Aonidiella aurantii

(Maskell, 1879)

Diagnosis
Scale cover of adult female in life circular, 1.5-2.0 mm diameter, flat, semi-translucent red-brown or red-grey with reniform orange female showing through it; exuviae darker, central AONAURL2.jpg . Scale cover of male in life elongate oval, 1.0-1.3 mm long and half as broad, greyish-brown with red-brown subterminal exuvae AONAUL.jpg (Davidson and Miller, 1990; Bedford, 1998) AOAUL.jpg. The adult male is winged (Ghauri, 1962). Colour photographs of all the life stages are given by Flanders et al., 1995.

Body of slide-mounted adult female 0.7-1.2 mm across; prosoma becoming highly sclerotized and expanded with maturity; eventually the lateral lobes may project further posteriorly than the apex of the pygidium AOAUO.jpg . (This reniform shape characterises the genera Aonidiella and Africonidia (Ben-Dov, 1990a), although Aspidiotus macfarlanei is also reniform (Williams and Watson, 1988). Abdominal segments lacking any prepygidial macroducts. Pygidium with paraphyses present, most or all no longer than median lobes; perivulvar pores absent; prevulvar apophyses and scleroses both present; and plates lateral to third lobes each fringed (not with a single fleshy process) AOAUP1.jpg .

Host range
Aonidiella aurantii occurs on host-plants belonging to at least 77 plant families (Borchsenius, 1966) but is mainly known as an important pest of Citrus. This species has been successfully reared on potato tubers and pumpkin (Li and Liao, 1990). Its host range includes species of: Abelmoschus esculentus, Acacia, Aleurites, Annona, Araucaria, Ardisia, Artocarpus altilis, Asparagus plumosus, Bauhinia, Bromeliaceae, Broussonetia, Buxus, Camellia sinensis, Capsicum frutescens, Carica papaya, Casuarina, Ceiba, Chaenomeles, Cinnamomum, Citrus spp., Cocos nucifera, Cucurbita pepo, Cycas revoluta, Erythrina, Eucalyptus, Euonymus japonica, Ficus carica, Gardenia, Gossypium, Grevillea, Hedera, Hibiscus, Ilex, Jasminum, Jodina, Juglans regia, Laurus, Ligustrum, Malus, Mangifera indica , Melia, Morus nigra, Musa sapientum, Nerium oleander, Olea europaea, Palmae, Passiflora edulis, Persea americana, Phoenix dactylifera, Pistacea, Psidium guajava, Punica granatum, Ricinus communis, Rosa, Rubus, Salix, Schefflera, Sida, Tamarindus, Vitis vinifera, Yucca, Ziziphus.

Affected plant stages: vegetative growing, flowering, fruiting and post-harvest stages

Affected plant parts: whole plant, even occasionally on roots, but especially on leaves, young branches AONAULL1.jpg and fruits AONAUL3.jpg

Biology and ecology
A. aurantii reproduces sexually and is viviparous. The sex ratio of females to males varies from 1:1 to 2.6:1 (Parry-Jones, 1936; Ebeling, 1959). Stofberg, 1937, gives a ratio of 1 female to 3 males. Males locate unmated females by following pheromones released by them. Identification and synthesis of the female pheromone of A. aurantii has been achieved (Roelofs et al., 1978). Detailed descriptions of the biology of A. aurantii are given by Quayle, 1941; and Ebeling, 1959.

There is a close correlation between temperature and development (Parry-Jones, 1936; Stofberg, 1937). The period from crawler to crawler for red scale females on leaves outdoors at Nelspruit, South Africa varies from 61 days (in mid-summer) to 138 days (in winter) according to Stofberg, 1937. Development is slower on leaves than on fruit; females on leaves also produce fewer crawlers (Parry-Jones, 1936).

On Citrus in Zimbabwe (Parry-Jones, 1936), the average duration of the first and second instars and the adult stage up to the commencement of reproduction by females is 13, 10 and 32 days respectively in early summer, and up to 24, 24 and 70 days respectively in winter. The total developmental period of the female is 55 to 118 days, and for the male, 26 -76 days. The life cycle takes longer under field conditions in California, USA (Nel, 1933; Quayle, 1941).

