEXS.jpg and A. nerii are similar in possessing marginal prepygidial ducts and short pygidial macroducts. However, in A. excisus the median lobes are strongly recessed into the pygidial margin ASPEXP.jpg. In A. nerii the median lobes are never strongly recessed ASPNEP.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 (under glass in cold areas) from Kenya, Pakistan, India (Lakshaweep Is, Madras, 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 leaves and stems of hosts in 11 plant families: Araceae, Caricaceae, Commelinaceae, Compositae, Convolvulaceae, Labiatae, Musaceae, Oleaceae, Orchidaceae, Rutaceae and Verbenaceae including species of Aglaonema, Carica, Citrus, Cyanotis, Euphorbia, Hoya, Lonicera and Thespesia (Rao and Kumar, 1952; Beardsley, 1966; Velasquez, 1971; Kawai, 1980; Nakahara, 1982; Williams and Watson, 1988; Davidson and Miller, 1990; Watson et al., 1995; Miller, 1996; Kondo, 2001; The Natural History Museum collection, London, UK). This species is a pest of bananas in Fiji (Chua and Wood, 1990). Scale cover of adult female circular or uneven, translucent white, slightly convex with subcentral yellow exuviae; scale cover of male similar to that of female but smaller and oval (Davidson and Miller, 1990). ASPEXL1.jpg The adult male is winged (Ghauri, 1962) ASPEXL2.jpg and ASPEXL3.jpg .
Oceanaspidiotus spinosus (Comstock) is a tropical species not included in the key, which could be mistaken for A. nerii OCSPIS.jpg . However, O. spinosus has dorsal macroducts about 15x as long as wide, with sclerotized orifices OCSPIP.jpg . In A. nerii, the dorsal macroducts are up to 6x as long as wide, and their orifices are not obviously sclerotized ASPNEP.jpg. Oceanaspidiotus spinosus OCSPINL2.jpg has been recorded from ?United Kingdom (England, under glass - but there have been no recent records), former USSR (Georgia; Russia (St Petersburg, under glass)), Germany, Italy, Georgia, Turkey, Spain including Canary Is., Algeria, Madagascar, Mozambique, South Africa, Tanzania, India, Israel, Nepal, Syria, China (Guangdong, Jiangsu), Japan (Kyushu, Ryukyu), Bahamas, Bermuda, Cuba, Puerto Rico, Mexico, USA (Alabama, California (but not collected for many years), District of Colombia (under glass), Florida, Georgia, Hawaii, Louisiana, Mississippi, New York, Texas, Virginia), Mexico, Bahamas, Bermuda, Brazil, Central America, Colombia, Costa Rica, Cuba, Dominican Republic, Argentina, Peru, Puerto Rico, Uruguay, Cook Is and Yap Is, on leaves, twigs or bark of hosts from at least 19 plant families: Asclepiadaceae, Cactaceae, Caprifoliaceae, Celastraceae, Cycadaceae, Euphorbiceae, Lauraceae, Liliaceae, Malvaceae, Moraceae, Magnoliaceae, Orchidaceae, Palmae, Platanaceae, Rosaceae, Sapindaceae, Taxaceae, Theaceae, Taxaceae and Vitaceae including species of Aleurites, Asparagus, Camellia, Cinnamomum, Dendrobium, Echinopsis, Euonymus, Euphorbia, Ficus, Harrisia, Laurus, Livistona, Magnolia, Mangifera indica, Nephelium, Opuntia, Palmae, Persea americana, Platanus, Rosa, Rubus, Smilax, Stapelia, Taxus, Trachycarpus, Viburnum and Vitis vinifera (Ferris, 1941a; Beardsley, 1966; Gerson and Tsur, 1970; 1976 and 1999 data from S. Kawai photographs; Nakahara, 1982; Takagi, 1984; Williams and Watson, 1988; Hodgson and Hilburn, 1991; Longo et al., 1995; Miller, 1996; Tao, 1999; Claps et al., 2001a; Kondo, 2001). Palms are preferred hosts. There are no records of this species causing damage, but heavy infestations can develop OCSPINL1.jpg . See also genus Oceanaspidiotus, kingdom Animalia.
