(Linnaeus, 1758)
Diagnosis
In life, scale cover of adult female circular, flat to moderately convex, 1.5-2.5 mm diameter, dark brown or bluish-black with reddish brown central exuviae CHRYAOL5.jpg ; sometimes there is a slightly raised, sub-central point that may be pale. Scale cover of male similar to that of the female but more elongate and sometimes slightly paler, reddish brown subcentral exuviae CHRYAOL6.jpg
CHRAOL.jpg . Body of living female yellow and up to 1.7 mm long. Adult male winged, about 0.8 mm long, with very long genitalia (Ghauri, 1962).
Body of slide-mounted adult female membranous, pyriform (never reniform); with front of head rounded; metathorax with an elongate, pointed marginal spur present on either side; only one prepygidial segment (abdominal segment II) bearing a cluster of more than five submarginal ducts on each side CHRAOS.jpg . Pygidium broad, subtended by an angle greater than 90°, with three pairs of rounded lobes; perivulvar pores present; paraphyses present only between third lobes, each paraphysis longer than a median lobe; first two plates lateral to third lobe fringed CHRAOP.jpg .
Host range
Chysomphalus aonidum is a highly polyphagous species with a preference for Citrus, particularly navel and Valencia oranges and grapefruits (Bedford, 1989). It has been recorded from hosts in 77 plant families, including crops, ornamentals, palms and forestry trees (Borchsenius, 1966); however, its host range is probably wider. In the laboratory, C. aonidum can be mass-reared for biological control purposes on Citrullus spp. (melons or watermelons) (Bedford, 1989) or on potato tubers or pumpkins (Li and Liao, 1990). In tests, Ceballos and Hernández, 1986, found mass-rearing most successful on Cucurbita pepo. Hosts include species of: Acacia, Acer, Agathis, Agave, Aleurites, Aloe, Amaranthus, Annona spp., Aralia, Arbutus, Areca, Artocarpus, Asparagus officinalis, Aspidistra, Barringtonia, Bauhinia, Begonia, Bougainvillea, Calophyllum, Camellia spp., Canna, Carica papaya, Cassia, Castanea, Chamaerops, Chrysalidocarpus, Chrysanthemum, Cinnamomum verum, Citrullus, Citrus spp., Cocos spp., Codiaeum, Coelogyne, Colocasia, Cordyline, Crataegus, Cucurbita, , Cucurbita pepo, Cupressus, Cycas spp., Cynodon, Cyperus, Dendrobium, Dianthus, Dictyosperma, Diospyros, Dodonaea, Dracaena, Dypsis, Elaeagnus, Eriobotrya, Erythrina, Eucalyptus, Eugenia, Euonymus, Euphorbia, Fagraea, Fatsia, Ficus spp., Gardenia, Gerbera, Gladiolus, Gossypium, Hedera, Heliconia, Hibiscus, Howea, Ilex spp., Ipomoea, Ixora, Jasminum, Latania, Lauraceae, Laurus nobilis, Leucopogon, Ligustrum, Lilium, Ludwigia, Macadamia, Magnolia, Malus domestica, Malus sylvestris, Mangifera indica, Maranta, Melaleuca, Morinda, Morus, Musa, Myrtus, Nerium, Nothofagus, Olea europaea, Oncidium, Orchidaceae, Palmae, Pandanus, Persea americana, Phoenix dactylifera, Phormium, Pinus, Plumeria, Podocarpus, Poncirus, Premna, Prunus, Psidium, Punica, Pyrus, Quercus, Rhamnus, Rhododendron, Rhus, Ricinus, Robinia, Rosa, Salix, Sanseviera, Santalum, Smilax, Solanum, Spondias, Strelitzia, Tamarindus, Tamarix, Taxus, Terminalia, Theaceae, Trachycarpus, Vanda, Vanilla, Viburnum, Vicia, Vitis vinifera, Xanthophyllum and Ziziphus.
Affected plant stages: vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: mainly on leaves CHRAOL2.jpg , occasionally on fruits and stems
Biology and ecology
Reproduction in C. aonidum is sexual; no evidence of parthenogenesis has been recorded. The sex ratio in C. aonidum has been found to be biased in favour of females (Nur, 1990). Each adult female lays about 50-150 oval eggs under the scale over a period of 1-8 weeks, depending on the part of the plant infested (those on the leaves being less fecund than those infesting fruits) (Rosen and DeBach, 1978). Eggs hatch under the scale and the first-instar nymphs or crawlers walk about to find a suitable feeding site before settling to a sessile lifestyle. The second-instar nymphs are the main feeding stage in both sexes. Development to adult takes 7-16 weeks according to temperature. In California, C. aonidum has up to 6 generations per year (Gill, 1997). In countries with a cold winter, such as Taiwan, there may be three distinct generations per year (Su, 1983), while in tropical conditions (and heated greenhouses) breeding is continuous and generations are asynchronous.