Females on Citrus fruit in Zimbabwe (Parry-Jones, 1936) each produce 66 - 143 crawlers in 33 - 48 days, depending on environmental conditions. The highest rate of production is almost 7 crawlers per day per female in summer, but only one crawler per day in winter.

All stages are present throughout the year in most of South Africa, without any dormant stage; however, there is a distinct sequence of four field generations in some parts of South Africa (Bedford, 1968). Stofberg, 1937, refers to four generations, with sometimes a partial fifth. Parry-Jones, 1936, calculated five generations in shade conditions and up to seven in the sun, but Bedford, 1998, found the same number of field generations throughout the tree, despite the different rate of growth on leaves and fruit. Grout and Richards, 1989, using sticky traps with a female pheromone in South Africa, found four to six generations per year on orange, and 5-7generations per year on lemon. According to Davidson and Miller, 1990, A. aurantii has two to three generations per year in California, six in Argentina and four in Cyprus.

After harvest, A. aurantii overwinters on twigs and leaves. Immatures that survive the winter mature in early summer and the crawlers infest the new growth and young green fruit.

Some ant species cause coincident infestations of red scale (DeBach et al., 1951; Steyn, 1954a, Steyn, 1954b; Annecke, 1958, Annecke, 1959; Compere, 1961; Bedford, 1968). They patrol the branches to obtain honeydew from soft scale insects, mealybugs, cottony cushion scale (Icerya purchasi) and aphids. Their restless activity deters parasitoids and predators. The honeydew-producing insects increase rapidly and provide more food for ants (as well as causing sooty mould) - but red scale (which provides nothing for the ants) also increases quite out of proportion to its normal low level in the absence of ants. Ants can thus be responsible for making A. aurantii a major pest, while it is a minor pest in orchards where ants are controlled (Bedford, 1998).

The dispersal phase of A. aurantii 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 (Ebeling, 1950) and transport of infested plant material by man. They emerge from under the female's scale and search for a suitable feeding site at which to settle permanently. The majority emerge before 12.00 h and settle within 6 hours, on older branches and leaves; but they are strongly phototactic and tend to move outwards onto the new growth and green fruit. They settle mostly on the upper surface of leaves, especially alongside the midrib or the larger veins. On very young fruit they usually settle in the depressions of the oil glands, but mature fruits can be completely encrusted with scales of all sizes. 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
Aonidiella aurantii inserts its mouthparts deep into plant tissue and sucks sap from parenchyma cells (Bedford, 1998). Saliva injected as the sales feed is very toxic to the leaves, twigs, branches and fruit of Citrus trees. The leaves develop a characteristic yellow spot under and around each female scale. Prolonged infestation may cause leaf drop and defoliation, and dieback of twigs and eventually large branches. Maturing fruit can become completely encrusted with scales; developing scales form prominent pits on young fruit which are still evident when the fruit matures. Such fruit tend to dry out and fall off. Even the trunk can become heavily infested, especially in young Citrus trees. Newly planted trees may be severely set back or even killed. Neglected heavy infestations lower the productivity of an orchard (Bedford, 1998). On other fruits, infestation may cause discolouration AONAUL3.jpg.

Economic impact
Aonidiella aurantii is potentially a severe pest of Citrus in California (USA), Australia, New Zealand, Mexico, Chile, Argentina, Brazil, Israel and islands of the eastern Mediterranean (Ebeling, 1959; Claps et al., 2001a), and South Africa (Bedford, 1998). The severity of attack varies, the descending order of susceptibility being lemons, grapefruit, navel oranges, mid-season and Valencia oranges (these two the same) and finally mandarin oranges. All can become heavily infested, especially if ants or insecticides disturb the natural balance (Bedford, 1998). On Citrus, the toxic saliva injected into the host can kill leaves, twigs and branches, and stunts and pits the fruit so that it falls or is unmarketable (Williams and Watson, 1988).

