Invasive Species Compendium

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Datasheet

Aspidiotus destructor
(coconut scale)

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Datasheet

Aspidiotus destructor (coconut scale)

Summary

  • Last modified
  • 10 December 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Aspidiotus destructor
  • Preferred Common Name
  • coconut scale
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Aspidiotus destructor (coconut scale); slide mounted adult.
TitleAdult
CaptionAspidiotus destructor (coconut scale); slide mounted adult.
Copyright©Natural History Museum, London
Aspidiotus destructor (coconut scale); slide mounted adult.
AdultAspidiotus destructor (coconut scale); slide mounted adult.©Natural History Museum, London
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Cocos nucifera, Solomon Islands. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
TitleAdult female
CaptionAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Cocos nucifera, Solomon Islands. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Copyright©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Cocos nucifera, Solomon Islands. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Adult femaleAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Cocos nucifera, Solomon Islands. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Carica papaya, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
TitleAdult female
CaptionAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Carica papaya, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Copyright©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Carica papaya, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Adult femaleAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Carica papaya, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Persea americana, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
TitleAdult female
CaptionAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Persea americana, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Copyright©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Persea americana, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Adult femaleAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Persea americana, Fiji. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Citrus maxima, Western Samoa. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
TitleAdult female
CaptionAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Citrus maxima, Western Samoa. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Copyright©CAB International
Aspidiotus destructor (coconut scale); anatomic details of an adult female taken from Citrus maxima, Western Samoa. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).
Adult femaleAspidiotus destructor (coconut scale); anatomic details of an adult female taken from Citrus maxima, Western Samoa. 1: General aspect. 2: Pygidium. 3: Antenna. 4: Anterior spiracle (not to scale).©CAB International

Identity

Top of page

Preferred Scientific Name

  • Aspidiotus destructor Signoret, 1869

Preferred Common Name

  • coconut scale

Other Scientific Names

  • Aspidiotus cocotis Newstead, 1893
  • Aspidiotus lataniae Green, 1896
  • Aspidiotus simillimus translucens Fernald
  • Aspidiotus translucens Cockerell & Robinson, 1915
  • Aspidiotus transparens Green, 1890
  • Aspidiotus vastatrix Leroy
  • Aspidiotus watanabei Takagi, 1969
  • Temnaspidiotus destructor (Signoret) Borchsenius

International Common Names

  • English: bourbon aspidiotus; bourbon scale; transparent scale
  • Spanish: cochinilla blanca-amarilla del coco; cochinilla del cocotero; escama blanca del cocotero; escama del cocotero; escama del pino; escama transparente
  • French: cochenille du cocotier

Local Common Names

  • Germany: Schildlaus, Kokospalmen-
  • Italy: Cocciniglia del cocco

EPPO code

  • ASPDDE (Aspidiotus destructor)
  • ASPDTR (Aspidiotus transparens)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hemiptera
  •                         Suborder: Sternorrhyncha
  •                             Unknown: Coccoidea
  •                                 Family: Diaspididae
  •                                     Genus: Aspidiotus
  •                                         Species: Aspidiotus destructor

Notes on Taxonomy and Nomenclature

Top of page Aspidiotus destructor was first described by Signoret in 1869. Williams and Watson (1988) list synonyms and discuss nomenclature.

Description

Top of page Jalaluddin and Mohanasundaram (1992) describe the morphology of different instars and the adult female and male of A. destructor. Williams and Watson (1988) provide a key and give detailed descriptions and illustrations of adult female morphology. Considerable variation occurs in the relative sizes and number of distinguishing features, such as median and second lobes, macroduct number and marginal setae (Williams and Watson, 1988).

Egg

The eggs are yellow and very small. They are laid under the scale around the body of the female.

Larva and Pupa

Females have two nymphal stages, while males have two feeding nymphal stages, followed by non-feeding pre-pupal and pupal stages (four immature stages altogether) (Tabibullah and Gabriel, 1973).

The first-instar larvae are about 1mm long, yellowish-brown, oval and translucent. Second-instar females become immobile and secrete a translucent wax scale cover. The second-instar males are smaller than the females. They group together, secrete a filamentous waxy material and become immobile. The male pre-pupal and pupal stages are spent under the scale produced by the second instar stage.