Chrysomphalus aonidum has a preference for humid environments and cannot tolerate freezing temperatures. It tends to prefer the lower and central parts of mature Citrus trees and rarely infests green wood (Rosen and DeBach, 1978). Male stages are rather more tolerant of lower humidity than females, so male scales are more often found on the upper surface of the leaf while females congregate on lower leaf surfaces (Bedford, 1989). Like other diaspidid scale insects, C. aonidum suffers increased mortality in heavy rain and reaches high population levels during dry weather.
The first instar stage is the sole dispersal stage. Each crawler walks to an exposed position on the plant, from which air currents may carry it as much as several tens of kilometres away (Greathead, 1990). Passing animals or people can also can carry the crawlers over great distances. Movement of infested planting material or produce is the main way in which C. aonidum has been introduced to other countries.
This species has 2n=8 chromosomes (Nur, 1990).
Symptoms
Chrysomphalus aonidum is a leaf-infesting species, but in high-density infestations it may spread to fruits, stems and trunks and may cause premature leaf and fruit drop and stem dieback. The scales appear as circular dark spots. An infestation appears as dark-purple to reddish-brown or black spots with paler margins, on both surfaces of shaded leaves of the host plant. Heavy infestations cause yellowing of the leaves CHRYAOL4.jpg
CHRYAOL7.jpg , followed by defoliation of part or all of the host. Chrysomphalus aonidum prefers shade and is therefore most common in the lower part of the canopy.
Economic impact
The most important crop damaged by C. aonidum is Citrus: damage has been recorded in the USA (Florida, Texas), Brazil, Mexico, Cuba, Puerto Rico, Central America, Trinidad, Colombia, Venezuela, Argentina, Paraguay, Uruguay, Italy and North Africa, Lebanon, Egypt, Israel, South Africa, India, Pakistan, Australia and China (Rosen and DeBach, 1978; Rose, 1990) and Western Samoa (Maddison, 1976). According to Bedford, 1989, in unsprayed Citrus orchards in South Africa C. aonidum used to cause almost complete defoliation of individual trees, which would produce hardly any crop the following season. Heavy infestation of fruits resulted in up to 100% culling at the packhouse. Expenditure on insecticide sprays to control the scale was high until the introduction of effective biological control of C. aonidum reducedinfestation by over 50%. Heavy infestation of [l][m]Host plants[/m][r]Citrus[/r]Citrus foliage in New Caledonia resulted in death of trees (Cohic, 1950). In 1976, it was estimated that C. aonidum caused an annual loss on Citrus in Texas of US3.85 million (Kosztarab, 1990).
Chrysomphalus aonidum is a pest of olive in Israel and Turkey (Argyriou, 1990). Danzig and Pellizzari, 1998, refer to the species as a dangerous pest in the Palaearctic region. In Poland, this species causes serious damage to ornamental plants in glasshouses (Labanowski, 1999). It has been recorded severely damaging pine seedlings in Papua New Guinea (Szent-Ivany and Stevens, 1966), and causes problems on several economically important hosts in Brazil (Claps et al., 2001a). Foldi, 2001, lists this species as an occasional pest in France. Chrysomphalus aonidum has been recorded causing serious damage to young tea in India (Das, 1974); as a serious pest of bananas in the Caribbean and Central America (Rosen and DeBach, 1978), and as an occasional pest of bananas in Israel, where heavy infestations developed in high temperatures and made the fruit unmarketable (Chua and Wood, 1990). It is a problem on coconuts in the Philippines (Rosen and DeBach, 1978) and was a serious pest of coconut in the Seychelles in the 1930s, causing yellow patches on infested pinnae and death of whole leaves, leading to reductions in yield in severe outbreaks (Vesey-Fitzgerald, 1940).
Detection and inspection methods
Examine plants closely for dark spots on the leaves, especially in shady parts of the plant. Good light conditions are essential and in poor light, a powerful flashlight is helpful. A large hand lens may assist in recognition of small, immature stages.