Oceanaspidiotus pangoensis (Doane and Ferris) OCPAS.jpg resembles O. spinosus, but differs in having the margins of the prosoma sclerotized in mature specimens, the anus situated near the middle of the pygidium, and only 0-8 perivulvar pores OCPAP.jpg; whereas in O. spinosus and Aspidiotus nerii, the prosoma remains membranous, the anus is situted closer to the posterior margin than to the middle of the pygidium, and at least 10 perivulvar pores are present. Oceanaspidiotus pangoensis is known from American Samoa, Fiji, Niue, Tonga and Western Samoa on species of Broussonetia, Bruguiera, Cocos nucifera, Cordia, Hernandia and Smilax (Reddy, 1970; Williams and Watson, 1988).
In Israel, Gerson and Hazan, 1979, described the parthenogenetic population on Pittosporum undulatum as a new species, A. paranerii Gerson and Hazan, but this can only be identified by a detailed life history study. Aspidiotus paranerii reproduced more slowly than the sexual population of A. nerii, and was less fecund.
Comments
DeBach and Rosen, 1991, considered Aspidiotus nerii to be native to the Mediterranean area. It now has a worldwide distribution, especially in the tropical and subtropical zones (Zahradník, 1990a) and occurs under glass in cold areas (Davidson and Miller, 1990). The wide geographical distribution of this pest is primarily due to the activities of man.
Europe
Belgium: The Natural History Museum collection, London, UK
Bulgaria: present, no further details (CIE, 1970)
Cyprus: present, no further details (Nakahara, 1982)
Europe: widespread (under glass in cold areas) (Nakahara, 1982)
Former USSR: present, no further details (CIE, 1970)
Armenia: under glass (Nakahara, 1982)
Azerbaijan: under glass (Nakahara, 1982)
Crimea: under glass (Nakahara, 1982)
Georgia, Republic of: present, no further details (CIE, 1970)
Krasnodar Territory: under glass (Nakahara, 1982)
Former Yugoslavia: present, no further details (CIE, 1970)
France: present, no further details (CIE, 1970; Foldi, 2001)
Corsica: present, no further details (CIE, 1970)
Gibraltar: present, no further details (CIE, 1970)
Greece: present, no further details (CIE, 1970; Alexandrakis and Bénassy, 1981)
Crete: present, no further details (Alexandrakis and Bénassy, 1981)
Italy: present, no further details (Longo et al., 1995)
Sardinia: present, no further details (Longo et al., 1995)
Sicily: present, no further details (Longo et al., 1995)
Malta: present, no further details (CIE, 1970)
Poland: The Natural History Museum collection, London, UK
Portugal: present, no further details (CIE, 1970)
Azores: present, no further details (CIE, 1970; Nakahara, 1982)
Madeira Is: present, no further details (Nakahara, 1982)
Romania: present, no further details (CIE, 1970)
Spain: widespread (Amparo Blay Golcoechea, 1993)
Balearic Is: present, no further details (Amparo Blay Golcoechea, 1993)
Canary Is: present, no further details (Amparo Blay Golcoechea, 1993)
Switzerland: present (Kozár et al., 1994; Kozár and Hippe, 1996)
United Kingdom: widespread under glass and outdoors in sheltered situations (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
Channel Is (Guernsey, Jersey): The Natural History Museum collection, London, UK
England: The Natural History Museum collection, London, UK
Scilly Is: The Natural History Museum collection, London, UK
Scotland: The Natural History Museum collection, London, UK
Wales: The Natural History Museum collection, London, UK
Asia
China: present, no further details (Nakahara, 1982)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Shaanxi: present, no further details (Tao, 1999)
Shandong: present, no further details (Tao, 1999)
India
Assam: The Natural History Museum collection, London, UK
Iran: present, no further details (Seghatoleslami, 1977; Nakahara, 1982)
Israel: present, no further details (Izraylevich et al., 1995)
Japan: present, cannot read any further details (Kawai, 1980)
Honshu: The Natural History Museum collection, London, UK
Jordan: present, no further details (CIE, 1970; Nakahara, 1982)
Korea: present, no further details (Nakahara, 1982)
Lebanon: present, no further details (CIE, 1970; Nakahara, 1982)
Saudi Arabia: present, no further details (CIE, 1970)
Syria: present, no further details (CIE, 1970)
Turkey: present, no further details (CIE, 1970)
Africa
Algeria: present, no further details (CIE, 1970; Tao, 1999)