Adults

The scale cover of the adult female is oval to circular, 1.5-2.0 mm across, fairly flat, very thin and translucent. The pale yellow exuviae are more or less central on the scale (Williams and Watson, 1988). The yellow adult female under the scale is 0.6-1.1 mm long.

The adult male scale cover is redder than the female's, smaller and more oval (Williams and Watson, 1988). The male has one pair of wings and is motile.

Distribution

Top of page A. destructor apparently originated in the Pacific islands (Burger and Ulenberg, 1990) but is now recorded in tropical and subtropical regions worldwide. It is present in nearly all countries where coconuts are grown. In the northern parts of its range, it is found only under glass (Danzig and Pellizzari, 1998). It has been recorded under glass at a few botanic gardens in the UK (C Malumphy, Central Science Laboratory, UK, personal communication).

Distribution Table

Top of page

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 10 Jan 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AngolaPresentUK, CAB International (1966)
BeninPresentUK, CAB International (1966)
BurundiPresentUK, CAB International (1966)
Cabo VerdePresentUK, CAB International (1966)
CameroonPresentUK, CAB International (1966)
Congo, Democratic Republic of thePresentBuyckx (1962)
Congo, Republic of thePresentUK, CAB International (1966)
Côte d'IvoirePresentUK, CAB International (1966)
EgyptPresentUK, CAB International (1966); Danzig and Pellizzari (1998)
EritreaPresentUK, CAB International (1966)
EthiopiaPresentUK, CAB International (1966)
GhanaPresentUK, CAB International (1966)
GuineaPresentUK, CAB International (1966)
Guinea-BissauPresentUK, CAB International (1966)
KenyaPresentUK, CAB International (1966)
MadagascarPresentUK, CAB International (1966)
MauritaniaPresentUK, CAB International (1966)
MauritiusPresentUK, CAB International (1966); CABI (Undated)
MozambiquePresentUK, CAB International (1966)
NigeriaPresentUK, CAB International (1966)
RéunionPresentUK, CAB International (1966)
RwandaPresentUK, CAB International (1966)
São Tomé and PríncipePresentUK, CAB International (1966); Fernandes (1974)
SenegalPresentUK, CAB International (1966)
SeychellesPresentUK, CAB International (1966)
Sierra LeonePresentUK, CAB International (1966)
SomaliaPresentUK, CAB International (1966)
South AfricaPresentUK, CAB International (1966)
SudanPresentUK, CAB International (1966)
TanzaniaPresentUK, CAB International (1966)
-Zanzibar IslandPresentUK, CAB International (1966)
TogoPresentUK, CAB International (1966)
UgandaPresentUK, CAB International (1966)
ZambiaPresentUK, CAB International (1966)
ZimbabwePresentNHM (1957)

