(Signoret, 1869)
Diagnosis
In life, scale cover of adult female 1.5-2.0 mm diameter, circular, convex; on stems, usually tan but can be darker; on leaves, grey or almost white; exuviae subcentral, yellow to yellow-brown (Davidson and Miller, 1990) HEMLAL5.jpg . If present, male scale cover elongate oval with yellow subterminal exuviae, smaller and sometimes paler than that of female. Body of adult female bright yellow (Davidson and Miller, 1990; Gill, 1997) HEMLAL3.jpg . Eggs yellow, elongate, each 0.15 mm long.
Body of slide-mounted adult female membranous and broadly pyriform HEMLAS.jpg . Pygidium with large median lobes close together, almost touching; second and third lobes very reduced, unsclerotized HEMLAM2.jpg ; anal opening large, situated near posterior margin of pygidium (less than 2.3 times its length from base of median lobes); paraphyses shorter than the lobes, present only on the margin between the third lobes; perivulvar pores present HEMLAP.jpg .
Host range
Hemiberlesia lataniae is a highly polyphagous species that has been recorded from (mostly woody) hosts belonging to 278 genera in 78 plant families (Davidson and Miller, 1990). It prefers members of the families Rosaceae, Palmae and Euphorbiaceae, and evergreen hosts such as members of the coniferous families Cupressaceae (Cupressus, Juniperus, Thuja) and Pinaceae (Cedrus) (Zahradník, 1990). Hosts include species of: Abutilon, Acacia, Actinidia deliciosa, Albizia, Aleurites, Antigonon, Ardisia, Areca, Artocarpus heterophyllus, Asparagus, Atriplex, Averrhoa, Avicennia, Balaka, Bambusa, Barringtonia, Bauhinia, Begonia, Bignonia, Cactaceae, Camellia, Canavalia, Capsicum frutescens, Carya illinoinensis, Casimiroa, Cassia, Casuarina, Cayaponia, Ceiba pentandra, Chloris, Chrysanthemum, Chrysophyllum, Citrus spp., Cocos spp., Coffea arabica, Cordia, Cordyline, Cucurbita, Cycas, Cydonia, Cymbidium, Cyrtosperma, Diospyros, Dodonaea, Elaeagnus, Elaeis guineensis, Elaeocarpus tonganus, Elephantopus, Elymus, Eriobotrya, Eucalyptus, Eugenia, Euonymus, Feijoa, Ficus spp., Fitchia, Garcinia, Gladiolus, Gomphrena, Gossypium, Heliconia, Hibiscus, Jasminum, Juniperus, Lantana, Latania, Laurus nobilis, Leucaena, Litsea, Macadamia, Malus spp., Mangifera indica, Manihot esculenta, Manilkara zapota, Melastoma, Melia, Mespilus, Mimosa, Morinda, Morus alba, Muehlenbeckia, Musa, Myristica, Nerium, Olea europaea, Ophryosporus, Palmae, Pandanus, Parinari, Passiflora edulis, Pemphis, Persea americana, Phoenix dactylifera, Phytolacca, Piper, Pithecellobium, Plumeria, Populus, Prosopis, Prunus, Psidium guajava, Ptychosperma, Punica granatum, Pyrus communis, Ravenala, Rhizophora, Rhododendron, Rosa, Salix, Sapotaceae, Saurauia, Scaevola, Scalesia, Schefflera, Sida, Solanum spp., Tecoma, Timonius, Trachycarpus, Trithrinax, Veitchia, Viburnum, Vitis vinifera, Yucca and Zygogynum.
Affected plant stages: vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: all aerial parts HEMLAL1.jpg , but especially young branches HEMLAL4.jpg and leaves
Biology and ecology
In California, H. lataniae is apparently parthenogenetic and occurs at low elevations (Gill, 1997); in this population, development from egg to egg-laying female is 56-65 days (McKenzie, 1935). The species is also unisexual in Israel (Gerson and Zor, 1973).
In many tropical countries H. lataniae is bisexual and males are common. It is bisexual in Maryland, USA, where in the female, the first instar lasts for about 14 days and the second instar for 2-3 weeks, depending on environmental conditions; each female lays 15-25 eggs, and overwintering is as second instars; and there are two generations per year (Stoetzel and Davidson, 1974). In Egypt, there are three generations per year (El-Minshawy et al., 1971); and in Israel there are four (Gerson and Zor, 1973).