Historically, A. aurantii has been the most notorious pest on Citrus in many Citrus-producing countries in the world. The literature on this pest is very extensive, most of the research work having been conducted in California, USA. Excessive populations can build up very rapidly, and considerable damage can result (Bedford, 1998). In severe infestations, developing scales form prominent pits on young fruit which are still evident when the fruit matures. Severe infestation can also reduce the final crop yield and the following year's crop, as a result of defoliation and dieback (Bedford, 1998).

According to Outspan International, no more than six scale insects of 1 mm diameter or larger (of any species) are allowed per average-sized orange for export grade orange fruit (count 88), or 10 insects per grapefruit (count 40). For lemons and soft Citrus, the figures are five and four scale insects on counts 113 and calibre 3, respectively, per average-sized fruit for export grade. As a population density of only 50 females per 100 fruit results in nearly 1% of crop rejection for export in the packhouse, any population increase above this level soon entails commercial losses. When red scale infestations are exceptionally heavy, rejection of fruit can reach 100%, at a scale density of 7700 per 100 fruit (Bedford, 1971). A high pressure unit for rinsing Citrus fruit in the packhouse has been developed to remove armoured scale insects such as A. aurantii, using a water jet at a pressure of 28 bar (Bedford, 1990).

Aonidiella aurantii has also been recorded damaging papaya (Carica papaya) in Taiwan (Chua and Wood, 1990), guava (Psidium guajava) in India (Hayes, 1970) and olive in countries around the Mediterranean and California; it caused serious damage on this crop in Morocco (Argyriou, 1990).

Detection and inspection methods
Inspect leaves, branches and fruit, aided by a x8 hand lens and a powerful torch if required, or using a dissection microscope in the laboratory. Look for reddish-orange, flattened, circular scale covers, each about 1.5-2.0 mm in diameter (adult females). Between these there are usually many immature scales which may be very small and white (first instars) and others which are greyish to red and intermediate in size.

The presence of red scale in an orchard can be determined by trapping the males using hanging traps with synthetic pheromones (Kozár, 1990). Male A. aurantii are also attracted to the colours white and yellow.

Phytosanitary risk
Aonidiella aurantii is included in the quarantine lists of many countries (Burger and Ulenberg, 1990). The species is easily transported on infested fruit and planting material.

Natural enemies
The sex pheromones of red scale, which are commercially available for use in monitoring the scale population, may act as kairomones by attracting parasitic wasps (Gieselmann and Rice, 1990).

A description of parasitized A. aurantii with colour photographs of the stages is given by Flanders et al., 1995.

Parasitoids:
- Aphelinus africanus, attacking: nymphs, in South Africa. Introduced: Argentina
- Aphytis chionaspis, attacking: nymphs, in China. Introduced: South Africa, Australia
- Aphytis chrysomphali, attacking: nymphs in Australia, Argentina
- Aphytis coheni, attacking: nymphs, in Israel. Introduced: South Africa
- Aphytis holoxanthus , attacking: nymphs, in Australia
- Aphytis lingnanensis, attacking: nymphs, in China, Taiwan and Japan. Introduced: USA (California), South Africa, Australia, 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: USA (California), South Africa, Swaziland, Australia, Mexico, Argentina, Israel
- Encarsia citrina, attacking: nymphs, in Japan. Introduced to: USA; Australia; Mediterranean Basin; Turkey; Italy, Indonesia (Bali), Tahiti, Fiji, Cook Islands, Africa
- Encarsia perniciosi, attacking: nymphs, in China, Taiwan. Introduced: USA (California), Australia
- Habrolepis rouxi, attacking: nymphs, in South Africa. Introduced: USA (California)
- Signiphora fax

Predators:
- Aleurodothrips fasciapennis, in China
- Chilocorus cacti, attacking: nymphs, adults, in USA. Introduced: South Africa, Swaziland
- 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
- Cybocephalus micans, in Israel
- Cryptolaemus montrouzieri
- Hemisarcoptes malus
- Rhyzobius lophanthae, attacking: nymphs, adults, in Australia. Introduced: South Africa, Swaziland, Chile

Fungal diseases:
- Nectria coccophila, in Brazil
- Myriangium duriaei, in Brazil

Distribution
See Aonidiella aurantii distribution.