Phytosanitary risk
Chrysomphalus aonidum is already fairly widespread, particularly in Citrus-growing regions of the world. As global warming progresses, its distribution is likely to extend north- and southwards with the expansion of Citrus-producing areas. Countries with new or developing Citrus industries just outside the currently known distribution of C. aonidum (CABIIE, 1988) will need to be vigilant to exclude the scale if possible. The scale may also become a problem on ornamental plants in these countries, and could become a pest of ornamentals under glass in temperate countries, as it has done in Hungary (Reiderné and Kozár, 1994). It is mentioned on quarantine lists (Burger and Ulenberg, 1990).
Natural enemies
Parasitoids:
- Ablerus perspeciosus in China
- Alaptus sp. in Brazil
- Aphytis chionaspis, attacking: eggs, larvae, nymphs, pupae, adults, in India, Israel, Suriname
- Aphytis chrysomphali, attacking: nymphs in Australia, Suriname, India
- Aphytis columbi, attacking: nymphs, in Australia
- Aphytis costalimai, in Argentina (Tucumán)
- Aphytis holoxanthus, attacking: nymphs, adults, in China, Hong Kong, Taiwan; introduced: Israel, Australia (Queensland), USA (California, Florida, Texas), Argentina, Brazil, Peru, Mexico, Cyprus, Egypt, South Africa
- Aphytis lingnanensis, attacking: nymphs, in China, Taiwan and Japan. Introduced: USA (California), South Africa, Australia, Mexico, Argentina, Chile, Cyprus, Israel, Morocco
- Aprostocetus purpureus
- Arrhenophagus chionaspidis
- Comperiella bifasciata, attacking: nymphs, in India, China and Japan. Introduced: USA (California), South Africa, Swaziland, Australia, Mexico, Argentina, Israel
- Comperiella pia, attacking: nymphs, in Australia (possibly host specific)
- Encarsia aurantii
- Encarsia citrina, attacking: nymphs, in Japan. Introduced to: USA; Australia; Mediterranean Basin; Turkey; Italy, Indonesia (Bali), Tahiti, Fiji, Cook Islands, Egypt, Africa, Brazil
- Encarsia herndoni, attacking: nymphs, adults
- Encarsia lounsburyi, attacking: nymphs, adults in Brazil, Cuba, Egypt
- Habrolepis pascuorum, in Egypt
- Metaphycus helvolus
- Pseudhomalopoda elongata, attacking: nymphs, in USA (Texas, Florida)
- Pseudhomalopoda prima, attacking: nymphs, in USA (Texas, Florida)
- Pteroptrix smithi, attacking: nymphs, in China, Hong Kong; introduced: Israel, Mexico
- Signiphora fax in Argentina
Predators:
- Aleurodothrips fasciapennis, attacking: eggs, nymphs, adults, in Indonesia; introduced to: Fiji
- Chilocorus bipustulatus, attacking: eggs, larvae, nymphs, pupae, adults, in Israel
- Chilocorus circumdatus, attacking: eggs, larvae, nymphs, pupae, adults, in India (Assam)
- Chilocorus distigma in East Africa, intorduced to: Seychelles
- Chilocorus kuwanae, attacking: nymphs, adults
- Chilocorus nigrita, attacking: eggs, larvae, nymphs, pupae, adults, in India; introduced: Seychelles, South Africa
- Chilocorus renipustulatus, attacking: nymphs, adults
- Chrysopa sp., in Brazil (Sao Paulo)
- Haplothrips cahirensis, in Egypt, Sudan
- Pentilia egena, in Brazil (Sao Paulo)
- Pharoscymnus horni, attacking: eggs, larvae, nymphs, pupae, adults, in India
- Rhyzobius lophanthae, attacking: eggs, larvae, nymphs, pupae, adults, introduced to South Africa
- Rhyzobius pulchellus, attacking: nymphs, adults, in Vanuatu, New Caledonia
- Scymnus severini, attacking: nymphs, adults
Pathogens:
- Beauveria bassiana in China
- Cladosporium cladosporoides in Egypt
- Fusarium coccidicola, in India (Assam)
- Fusarium coccophilum [Nectria flammea] in China
- Podonectria coccicola in India, China
- Verticillium sp. in China
Distribution
See Chrysomphalus aonidum distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species. This species is indistinguishable from C. bifasciculatus in the field (Gill, 1997). On ornamental plants, it would be possible to confuse C. aonidum in life with species of Lindingaspis and Melanaspis, but the slide-mounted females are quite distinct.