In South Africa, heavy infestations of H. lataniae have been observed on avocado fruit hanging near the ground, compared to higher up in the tree (Steyn, 1995); this may indicate some preference for shade.
Dispersal is by the active crawling of the first instar, and passive transport by wind and animal agents, including man. Dry weather favours dispersal and establishment; heavy rain causes high mortality of crawlers.
Symptoms
The presence of H. lataniae can be detected by abnormal colouring and some distortion of leaves, twigs and fruit, and as pitting of the bark of stems. Infested mango fruit turn pale green or yellow where the scales feed; the discoloration remains on the fruit skin even if the scales are removed. In South Africa, if there are more than ten of these spots per fruit on 10% or more of the shipment, it cannot be exported (Daneel, 1998). Heavy infestation can cause dieback of twigs and branches (Kosztarab, 1996).
According to Ebeling, 1959, feeding on fruit by H. lataniae irritates the flesh of the avocado cultivar Fuerte and possibly other thin-skinned cultivars, causing formation of nodules on the inside of the peel and corresponding depressions in the flesh beneath.
Economic impact
Hemiberlesia lataniae is considered to be a serious pest in many areas of the world, including Israel, the Palearctic region, the former USSR and USA (Florida and Hawaii) (Danzig and Pellizzari, 1998; Miller and Davidson, 1990). It is a pest of avocado, banana, Citrus, coconut, guava and mango in Brazil (Foldi, 1988; Chua and Wood, 1990; Claps et al., 2001a). On avocado in California, it causes culling or grade reduction of fruit due to pitting and other deformation of the outer flesh of thin-skinned varieties, and it is regarded as a serious pest of this crop in Israel. Hemiberlesia lataniae is a serious pest of palms and ornamentals in Florida and causes problems on ornamentals and kiwi fruit in California (Gill, 1997). In Argentina it is considered to be a pest on olives in several provinces, and damages them in Peru (Canales Canales and Valdivieso, 1999), Israel, Turkey, Egypt and seriously in Chile (Argyriou, 1990). In Chile H. lataniae is important on kiwifruit (Actinidia) in Maule province (Claps et al., 2001a). On grapevine in South Africa, it attacks the main stem, branches and shoots of the vines, causing dieback of bearing stems, as well as older wood and occasionally whole vines (Swart and De Klerk, 1986). It is common on tea in Sri Lanka (Muraleedharan, 1983), infesting the stem, branches and leaves, making the bushes weak and unproductive; persistent attacks may kill pruned bushes (Chua and Wood, 1990). Charles and Henderson, submitted, record H. lataniae as a pest on economically important plants in New Zealand, and that it occurs on native plants there also. Foldi, 2001, lists it as an economically important pest in France.
Detection and inspection methods
Inspect the stem, branches, leaves and fruit for the presence of greyish or brownish scale covers up to 2 mm long. A magnifying glass or hand lens and good lighting are important to ensure detection.
Phytosanitary risk
According to Burger and Ulenberg, 1990, H. lataniae is a quarantine pest in several countries. Introduction to new countries is often via importation of infested plant material, especially ornamental plants. Hemiberlesia lataniae is frequently intercepted at plant quarantine inspection (Williams and Watson, 1988).
Natural enemies
In most areas where H. lataniae is present, natural enemies usually control populations in field conditions. However, this will only occur where ants do not disturb the balance of natural enemies and pesticides are not applied indiscriminately. Stoetzel and Davidson, 1974, studied its natural enemies in Maryland.
Parasitoids:
- Aphytis spp., attacking: nymphs, adults, in Australia
- Aphytis chilensis, attacking: nymphs, adults, in Chile
- Aphytis melinus, attacking: nymphs, adults, in Australia (introduced)
- Aphytis mytilaspidis, in Crete
- Aphytis proclia
- Cheletomimus berlesei, attacking: nymphs, adults, in USA
- Comperiella lemniscata, attacking: nymphs, adults, in Australia (introduced)
- Habrolepis obscura
- Signiphora fax, attacking: nymphs, adults, in USA
- Signiphora flavella, attacking: nymphs, adults, in USA, Australia, Chile
- Signiphora perpauca, attacking: nymphs, adults, in USA, Australia
Predators:
- Cheletomimus berlesei
- Chilocorus sp.