Asia

AzerbaijanPresentUK, CAB International (1966); Danzig and Pellizzari (1998)
BangladeshPresentAPPPC (1987)
BhutanPresentNHM (1985)
British Indian Ocean TerritoryPresentUK, CAB International (1966)
BruneiPresentWaterhouse (1993)
CambodiaPresentUK, CAB International (1966); Waterhouse (1993)
ChinaPresentCABI (Undated a)Present based on regional distribution.
-FujianPresentUK, CAB International (1966); Tao (1999)
-GuangdongPresentUK, CAB International (1966); Tao (1999)
-GuangxiPresentUK, CAB International (1966); Tao (1999)
-HainanPresentUK, CAB International (1966); Tao (1999)
-HubeiPresentTao (1999)
-HunanPresentZhou et al. (1993)
-JiangsuPresentUK, CAB International (1966)
-JiangxiPresentTao (1999)
-ShandongPresentUK, CAB International (1966); Tao (1999)
-SichuanPresentTao (1999)
-ZhejiangPresentUK, CAB International (1966); Tao (1999)
GeorgiaPresentUK, CAB International (1966); Danzig and Pellizzari (1998)
Hong KongPresentUK, CAB International (1966)
IndiaPresentCABI (Undated a)Present based on regional distribution.
-Andaman and Nicobar IslandsPresentNHM (1992)
-Andhra PradeshPresentUK, CAB International (1966)
-AssamPresentUK, CAB International (1966)
-BiharPresentUK, CAB International (1966)
-GujaratPresentUK, CAB International (1966)
-Jammu and KashmirPresentUK, CAB International (1966)
-KarnatakaPresentUK, CAB International (1966)
-KeralaPresentUK, CAB International (1966)
-LakshadweepPresentNHM (1940); UK, CAB International (1966)
-Madhya PradeshPresentUK, CAB International (1966)
-MaharashtraPresentUK, CAB International (1966)
-OdishaPresentUK, CAB International (1966)
-PunjabPresentUK, CAB International (1966)
-SikkimPresentUK, CAB International (1966)
-Tamil NaduPresentUK, CAB International (1966)
-Uttar PradeshPresentUK, CAB International (1966)
-West BengalPresentUK, CAB International (1966)
IndonesiaPresentWaterhouse (1993)
-Irian JayaPresentCABI (Undated)Original citation: Williams and Watson (1988)
-JavaPresentUK, CAB International (1966)
-Lesser Sunda IslandsPresentUK, CAB International (1966)
-SumatraPresentUK, CAB International (1966)
IranPresentUK, CAB International (1966); Danzig and Pellizzari (1998)
JapanPresentKawai (1980)
-Bonin IslandsPresentNakahara (1982)
-HonshuPresentUK, CAB International (1966)
MalaysiaPresentWaterhouse (1993)
-Peninsular MalaysiaPresentUK, CAB International (1966)
-SabahPresentUK, CAB International (1966)
-SarawakPresentUK, CAB International (1966)
MaldivesPresentWatson et al. (1995)
MyanmarPresentUK, CAB International (1966); Waterhouse (1993)
NepalPresentNHM (1967)
North KoreaPresentDanzig and Pellizzari (1998)
OmanPresent, WidespreadKinawy (1991)
PakistanPresentUK, CAB International (1966)
PhilippinesPresentUK, CAB International (1966); Velasquez (1971); Waterhouse (1993)
Saudi ArabiaPresentNHM (1969); Danzig and Pellizzari (1998)
SingaporePresentAPPPC (1987); Waterhouse (1993)
Sri LankaPresentUK, CAB International (1966)
TaiwanPresentNHM (1930); Takagi (1969); Wong et al. (1999)
ThailandPresentAPPPC (1987); Waterhouse (1993)
VietnamPresentUK, CAB International (1966); Waterhouse (1993)
YemenPresentNHM (1958)

Europe

FrancePresentGermain and Matile-Ferrero (2005)
GermanyPresentDanzig and Pellizzari (1998)
ItalyPresentLongo et al. (1995)
PortugalPresentCABI (Undated a)Present based on regional distribution.
-MadeiraPresentUK, CAB International (1966)
RussiaPresentUK, CAB International (1966); Danzig and Pellizzari (1998)
SpainPresentCABI (Undated a)Present based on regional distribution.
-Canary IslandsPresentUK, CAB International (1966)

North America

Antigua and BarbudaPresentUK, CAB International (1966)
BahamasPresentNHM (1968)
BarbadosPresentUK, CAB International (1966); Bennett and Alam (1985)
BelizePresentUK, CAB International (1966)
Cayman IslandsPresentUK, CAB International (1966)
Costa RicaPresentUK, CAB International (1966)
CubaPresentUK, CAB International (1966)
DominicaPresentUK, CAB International (1966)
Dominican RepublicPresentUK, CAB International (1966)
GrenadaPresentUK, CAB International (1966)
GuadeloupePresentUK, CAB International (1966)
GuatemalaPresentUK, CAB International (1966)
HaitiPresentUK, CAB International (1966)
HondurasPresentUK, CAB International (1966)
JamaicaPresentUK, CAB International (1966)
MartiniquePresentUK, CAB International (1966)
MexicoPresentUK, CAB International (1966); Miller (1996)
MontserratPresentUK, CAB International (1966)
Netherlands AntillesPresentUK, CAB International (1966)
NicaraguaPresentUK, CAB International (1966)
PanamaPresentUK, CAB International (1966)
Puerto RicoPresentUK, CAB International (1966)
Saint Kitts and NevisPresentUK, CAB International (1966)
Saint LuciaPresentUK, CAB International (1966)
Saint Vincent and the GrenadinesPresentUK, CAB International (1966)
Trinidad and TobagoPresentUK, CAB International (1966)
U.S. Virgin IslandsPresentUK, CAB International (1966)
United StatesPresentCABI (Undated a)Present based on regional distribution.
-ConnecticutPresentNakahara (1982)
-FloridaPresentUK, CAB International (1966)
-GeorgiaPresentNakahara (1982)
-HawaiiPresentCABI (Undated)Original citation: Heu, 2002
-PennsylvaniaPresentNakahara (1982)