Chrysomphalus ansei (Green) could be misidentified as C. aonidum, but differs in possessing no more than 4 marginal macroducts on each side of each prepygidial abdominal segment. In contrast, C. aonidum possesses one prepygidial abdominal segment with a group of more than 5 marginal macroducts on each side CHRAOS.jpg. Chrysomphalus ansei is known only from the Seychelles on coconut and Litsea.
Chrysomphalus propsimus Banks could be misidentified as C. aonidum, but differs in possessing no more than 4 marginal macroducts on each side of each prepygidial abdominal segment. In contrast, C. aonidum possesses one prepygidial abdominal segment with a group of more than 5 marginal macroducts on each side CHRAOS.jpg. Chrysomphalus propsimus is known from the Philippines, Indonesia (Sumatra), Kiribati, Tuvalu and Jamaica on species of Calamus, Cocos nucifera and Pandanus (Williams and Watson, 1988; The Natural History Museum collection, London, UK).
Comments
Chrysomphalus aonidum is a tropical species, apparently native to the Oriental region (Gill, 1997) but it has been dispersed widely in tropical and sub-tropical parts of the world, mostly in association with the Citrus industry. In northern countries in its range, this species occurs under glass. In 1997 there were no known infestations in California (Gill, 1997).
Europe
Belgium: present, no further details (Danzig and Pellizzari, 1998)
Croatia: present, no further details (EPPO, 1999)
Cyprus: present, no further details (CABIIE, 1988a)
Former Yugoslavia: present, no further details (CABIIE, 1988a)
France: present, no further details (Foldi, 2001)
Germany: present, no further details (Danzig and Pellizzari, 1998)
Greece: present, no further details (CABIIE, 1988a)
Italy: under glass (Longo et al., 1995; Danzig and Pellizzari, 1998)
Malta: present, no further details (CABIIE, 1988a)
Netherlands: present, no further details (Danzig and Pellizzari, 1998)
Portugal
Madeira: present, no further details (CABIIE, 1988a)
Romania: present, no further details (CABIIE, 1988a)
Spain: present, no further details (CABIIE, 1988a)
Canary Islands: present, no further details (CABIIE, 1988a)
United Kingdom: restricted to a few botanical gardens, under glass (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
England: The Natural History Museum collection, London, UK
Scotland: The Natural History Museum collection, London, UK
Wales: (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
Asia
Bhutan: present, no further details (CABIIE, 1988a)
Chagos Archipelago: present, no further details (CABIIE, 1988a)
China
Fujian: present, no further details (CABIIE, 1988a; Tao, 1999)
Guangdong: present, no further details (CABIIE, 1988a; Tao, 1999)
Guangxi: present, no further details (CABIIE, 1988a)
Guizhou: present, no further details (Tao, 1999)
Hainan: present, no further details (Tao, 1999)
Hebei: present, no further details (CABIIE, 1988a)
Hong Kong: present, no further details (CABIIE, 1988a; Tao, 1999)
Hunan: present, no further details (CABIIE, 1988a)
Inner Mongolia: present, no further details (Tao, 1999)
Jiangsu: present, no further details (CABIIE, 1988a; Tao, 1999)
Jiangxi: present, no further details (CABIIE, 1988a; Tao, 1999)
Shandong: present, no further details (CABIIE, 1988a; Tao, 1999)
Sichuan: present, no further details (CABIIE, 1988a)
Suchuan: present, no further details (Tao, 1999)
Yunnan: present, no further details (CABIIE, 1988a)
Zhejiang: present, no further details (Tao, 1999; Wang et al., 2000)
India
Andhra Pradesh: present, no further details (Dhileepan, 1991)
Assam: present, no further details (CABIIE, 1988a)
Gujarat: present, no further details (CABIIE, 1988a)
Karnataka: present, no further details (CABIIE, 1988a)
Kerala: present, no further details (Dhileepan, 1991)
Madhya Pradesh: present, no further details (CABIIE, 1988a)
Maharashtra: present, no further details (CABIIE, 1988a)
New Delhi: The Natural History Museum collection, London, UK
Tamil Nadu: present, no further details (CABIIE, 1988a)
Tripura: present, no further details (CABIIE, 1988a)
West Bengal: present, no further details (CABIIE, 1988a)
Indonesia
Java: present, no further details (CABIIE, 1988a)