- Chilocorus stigma
- Cycloneda sanguinea
- Dentifibula obtusilobae
- Eremochrysa californica
- Hemisarcoptes spp.
- Hemisarcoptes coccophagus, attacking all stages apart from eggs, widespread including Israel; Introduced: New Zealand
- Hemisarcoptes malus
- Rhyzobius lophanthae
- Rhyzobius pulchellus
- Signiphora spp.
- Watsoniella flavipes
Pathogens:
- Nectria flammea, attacking: nymphs, adults, in USA
Distribution
See Hemiberlesia lataniae distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species. Gill, 1997, says that H. lataniae cannot be separated from H. rapax in the field; H. palmae is also similar.
The leaf form of Diaspidiotus ancylus (Putnam) (convex scale, Putnam scale, cochenille de Putnam) DIASANS.jpg could be confused with H. lataniae, but in slide-mounted adult females, D. ancylus differs in possessing a relatively small anal opening situated more than 2.5x its length away from the bases of the median lobes DIASANP.jpg, and mature females often have slight sclerotization of the prosoma DIASANS.jpg. In contrast, H. lataniae has a relatively large anal opening situated less than 2.3x its length away from the bases of the median lobes HEMLAP.jpg , and the prosoma remains membranous throughout life HEMLAS.jpg . Diaspidiotus ancylus is a polyphagous species known from Canada (Ontario), USA (continental including Arizona, California, Indiana, Iowa, New Mexico, New York, Tennessee), Mexico, Central America, Brazil (Sao Paulo), Chile (first to eighth regions), Spain, Portugal, Germany, South Africa and Australia; it is often found on the bark of branches and trunk, and sometimes on leaves, of basswood, elm and maple. Hosts include species of Acer, Aesculus, Betula, Carya, Catalpa, Celtis, Cephalanthus, Crataegus, Cydonia, Fagus, Fraxinus, Hydrangea, Juglans, Liriodendron, Maclura, Malus sylvestris, Muehlenbeckia, Olea europaea, Oxalis, Persea americana, Populus spp., Prunus spp., Pyrus, Quercus, Ribes, Robinia spp., Salix, Sorbus, Tilia, Ulmus, Viburnum and Vitis vinifera (Munting, 1971a; Konstantinova, 1976; Nakahara, 1982; Zahradník, 1990a; Gill, 1997; Kosztarab, 1996; Miller, 1996; Danzig and Pellizzari, 1998; Claps et al., 2001a; CSIRO, 2001; The Natural History Museum collection, London, UK). Immature stages of D. ancylus can be mistaken for D. perniciosus (Claps et al., 2001a). This species has been a pest on walnuts, cranberries, blueberries and on elms and other ornamentals in the USA in the past; it is probably native to North America (Gill, 1997). Diaspidiotus ancylus was listed as a pest of deciduous fruit trees of regional importance by Kozár, 1990b; heavy infestations can kill twigs and branches (Kosztarab, 1996). One or two generations per year have been recorded in the USA, and overwintering is as adult females (second instar females in Chile) (Kosztarab, 1996). This species does not occur in the former USSR, but featured on the list of quarantine pests for this region (Konstantinova, 1976).
Comments
Hemiberlesia lataniae is one of the commonest cosmopolitan species of Diaspididae (Williams and Watson, 1988), but its area of origin is unknown (Gill, 1997). It was originally described from a palm, Latania borbonica, in 1869 (Quayle, 1938). This palm was a very popular ornamental plant which was distributed worldwide, carrying the pest with it. Now it is widespread throughout the tropical and subtropical areas of the world, and under glass in temperate regions (Nakahara, 1982).
Europe
Austria: establishment uncertain (The Natural History Museum collection, London, UK)
Bulgaria: present, no further details (CIE, 1976)
Cyprus: present, no further details (Danzig and Pellizzari, 1998)
Former USSR: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Russian Federation: present, no further details (Miller and Davidson, 1990)
Former Yugoslavia: present, no further details (Danzig and Pellizzari, 1998)
France: present, no further details (Danzig and Pellizzari, 1998; Foldi, 2001)
Germany: present, no further details (Danzig and Pellizzari, 1998)
Greece: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Crete: present, no further details (CIE, 1976)
Italy: present in the south (Longo et al., 1995; Danzig and Pellizzari, 1998)
Sicily: present, no further details (Longo et al., 1995)
Malta: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Portugal: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Madeira: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Romania: present, no further details (CIE, 1976)
Spain: fairly widespread (Amparo Blay Golcoechea, 1993)
Canary Is: present, no further details (Amparo Blay Golcoechea, 1993; Danzig and Pellizzari, 1998)
United Kingdom: restricted to a few botanical collections under glass (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
England: The Natural History Museum collection, London, UK
Wales: (C.P. Malumphy, Central Science Laboratory, UK, pers. comm.)