Microscopic examination of slide-mounted adult females is required for authoritative identification to species. Aonidiella aurantii is very similar to A. citrina and could be confused with A. inornata McKenzie, 1938, as all three species lack perivulvar pores. Aonidiella aurantii possesses prevulvar apophyses and scleroses AOAUP.jpg, whereas A. citrina possesses only prevulvar apophyses AOCIP.jpg and A. inornata lacks prevulvar apophyses and scleroses entirely AOINORP.jpg. The precise identity of A. inornata requires research, as its morphology is rather variable; Williams and Watson, 1988, remark that it may be a synonym of A. comperei McKenzie, 1938 or A. eremocitri McKenzie, 1937a (these species are discussed below). A key to separate five economically important species of Aonidiella in the tropics is given by Williams and Watson, 1988 (although this key does not include A. citrina). McKenzie, 1938, provided a key to separate the species of Aonidiella.

According to the literature, A. inornata AOINORP.jpg is fairly polyphagous on the leaves and fruit of hosts including species of Allamanda, Annesijoa, Areca, Artocarpus, Astronia, Barringtonia, Bischofia, Campnosperma, Carica, Citrus, Cocos, Cordyline, Cycas, Dracaena, Elettaria, Euphorbia, Hedyotis, Jasminum, Mangifera, Melaleuca, Metroxylon, Musa, Nerium, Pandanus, Piper, Plumeria and Vitis vinifera. It is known from India (Uttar Pradesh), Maldives, Indonesia (including Irian Jaya), Laos, West Malaysia, Sarawak, Brunei Darussalam, Thailand, China (Hainan, Hebei, Hong Kong, Inner Mongolia, Liaoning, Yunnan), South-East Asia, Taiwan, Japan, Okinawa, Philippines, Papua New Guinea, Australia (Queensland), Guam, Caroline Is, Marshall Is, Palau, Is, Pohnpei, Truk, Wake Is, Yap Is, Kiribati, Vanuatu, Fiji, Western Samoa, USA (Hawaii), Cayman Is, Puerto Rico and East Africa (Pemba I., Tanzania) (Takagi, 1969; Velasquez, 1971; Kawai, 1980; Nakahara, 1982; Williams and Watson, 1988; Watson et al., 1995; Danzig and Pellizzari, 1998; Tao, 1999; Heu, 2002; The Natural History Museum collection, London, UK). AOINORS.jpg

Aonidiella comperei McKenzie (false yellow scale) AOCOMPP.jpg and A. eremocitri McKenzie AONERP1.jpg are superficially similar to A. aurantii but possess a few perivulvar pores on either side of the vulva, and lack the prevulvar scleroses and apophyses that are present in A. aurantii AOAUP.jpg. Aonidiella comperei AOCOMPS.jpg has been recorded from Barbados, Brazil (Alagôas, Guanabara, Paraíba, Pernambuco, Rio de Janeiro), Tanzania, China (Inner Mongolia), Taiwan, India (Lakshadweep Is, Maharashtra), Indonesia (Sulawesi), Maldive Is, Sri Lanka, Malaysia, Sarawak, Philippines, Mariana Is, Palau, Yap Is, Caroline Is, Truk Is, Marshall Is, Kiribati, Papua New Guinea and Australia (Queensland) on hosts from 8 plant families - Celastraceae, Compositae, Ebenaceae, Euphorbiaceae, Moraceae, Musaceae, Palmae and Rubiaceae (Borchsenius, 1966; Beardsley, 1966; Silva et al., 1968; Velasquez, 1971; Bennett and Alam, 1985; Williams and Watson, 1988; Watson et al., 1995; Claps et al., 2001a; The Natural History Museum collection, London, UK), including species of Annesijoa, Carica papaya, Citrus sinensis, Diospyros, Ficus, Melissa, Morinda, Musa, Pluchea, Rosa, Tamarindus and Vitis vinifera, and may have a wider host range. AONCOMP1.jpg

A. eremocitri AONERS.jpg has been recorded from Malaysia (Sarawak), Australia, Papua New Guinea, Solomon Is, Palau Is, Vanuatu and Fiji, on hosts from 9 plant families - Compositae, Ebenaceae, Euphorbiaceae, Moraceae, Musaceae Orchidaceae, Palmae, Rubiaceae and Rutaceae including species of Barringtonia, Bischofia, Citrus, Cocos nucifera and Eremocitrus (Borchsenius, 1966; Beardsley, 1966; Bennett and Alam, 1985; Williams and Watson, 1988, Tao, 1999); it probably has a wider host range.