Oceania

American SamoaPresentUK, CAB International (1966); CABI (Undated);
AustraliaPresentCABI (Undated a)Present based on regional distribution.
-Northern TerritoryPresentUK, CAB International (1966); CSIRO (2001)
-QueenslandPresentCSIRO (2001)
Federated States of MicronesiaPresentSuta and Esguerra (1993); CABI (Undated)
FijiPresent, WidespreadUK, CAB International (1966); CABI (Undated);
French PolynesiaPresent, WidespreadUK, CAB International (1966); CABI (Undated);
GuamPresentUK, CAB International (1966); CABI (Undated)
Marshall IslandsPresentUK, CAB International (1966); CABI (Undated)
New CaledoniaPresentUK, CAB International (1966); CABI (Undated)
Northern Mariana IslandsPresentUK, CAB International (1966); CABI (Undated)
PalauPresentUK, CAB International (1966); CABI (Undated)
Papua New GuineaPresentUK, CAB International (1966); CABI (Undated)
SamoaPresent, WidespreadUK, CAB International (1966); CABI (Undated)
Solomon IslandsPresentUK, CAB International (1966); CABI (Undated)
TuvaluPresent, LocalizedIPPC (2005); CABI (Undated)
VanuatuPresentCABI (Undated)Original citation: NAPPO 15:2
Wallis and FutunaPresentUK, CAB International (1966); CABI (Undated)

South America

BrazilPresentCABI (Undated a)Present based on regional distribution.
-AlagoasPresentUK, CAB International (1966); Foldi (1988)
-AmazonasPresentFoldi (1988); Claps et al. (2001)
-BahiaPresentUK, CAB International (1966); Claps et al. (2001)
-CearaPresentUK, CAB International (1966); Claps et al. (2001)
-Fernando de NoronhaPresentFoldi (1988); Claps et al. (2001)
-MaranhaoPresentFoldi (1988); Claps et al. (2001)
-ParaPresentFoldi (1988); Claps et al. (2001)
-ParaibaPresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-PernambucoPresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-PiauiPresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-Rio de JaneiroPresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-Rio Grande do NortePresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-Santa CatarinaPresentFoldi (1988); Claps et al. (2001)
-Sao PauloPresentUK, CAB International (1966); Foldi (1988); Claps et al. (2001)
-SergipePresentUK, CAB International (1966)
ChilePresentCABI (Undated a)Present based on regional distribution.
-Easter IslandPresentCharlin C. (1973); Claps et al. (2001)
ColombiaPresentUK, CAB International (1966); Kondo (2001)
EcuadorPresentUK, CAB International (1966); Kondo (2001)
-Galapagos IslandsPresentCABI (Undated)Original citation: CIE, 1966
GuyanaPresentUK, CAB International (1966)
PeruPresentUK, CAB International (1966)
SurinamePresentUK, CAB International (1966)
VenezuelaPresentUK, CAB International (1966)

Hosts/Species Affected

Top of page A. destructor is a highly polyphagous species. Davidson and Miller (1990) recorded it from hosts belonging to 75 genera in 44 plant families, but its host range is probably wider than this. Its hosts are typically perennial species and include many species of fruit trees, such as avocado, breadfruit, mango, guava and papaya.

Coconut is its favourite host; the undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. Older trees (over 4 years) or trees on well-drained soil are seldom seriously infested.