Asia
Bangladesh: The Natural History Museum collection, London, UK
Brunei Darussalam: The Natural History Museum collection, London, UK
Chagos Archipelago: The Natural History Museum collection, London, UK
China
Fujian: present, no further details (Tao, 1999)
Guangdong: present, no further details (Tao, 1999)
Guangxi: present, no further details (Tao, 1999)
Hong Kong: present, no further details (CIE, 1976)
Hubei: present, no further details (CIE, 1976)
Inner Mongolia: present, no further details (Tao, 1999)
Jiangsu: present, no further details (Tao, 1999)
Jiangxi: present, no further details (Tao, 1999)
Yunnan: present, no further details (Tao, 1999)
Zhejiang: present, no further details (Tao, 1999)
India
Andaman and Nicobar Is: The Natural History Museum collection, London, UK
Andhra Pradesh: present, no further details (Dhileepan, 1991)
Assam: The Natural History Museum collection, London, UK
Bihar: present, no further details (CIE, 1976)
Gujarat: The Natural History Museum collection, London, UK
Karnataka: present, no further details (Dhileepan, 1991)
Kerala: present, no further details (Dhileepan, 1991)
Maharashtra: present, no further details (Dhileepan, 1991)
Orissa: The Natural History Museum collection, London, UK
Sikkim: The Natural History Museum collection, London, UK
Tamil Nadu: present, no further details (CIE, 1976)
West Bengal: present, no further details (CIE, 1976)
Indonesia: present, no further details (CIE, 1976)
Irian Jaya: present (Williams and Watson, 1988)
Java: present, no further details (CIE, 1976)
Sumatra: present, no further details (CIE, 1976)
Iran: present, no further details (Seghatoleslami, 1977; Danzig and Pellizzari, 1998)
Iraq: present, no further details (Danzig and Pellizzari, 1998)
Israel: present, no further details (Argyriou, 1990; Izraylevich et al., 1995)
Japan: present, no further details (Kawai, 1980; Nagarkatti and Sankaran, 1990; Tao, 1999)
Honshu: present, no further details (CIE, 1976)
Jordan: present, no further details (CIE, 1976)
Lebanon: present, no further details (CIE, 1976)
Malaysia: present, no further details (Chua and Wood, 1990)
West Malaysia: present, no further details (CIE, 1976)
Maldive Is: present, no further details (Watson et al., 1995)
Oman: The Natural History Museum collection, London, UK
Pakistan: present, no further details (CIE, 1976)
Philippines: present, no further details (Velasquez, 1971)
Singapore: present, no further details (CIE, 1976)
Sri Lanka: present, no further details (Nagarkatti and Sankaran, 1990; Chua and Wood, 1990)
Syria: present, no further details (CIE, 1976)
Taiwan: present, no further details (Wang and Su, 1989; Wong et al., 1999)
Thailand: present, no further details (CIE, 1976)
Turkey: present, no further details (Argyriou, 1990; Danzig and Pellizzari, 1998)
Africa
Aldabra: The Natural History Museum collection, London, UK
Algeria: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Angola: present, no further details (CIE, 1976)
Ascension I.: The Natural History Museum collection, London, UK
Benin: present, no further details (CIE, 1976)
Cameroon: present, no further details (CIE, 1976)
Cape Verde: present, no further details (CIE, 1976)
Congo Democratic Republic: present, no further details (CIE, 1976)
Côte d'Ivoire: present, no further details (CIE, 1976)
Egypt: present, no further details (El-Minshawy et al., 1974a; Williams and Greathead, 1990)
Eritrea: present, no further details (CIE, 1976)
Ethiopia: present, no further details (CIE, 1976)
Ghana: present, no further details (CIE, 1976)
Guinea: present, no further details (CIE, 1976)
Kenya: present, no further details (CIE, 1976; Nagarkatti and Sankaran, 1990)
Libya: present, no further details (CIE, 1976)
Madagascar: present, no further details (CIE, 1976)
Malawi: present, no further details (CIE, 1976)
Mali: present, no further details (CIE, 1976)