It is possible for inexperienced identifiers to confuse Aspidiotus macfarlanei Williams and Watson with Aonidiella aurantii. Aspidiotus macfarlanei also develops a sclerotized, reniform body at maturity ASPMACS.jpg (Williams and Watson, 1988), but has marginal prepygidial macroducts and lacks paraphyses on the pygidium ASPMACP.jpg. Aonidiella aurantii lacks marginal prepygidial macroducts, and possesses paraphyses AOAUP1.jpg. Aspidiotus macfarlanei has been recorded from the Solomon Islands on Carica papaya and Cocos nucifera.

Comments
Aonidiella aurantii is thought to be indigenous to South China (Longo et al., 1995), but the species has been dispersed widely by the transport of infested plant material by man. Davidson and Miller (1990) summarize its distribution as occurring in the tropics and in greenhouses in colder areas. Hodgson and Hilburn, 1991, remark that the species does not appear to have become established in Bermuda.

Europe
Cyprus: present, no further details (Danzig and Pellizzari, 1998)
France: present, no further details (Foldi, 2001)
Former USSR
Azerbaijan: present, no further details (Danzig and Pellizzari, 1998)
Black Sea coast: present, no further details (Nakahara, 1982)
Transcaucasus: present, no further details (Nakahara, 1982)
Former Yugoslavia: present, no further details (Danzig and Pellizzari, 1998)
Greece: present, no further details (IIE, 1996)
Crete: present, no further details (IIE, 1996)
Italy: in the south (Longo et al., 1995; IIE, 1996)
Sardinia: present, no further details (Longo et al., 1995)
Sicily: present, no further details (Longo et al., 1995)
Malta: present, no further details (IIE, 1996)
Portugal: present, no further details (Danzig and Pellizzari, 1998)
Madeira: present, no further details (IIE, 1996)
Romania: present, no further details (Danzig and Pellizzari, 1998)
Spain: present in Alicante, Madrid and Valencia (Amparo Blay Golcoechea, 1993; IIE, 1996)
Canary Islands: present, no further details (Amparo Blay Golcoechea, 1993; IIE, 1996)

Asia
Afghanistan: present, no further details (IIE, 1996)
Bangladesh: present, no further details (APPPC, 1987; IIE, 1996)
Bhutan: present, no further details (IIE, 1996)
Chagos Archipelago: present, no further details (IIE, 1996)
China
Beijing: present, no further details (IIE, 1996)
Fujian: present, no further details (Tao, 1999)
Guangdong: present, no further details (Tao, 1999)
Guangxi: present, no further details (IIE, 1996)
Hebei: present, no further details (IIE, 1996)
Hong Kong: present, no further details (Tao, 1999)
Hunan: present, no further details (IIE, 1996)
Inner Mongolia: present, no further details (Tao, 1999)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Shandong: present, no further details (Tao, 1999)
Sichuan: present, no further details (IIE, 1996)
Yunnan: present, no further details (Tao, 1999)
Zhejiang: present, no further details (IIE, 1996)
India
Andhra Pradesh: present, no further details (IIE, 1996)
Assam: present, no further details (IIE, 1996)
Bihar: present, no further details (IIE, 1996)
Delhi: present, no further details (IIE, 1996)
Himachal Pradesh: present, no further details (IIE, 1996)
Karnataka: present, no further details (IIE, 1996)
Kerala: present, no further details (IIE, 1996)
Madhya Pradesh: present, no further details (IIE, 1996)
Maharashtra: present, no further details (IIE, 1996)
Manipur: present, no further details (IIE, 1996)
Meghalaya: present, no further details (IIE, 1996)
Orissa: present, no further details (IIE, 1996)
Punjab: present, no further details (IIE, 1996)
Sikkim: present, no further details (IIE, 1996)
Tamil Nadu: present, no further details (IIE, 1996)
Tripura: present, no further details (IIE, 1996)
Uttar Pradesh: present, no further details (IIE, 1996)
West Bengal: present, no further details (IIE, 1996)
Indonesia
Irian Jaya: present, no further details (IIE, 1996)
Java: present, no further details (IIE, 1996)
Sumatra: present, no further details (IIE, 1996)
Iran: present, no further details (Seghatoleslami, 1977; Abivardi, 2001)
Iraq: present, no further details (IIE, 1996)
Israel: present, no further details (Rose, 1990; IIE, 1996)
Japan: present (Kawai, 1980; Tao, 1999)
Jordan: present, no further details (IIE, 1996)
Korea: present, no further details (Danzig and Pellizzari, 1998)
Kuwait: present, no further details (IIE, 1996)
Lebanon: present, no further details (IIE, 1996)
Malaysia
West Malaysia: present, no further details (IIE, 1996)
Sabah: present, no further details (IIE, 1996)
Sarawak: present, no further details (IIE, 1996)
Maldive Is: present, no further details (Watson et al., 1995)
Nepal: present, no further details (IIE, 1996)
Oman: The Natural History Museum collection, London, UK
Pakistan: present (IIE, 1996)
Philippines: present, no further details (Velasquez, 1971; IIE, 1996)
Saudi Arabia: present, no further details (IIE, 1996)
Sri Lanka: present, no further details (IIE, 1996)
Syria: present, no further details (Rose, 1990; IIE, 1996)
Taiwan: present, no further details (Wong et al., 1999)
Thailand: present, no further details (IIE, 1996)
Turkey: present, no further details (Rose, 1990; IIE, 1996)
Vietnam: present, no further details (IIE, 1996)
Yemen: present, no further details (IIE, 1996)