Sulawesi: present, no further details (CABIIE, 1988a)
Sumatra: present, no further details (CABIIE, 1988a)
Israel: present, no further details (CABIIE, 1988a)
Japan: present (Kawai, 1980; Tao, 1999)
Honshu: present, no further details (CABIIE, 1988a)
Kyushu: present, no further details (CABIIE, 1988a)
Shikoku: present, no further details (CABIIE, 1988a)
Jordan: present, no further details (CABIIE, 1988a)
Korea, DPR: present, no further details (CABIIE, 1988a)
Korea, Republic of: present, no further details (CABIIE, 1988a)
Lebanon: present, no further details (CABIIE, 1988a)
Malaysia
West Malaysia: present, no further details (CABIIE, 1988a)
Sabah: present, no further details (CABIIE, 1988a)
Myanmar: present, no further details (CABIIE, 1988a; Waterhouse, 1993)
Oman: present, no further details (EPPO, 1999)
Pakistan: present, no further details (CABIIE, 1988a)
Philippines: present, no further details (Velasquez, 1971; CABIIE, 1988)
Ryukyu Archipelago: present, no further details (CABIIE, 1988a)
Saudi Arabia: present, no further details (CABIIE, 1988a)
Singapore: present, no further details (CABIIE, 1988a)
Sri Lanka: present, no further details (CABIIE, 1988a; Tao, 1999)
Syria: present, no further details (CABIIE, 1988a; Danzig and Pellizzari, 1998)
Taiwan: present, no further details (CABIIE, 1988a; Wong et al., 1999)
Thailand: present, no further details (Waterhouse, 1993)
Turkey: present, no further details (CABIIE, 1988a)
Vietnam: present, no further details (Waterhouse, 1993)
Yemen: present in both North and South (CABIIE, 1988a)
Africa
Algeria: present, no further details (CABIIE, 1988a; Danzig and Pellizzari, 1998)
Burundi: present, no further details (CABIIE, 1988a)
Comoro Is: present, no further details (CABIIE, 1988a)
Egypt: present, no further details (El-Minshawy et al., 1974a; CABIIE, 1988a)
Ethiopia: present, no further details (CABIIE, 1988a)
Guinea: present, no further details (CABIIE, 1988a)
Kenya: present, no further details (CABIIE, 1988a)
Madagascar: present, no further details (CABIIE, 1988a)
Malawi: present, no further details (CABIIE, 1988a)
Mauritius: present, no further details (CABIIE, 1988a; Williams and Williams, 1988)
Morocco: present, no further details (CABIIE, 1988a; Danzig and Pellizzari, 1998)
Mozambique: present, no further details (CABIIE, 1988a)
Nigeria: present, no further details (CABIIE, 1988a)
Réunion: present, no further details (CABIIE, 1988a; Williams and Williams, 1988)
Rodrigues: present, no further details (CABIIE, 1988a; Williams and Williams, 1988)
Senegal: present, no further details (CABIIE, 1988a)
Seychelles: present, no further details (CABIIE, 1988a)
South Africa: present, no further details (CABIIE, 1988a)
Sudan: present, no further details (CABIIE, 1988a)
Tanzania: present, no further details (Bohlen, 1973; CABIIE, 1988a)
Zanzibar: present, no further details (CABIIE, 1988a)
Tunisia: present, no further details (CABIIE, 1988a)
Uganda: present, no further details (CABIIE, 1988a)
Zambia: present, no further details (CABIIE, 1988a)
Zimbabwe: present, no further details (CABIIE, 1988a)
Western Hemisphere
Argentina
Buenos Aires: present, no further details (Claps et al., 2001a)
Catamarca: present, no further details (Claps et al., 2001a)
Chaco: present, no further details (Claps et al., 2001a)
Corrientes: present, no further details (Claps et al., 2001a)
Entre Rios: present, no further details (CABIIE, 1988a)
Jujuy: present, no further details (CABIIE, 1988a)
La Rioja: present, no further details (Claps et al., 2001a)
Salta: present, no further details (CABIIE, 1988a)
Santa Fe: present, no further details (CABIIE, 1988a)
Tucumán: present, no further details (Claps et al., 2001a)
Barbados: present, no further details (Bennett and Alam, 1985; CABIIE, 1988)
Bermuda: fairly common (Hodgson and Hilburn, 1991)
Brazil: widely distributed (Claps et al., 2001a)
Acre: present, no further details (CABIIE, 1988a)
Amazonas: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Bahia: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Cerrados: present, no further details (Murakami et al., 1984)
Guanabara: present, no further details (Silva et al., 1968)