Mauritius: present, no further details (CIE, 1976; Williams and Williams, 1988)
Morocco: present, no further details (CIE, 1976; Danzig and Pellizzari, 1998)
Mozambique: present, no further details (CIE, 1976)
Nigeria: present, no further details (CIE, 1976)
Principe: present, no further details (CIE, 1976)
Réunion: present, no further details (CIE, 1976; Williams and Williams, 1988)
Rodrigues: present, no further details (CIE, 1976; Williams and Williams, 1988)
Sao Tomé: present, no further details (CIE, 1976)
Seychelles: present, no further details (CIE, 1976)
Sierra Leone: present, no further details (CIE, 1976)
Somalia: present, no further details (CIE, 1976)
South Africa: widespread (Mitchell and Ironside, 1982; Daneel et al., 1994; Daneel, 1998)
St Helena: present, no further details (CIE, 1976)
Sudan: present, no further details (CIE, 1976)
Tanzania: present, no further details (CIE, 1976; Mitchell and Ironside, 1982)
Zanzibar: present, no further details (Quayle, 1938; The Natural History Museum collection, London, UK)
Togo: present, no further details (CIE, 1976)
Tunisia: present, no further details (CIE, 1976)
Uganda: present, no further details (CIE, 1976)
Zambia: The Natural History Museum collection, London, UK
Zimbabwe: present, no further details (CIE, 1976)
Western Hemisphere
Argentina: frequent (Claps et al., 2001a)
Buenos Aires: present, no further details (Claps et al., 2001a)
Corrientes: present, no further details (Claps et al., 2001a)
Entre Rios: present, no further details (Claps et al., 2001a)
Mendoza: present, no further details (Claps et al., 2001a)
Misiones: present, no further details (Claps et al., 2001a)
Salta: present, no further details (Claps et al., 2001a)
Santiago del Estero: present, no further details (Claps et al., 2001a)
Tucumán: present, no further details (Claps et al., 2001a)
Bahamas: present, no further details (CIE, 1976)
Barbados: present, no further details (CIE, 1976; Bennett and Alam, 1985)
Bermuda: common and widespread (Hodgson and Hilburn, 1991)
Brazil: widely distributed (Claps et al., 2001a)
Amazonas: present, no further details (Foldi, 1988; Claps et al., 2001a)
Espirito Santo: present, no further details (CIE, 1976; Foldi, 1988; Claps et al., 2001a)
Guanabara: present, no further details (Foldi, 1988)
Minas Gerais: present, no further details (CIE, 1976; Foldi, 1988; Claps et al., 2001a)
Pará: present, no further details (CIE, 1976; Foldi, 1988)
Paraná: present, no further details (Foldi, 1988; Claps et al., 2001a)
Rio de Janeiro: present, no further details (Foldi, 1988; Claps et al., 2001a)
Rio Grande do Sul: present, no further details (Foldi, 1988; Claps et al., 2001a)
Sao Paulo: present, no further details (Foldi, 1988; Claps et al., 2001a)
Chile
Antofagasta: present, no further details (Claps et al., 2001a)
Atacama: present, no further details (Claps et al., 2001a)
Coqimbo: present, no further details (Claps et al., 2001a)
Easter Island: present, no further details (Charlín, 1973; 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 (Kondo, 2001)
Costa Rica: The Natural History Museum collection, London, UK
Cuba: present, no further details (CIE, 1976)
Dominica: present, no further details (CIE, 1976)
Ecuador: The Natural History Museum collection, London, UK
Galapagos Is: present, no further details (CIE, 1976)
Guatemala: present, no further details (CIE, 1976)
Guyana: present, no further details (CIE, 1976)
Haiti: present, no further details (CIE, 1976)
Honduras: present, no further details (CIE, 1976)
Jamaica: present, no further details (CIE, 1976)
Mexico: widespread (CIE, 1976; Miller, 1996)
Nicaragua: The Natural History Museum collection, London, UK
Panama: present, no further details (CIE, 1976)
Peru: present, no further details (CIE, 1976; Canales Canales and Valdivieso, 1999)
Puerto Rico: present, no further details (CIE, 1976; Medina Gaud et al., 1987)