Africa
Algeria: present, no further details (IIE, 1996)
Angola: present, no further details (IIE, 1996)
Congo Democratic Republic: present, no further details (IIE, 1996)
Egypt: present, no further details (El-Minshawy et al., 1974a; IIE, 1996)
Ethiopia: present, no further details (IIE, 1996)
Guinea: present, no further details (IIE, 1996)
Kenya: present, no further details (IIE, 1996)
Madagascar: present, no further details (IIE, 1996)
Malawi: present, no further details (IIE, 1996)
Mauritius: present, no further details (Williams and Williams, 1988)
Morocco: present, no further details (IIE, 1996)
Mozambique: present, no further details (IIE, 1996)
Réunion: present, no further details (Williams and Williams, 1988)
South Africa: present, no further details (Bedford, 1990; IIE, 1996)
St Helena: present, no further details (IIE, 1996)
Sudan: present, no further details (IIE, 1996)
Swaziland: present, no further details (IIE, 1996)
Tanzania: present, no further details (Bohlen, 1973; IIE, 1996)
Tunisia: present, no further details (IIE, 1996)
Uganda: present, no further details (IIE, 1996)
Zambia: present, no further details (IIE, 1996)
Zimbabwe: present, no further details (IIE, 1996)

Western Hemisphere
Antigua: present, no further details (Rose, 1990; IIE, 1996)
Argentina: widespread and abundant (Claps et al., 2001a)
Corrientes: present, no further details (IIE, 1996)
Entre Rios: present, no further details (IIE, 1996)
Jujuy: present, no further details (IIE, 1996)
Misiones: present, no further details (IIE, 1996)
Paraná Delta: present, no further details (IIE, 1996)
Tucumán: present, no further details (Claps and Terán, 2001)
Bahamas: present, no further details (IIE, 1996)
Barbados: present, no further details (Bennett and Alam, 1985)
Bolivia: present, no further details (IIE, 1996)
Brazil: abundant (Claps et al., 2001a)
Alagôas: present, no further details (Claps et al., 2001a)
Bahia: present, no further details (IIE, 1996)
Ceará: present, no further details (Claps et al., 2001a; IIE, 1996)
Guanabara: present, no further details (Silva et al., 1968)
Maranhao: 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; IIE, 1996)
Paraná: present, no further details (Claps et al., 2001a)
Pernambuco: present, no further details (Claps et al., 2001a)
Rio Grande do Norte: present, no further details (Claps et al., 2001a; IIE, 1996)
Rio Grande do Sul: present, no further details (Claps et al., 2001a; IIE, 1996)
Rio de Janeiro: present, no further details (Claps et al., 2001a; IIE, 1996)
Santa Catarina: present, no further details (Claps et al., 2001a; IIE, 1996)
Sao Paulo: present, no further details (Claps et al., 2001a; IIE, 1996)
Chile: abundant (Claps et al., 2001a)
Antofagasta: present, no further details (Claps et al., 2001a)
Atacama: present, no further details (Claps et al., 2001a)
Coquimbo: present, no further details (Claps et al., 2001a)
Maule: present, no further details (Claps et al., 2001a)
O'Higgins: present, no further details (Claps et al., 2001a)
Santiago: present, no further details (Claps et al., 2001a)