Cape Verde Is: The Natural History Museum collection, London, UK
Congo Democratic Republic: present, no further details (CIE, 1970; Nakahara, 1982)
Egypt: present, no further details (CIE, 1970; El-Minshawy et al., 1974a)
Eritrea: The Natural History Museum collection, London, UK
Ethiopia: present, no further details (Nakahara, 1982)
Kenya: present, no further details (CIE, 1970; Nakahara, 1982)
Libya: present, no further details (CIE, 1970)
Madagascar: present, no further details (CIE, 1970; Nakahara, 1982)
Malawi: present, no further details (CIE, 1970; Nakahara, 1982)
Morocco: present, no further details (CIE, 1970)
Mozambique: present, no further details (Nakahara, 1982)
Nigeria: present, no further details (CIE, 1970; Nakahara, 1982)
North Africa: present, no further details (Nakahara, 1982)
South Africa: present, no further details (CIE, 1970; Nakahara, 1982)
South-West Africa: present (Nakahara, 1982; The Natural History Museum collection, London, UK)
Tanzania: present, no further details (CIE, 1970; Nakahara, 1982)
Tunisia: present, no further details (CIE, 1970)
Uganda: present, no further details (CIE, 1970; Nakahara, 1982)
Zimbabwe: present, no further details (CIE, 1970; Nakahara, 1982)
Western Hemisphere
Antigua: present, no further details (Crouzel, 1973; Nakahara, 1982)
Argentina: very common (Claps et al., 2001a)
Buenos Aires: present, no further details (Claps et al., 2001a)
Catamarca: present, no further details (Claps et al., 2001a)
Córdoba: present, no further details (Claps et al., 2001a)
La Rioja: present, no further details (Claps et al., 2001a)
Mendoza: present, no further details (Claps et al., 2001a)
San Juan: 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)
Bermuda: quite common (Hodgson and Hilburn, 1991)
Bolivia: present, no further details (Nakahara, 1982)
Brazil: present, no further details (Nakahara, 1982)
Guanabara: present, no further details (Silva et al., 1968)
Minas Gerais: 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)
Sao Paulo: present, no further details (Claps et al., 2001a)
Canada: present, no further details (Nakahara, 1982)
Chile: widely distributed (Claps et al., 2001a)
Antofagasta: present, no further details (Claps et al., 2001a)
Atacama: present, no further details (Claps et al., 2001a)
Biobió: present, no further details (Claps et al., 2001a)
Coquimbo: present, no further details (Claps et al., 2001a)
Easter Island: present, no further details (Charlín, 1973; 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: The Natural History Museum collection, London, UK
Tarapacá: present, no further details (Claps et al., 2001a)
Valparaiso: present, no further details (Claps et al., 2001a)
Colombia: present, no further details (Nakahara, 1982; Kondo, 2001)
Costa Rica: present, no further details (Nakahara, 1982)
Cuba: present, no further details (Nakahara, 1982)
Ecuador: present, no further details (Nakahara, 1982; The Natural History Museum collection, London, UK)
Dominican Republic: present, no further details (Nakahara, 1982)
Grenada: present, no further details (Nakahara, 1982)
Guatemala: present, no further details (Nakahara, 1982)
Gaudeloupe: present, no further details (Nakahara, 1982)
Haiti: present, no further details (Nakahara, 1982)
Jamaica: present, no further details (Nakahara, 1982)
Mexico: widespread (Nakahara, 1982; Miller, 1996)
Panama: present, no further details (Nakahara, 1982)
Peru: present, no further details (Canales Canales and Valdivieso, 1999)
Puerto Rico: present, no further details (Nakahara, 1982)
Uruguay: present, no further details (CIE, 1970; Nakahara, 1982)
USA: widespread, under glass in colder areas (CIE, 1970; Nakahara, 1982)
California: widespread and common (Gill, 1997)
Hawaii: present on Oahu and Maui (Heu, 2002)
Oceania
Australia:
Lord Howe I.: present, no further details (Williams and Watson, 1988)
New South Wales: present, no further details (CIE, 1970; CSIRO, 2001)
Queensland: present, no further details (CIE, 1970; CSIRO, 2001)
Tasmania: present, no further details (CSIRO, 2001; The Natural History Museum collection, London, UK)
Victoria: The Natural History Museum collection, London, UK
New Caledonia: present, no further details (Williams and Watson, 1988)
New Zealand: present, no further details (Hill et al., 1993; Charles and Henderson, submitted)
Norfolk I.: present, no further details (Williams and Watson, 1988)