Maranhao: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Mato Grosso: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Minas Gerais: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Pará: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Paraíba: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Paraná: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Pernambuco: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Rio de Janeiro: present, no further details (CABIIE, 1988a; 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)
Santa Catarina: present, no further details (CABIIE, 1988a; Claps et al., 2001a)
Sao Paulo: present, no further details (Watanabe et al., 2000a; Claps et al., 2001a)
Cayman Islands: present, no further details (CABIIE, 1988a)
Chile: present, no further details (CABIIE, 1988a)
Antofagasta: present, no further details (Claps et al., 2001a)
Easter I.: present, no further details (Charlín, 1973)
Tarapacá: present, no further details (Claps et al., 2001a)
Colombia: present, no further details (CABIIE, 1988a; Kondo, 2001)
Cuba: present, no further details (CABIIE, 1988a)
Dominica: present, no further details (CABIIE, 1988a)
Dominican Republic: present, no further details (CABIIE, 1988a)
El Salvador: present, no further details (CABIIE, 1988a)
French Guiana: present, no further details (CABIIE, 1988a)
Guadeloupe: present, no further details (CABIIE, 1988a)
Guatemala: present, no further details (CABIIE, 1988a)
Guyana: present, no further details (CABIIE, 1988a)
Haiti: present, no further details (CABIIE, 1988a)
Honduras: present, no further details (CABIIE, 1988a)
Jamaica: present, no further details (CABIIE, 1988a)
Martinique: present, no further details (CABIIE, 1988a)
Mexico: widespread (Miller, 1996)
Montserrat: present, no further details (CABIIE, 1988a)
Panama: present, no further details (CABIIE, 1988a)
Paraguay: present, no further details (CABIIE, 1988a)
Puerto Rico: present, no further details (CABIIE, 1988a)
Saint Lucia: present, no further details (CABIIE, 1988a)
Saint Vincent: present, no further details (CABIIE, 1988a)
Suriname: present, no further details (CABIIE, 1988a)
Trinidad: present, no further details (CABIIE, 1988a)
Uruguay: present, no further details (CABIIE, 1988a)
USA: present in most states, under glass in colder areas (Nakahara, 1982; CABIIE, 1988a) but absent from California (Gill, 1997)
District of Columbia: present, no further details (CABIIE, 1988a)
Florida: present, no further details (Beardsley, 1966)
Hawaii: widespread (Heu, 2002)
Maryland: present, no further details (CABIIE, 1988a)
Mississippi: present, no further details (CABIIE, 1988a)
Texas: present, no further details (CABIIE, 1988a)
Virginia: present, no further details (CABIIE, 1988a)
United States Virgin Islands: present, no further details (CABIIE, 1988a)
Venezuela: present, no further details (CABIIE, 1988a)
Oceania
American Samoa: present, no further details (CABIIE, 1988a)
Australia: present, no further details (Tao, 1999)
New South Wales: present, no further details (CABIIE, 1988a; CSIRO, 2001)
Northern Territory: present, no further details (CABIIE, 1988a; CSIRO, 2001)
Queensland: present, no further details (CABIIE, 1988a; CSIRO, 2001)
Tasmania: present, no further details (CSIRO, 2001)
Bonin Is: present (Beardsley, 1966)
Caroline Islands: present, no further details (CABIIE, 1988a)
Cook Is: present, no further details (Maddison, 1976)
Federated States of Micronesia: present, no further details (EPPO, 1999)
Fiji: present (Williams and Watson, 1988)
Kiribati: present (Williams and Watson, 1988)
Nauru: The Natural History Museum collection, London, UK
New Caledonia: present (Williams and Watson, 1988)
Niue: present, no futher details (Maddison, 1976)
Ogasawara-shoto: present, no further details (CABIIE, 1988a)
Papua New Guinea: present (Williams and Watson, 1988)
Pohnpei: present, no further details (Beardsley, 1966)
Samoa: present, no further details (CABIIE, 1988a)
Solomon Is: present (Williams and Watson, 1988)
Tahiti: present (Doanne and Hadden, 1909; Cohic, 1955)
Tonga: present (Williams and Watson, 1988)
Tuvalu: present (Williams and Watson, 1988)
Western Samoa: present, no further details (Reddy, 1970; Williams and Watson, 1988)