Tarapacá: present, no further details (Claps et al., 2001a)
Valparaiso: present, no further details (Claps et al., 2001a)
Colombia: present, no further details (IIE, 1996)
Dominica: present, no further details (IIE, 1996)
Dominican Republic: present, no further details (Nakahara, 1982)
French Guiana: present, no further details (IIE, 1996)
Guadeloupe: present, no further details (IIE, 1996)
Guyana: present, no further details (IIE, 1996)
Honduras: present, no further details (IIE, 1996)
Jamaica: present, no further details (IIE, 1996)
Martinique: present, no further details (IIE, 1996)
Mexico: widespread (Miller, 1996; IIE, 1996)
Montserrat: present, no further details (IIE, 1996)
Nicaragua: present, no further details (Nakahara, 1982)
Panama: present, no further details (Nakahara, 1982)
Paraguay: present, no further details (Rose, 1990; IIE, 1996)
Puerto Rico: present, no further details (IIE, 1996)
St Lucia: present, no further details (IIE, 1996)
St Vincent: present, no further details (IIE, 1996)
Trinidad: present, no further details (IIE, 1996)
Uruguay: present, no further details (IIE, 1996)
USA
Alabama: present, no further details (Nakahara, 1982)
Arizona: present, no further details (IIE, 1996)
California: widespread (Gill, 1997)
District of Colombia: (under glass) (Nakahara, 1982)
Florida: present, no further details (IIE, 1996)
Georgia: present, no further details (Nakahara, 1982)
Kansas: present, no further details (Nakahara, 1982)
New Jersey: present, no further details (Nakahara, 1982)
Missouri: present, no further details (Nakahara, 1982)
Pennsylvania: present, no further details (Nakahara, 1982)
Oregon: present, no further details (Nakahara, 1982)
Texas: present, no further details (IIE, 1996)

Oceania
Australia
New South Wales: present, no further details (IIE, 1996; CSIRO, 2001)
Northern Territory: present, no further details (IIE, 1996; CSIRO, 2001)
Queensland: present, no further details (IIE, 1996; CSIRO, 2001)
South Australia: present, no further details (IIE, 1996; CSIRO, 2001)
Tasmania: present, no further details (IIE, 1996)
Victoria: present, no further details (IIE, 1996; CSIRO, 2001)
Western Australia: present, no further details (IIE, 1996; CSIRO, 2001)
Bonin Is: present, no further details (Beardsley, 1966)
Cook Islands: present, no further details (IIE, 1996)
Fiji: present, no further details (IIE, 1996)
Guam: present, no further details (Nakahara, 1982)
New Britain: present, no further details (IIE, 1996)
New Caledonia: present, no further details (IIE, 1996)
New Zealand: present, no further details (APPPC, 1987; Rose, 1990; IIE, 1996)
Niue: present, no further details (IIE, 1996)
Papua New Guinea: present, no further details (APPPC, 1987; IIE, 1996)
Russell Is: present, no further details (Nakahara, 1982)
Samoa: present, no further details (IIE, 1996)
Society Is: present, no further details (IIE, 1996)
Solomon Islands: present, no further details (APPPC, 1987; IIE, 1996)
Tonga: present, no further details (IIE, 1996)
Vanuatu: present, no further details (IIE, 1996)
Wallis Is: present, no further details (IIE, 1996)