Trinidad: present, no further details (CIE, 1976)
USA
Alabama: present, no further details (Nakahara, 1982)
Arkansas: present, no further details (Nakahara, 1982)
Arizona: present, no further details (Nakahara, 1982)
California: widespread at low altitudes (Gill, 1997)
Colorado: present, no further details (Nakahara, 1982)
District of Colombia: present, no further details (Nakahara, 1982)
Delaware: present, no further details (Nakahara, 1982)
Florida: present, no further details (Dekle, 1976; Howard and Oliver, 1985; Miller and Davidson, 1990)
Georgia: present, no further details (Nakahara, 1982)
Hawaii: present on Oahu, Maui, Kauai and Molokai (Heu, 2002)
Iowa: present, no further details (Nakahara, 1982)
Illinois: present, no further details (Nakahara, 1982)
Indiana: present, no further details (Nakahara, 1982)
Kansas: present, no further details (Nakahara, 1982)
Louisiana: present, no further details (CIE, 1976; Howard and Oliver, 1985)
Massachusetts: present, no further details (Nakahara, 1982)
Maryland: present, no further details (Stoetzel and Davidson, 1974)
Michigan: present, no further details (Nakahara, 1982)
Missouri: present, no further details (Mitchell and Ironside, 1982)
Mississippi: present, no further details (Nakahara, 1982)
North Carolina: present, no further details (Nakahara, 1982)
New Jersey: present, no further details (Nakahara, 1982)
New Mexico: The Natural History Museum collection, London, UK
New York: present, no further details (Nakahara, 1982)
Oklahoma: present, no further details (Nakahara, 1982)
Pennsylvania: present, no further details (Nakahara, 1982)
South Carolina: present, no further details (Nakahara, 1982)
Tennessee: present, no further details (Nakahara, 1982)
Texas: present, no further details (Nakahara, 1982)
Virginia: present, no further details (CIE, 1976; Nakahara, 1982)
Virgin Islands: present, no further details (CIE, 1976)
Venezuela: present, no further details (CIE, 1976)
Oceania
Australia
New South Wales: present, no further details (CSIRO, 2001)
Northern Territory: The Natural History Museum collection, London, UK
Queensland: present, no further details (Mitchell and Ironside, 1982; CSIRO, 2001)
Victoria: The Natural History Museum collection, London, UK
Bonin Is: present, no further details (Beardsley, 1966)
Caroline Islands: present, no further details (CIE, 1976)
Cook Is: present (Williams and Watson, 1988)
Fiji: present (Williams and Watson, 1988; Chua and Wood, 1990)
Gambier Is: present (Williams and Watson, 1988)
Johnston Island: present, no further details (CIE, 1976)
Kiribati: present (Williams and Watson, 1988)
Kusaie: present, no further details (Beardsley, 1966)
Marquesas Is: present (Williams and Watson, 1988)
Marshall Is: present, no further details (Beardsley, 1966)
New Caledonia: present (CIE, 1976; Williams and Watson, 1988)
New Zealand: present, no further details (Hill et al., 1993; Charles et al., 1995; Charles and Henderson, submitted)
Niue: present, no further details (Williams and Watson, 1988)
Palau: present, no further details (Beardsley, 1966)
Papua New Guinea: present (Williams and Watson, 1988; Chua and Wood, 1990)
Pohnpei: present, no further details (Beardsley, 1966)
Samoa: present, no further details (CIE, 1976; Williams and Watson, 1988)
Solomon Is: present (CIE, 1976; Williams and Watson, 1988)
South Mariana Is: present (Beardsley, 1966)
Tahiti: present, not further details (Doanne and Hadden, 1909)
Tokelau: present (Williams and Watson, 1988)
Tonga: present (CIE, 1976; Williams and Watson, 1988)
Truk Is: present (Beardsley, 1966)
Tuvalu: present (Williams and Watson, 1988)
Vanuatu: present (Williams and Watson, 1988)
Volcano Is: present, no further details (Beardsley, 1966)
Wake I.: present, no further details (Beardsley, 1966)
Western Samoa: present (Williams and Watson, 1988)
Yap Is: present, no further details (Beardsley, 1966)