Host Plants and Other Plants Affected

Top of page
Plant nameFamilyContext
ActinidiaActinidiaceaeMain
AleuritesEuphorbiaceaeOther
AllamandaApocynaceaeOther
AlpiniaZingiberaceaeOther
AnnonaAnnonaceaeOther
Annona cherimola (cherimoya)AnnonaceaeOther
Annona muricata (soursop)AnnonaceaeOther
Artocarpus altilis (breadfruit)MoraceaeOther
BrassicaBrassicaceaeOther
CamelliaTheaceaeOther
Camellia sinensis (tea)TheaceaeOther
Capsicum (peppers)SolanaceaeOther
Carica papaya (pawpaw)CaricaceaeOther
Cassia (sennas)FabaceaeOther
Ceiba pentandra (kapok)BombacaceaeOther
Cinnamomum verum (cinnamon)LauraceaeOther
CitrusRutaceaeOther
Cocos nucifera (coconut)ArecaceaeMain
Colocasia esculenta (taro)AraceaeOther
Cucumis (melons, cucuimbers, gerkins)CucurbitaceaeOther
Dioscorea (yam)DioscoreaceaeOther
Elaeis guineensis (African oil palm)ArecaceaeMain
Eucalyptus deglupta (kamarere)MyrtaceaeOther
EugeniaMyrtaceaeOther
Euphorbia (spurges)EuphorbiaceaeOther
FicusMoraceaeOther
Ficus carica (common fig)MoraceaeOther
Hevea brasiliensis (rubber)EuphorbiaceaeOther
Hibiscus (rosemallows)MalvaceaeOther
Jasminum (jasmine)OleaceaeOther
Mangifera indica (mango)AnacardiaceaeMain
Musa (banana)MusaceaeMain
Myristica fragrans (nutmeg)MyristicaceaeOther
Pandanus (screw-pine)PandanaceaeOther
Passiflora (passionflower)PassifloraceaeOther
Persea americana (avocado)LauraceaeOther
Phoenix (date palm)ArecaceaeOther
Phoenix dactylifera (date-palm)ArecaceaeOther
Physalis (Groundcherry)SolanaceaeOther
Piper (pepper)PiperaceaeOther
Piper betle (betel pepper)PiperaceaeOther
Piper nigrum (black pepper)PiperaceaeOther
Plumeria (frangipani)ApocynaceaeOther
Prunus persica (peach)RosaceaeOther
Psidium guajava (guava)MyrtaceaeOther
Raphanus (radish)BrassicaceaeOther
Rhizophora (mangrove)RhizophoraceaeOther
Saccharum officinarum (sugarcane)PoaceaeOther
Solanum (nightshade)SolanaceaeOther
Solanum lycopersicum (tomato)SolanaceaeOther
Spondias purpurea (red mombin)AnacardiaceaeOther
Syzygium aromaticum (clove)MyrtaceaeOther
Tamarindus indica (Indian tamarind)FabaceaeOther
Theobroma cacao (cocoa)MalvaceaeOther
Vigna unguiculata (cowpea)FabaceaeOther
Vitis vinifera (grapevine)VitaceaeOther
Xanthosoma sagittifolium (elephant ear)AraceaeOther
Zingiber officinale (ginger)ZingiberaceaeOther

Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

Top of page On leaves, A. destructor causes yellow spots to develop beneath the insects, due to the toxicity of saliva injected in to plant tissues while feeding. Entire leaves may turn yellow to brown and fall, and fruits may be discoloured, stunted or fall prematurely. The bright yellow colour of affected coconut palms is clearly visible from a great distance. The undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. In extreme cases, the leaves dry out, entire fronds drop off and the crown dies.

List of Symptoms/Signs

Top of page
SignLife StagesType
Fruit / discoloration
Fruit / external feeding
Fruit / lesions: black or brown
Leaves / abnormal colours
Leaves / abnormal leaf fall
Leaves / necrotic areas
Stems / external feeding

Biology and Ecology

Top of page A. destructor reproduces sexually. Males locate unmated females by following pheromones released by them. The life cycle of A. destructor typically lasts for 32-34 days. In one study the life cycle was found to be 32 days for females and 27 days for males (Tabibullah and Gabriel, 1973; Taylor, 1935, also did an in-depth study).

Each female deposits 20-50 eggs under her scale cover over a few days. In China on Actinidia, the average number of eggs laid by one female was 32-42 (Zhou et al., 1993). At room temperature (26-28°C), the egg stage lasted for 5 days, the larval stage lasted 17 days, the pre-oviposition stage in adult females lasted 25 days, the female generation lasted 44 days and the male generation lasted 38 days (Zhou et al., 1993). In the Philippines, on coconut, the egg stage lasted for 8 days in both sexes (Tabibullah and Gabriel, 1973). After hatching, the nymphs crawl under the scale edge out into the open and colonize the undersides of leaves and tender shoots. They drop off the leaves easily, so mortality is high during heavy rain.

In China, A. destructor produced three generations annually, with the fertilized females overwintering on the stems of Actinidia trees (Zhou et al., 1993). In Japan on tea plants, A. destructor had only one generation per year (Murakami, 1970). However, in tropical conditions in Trinidad reproduction is continuous (Goberdhan, 1962).

The dispersal phase of A. destructor is the first instar, or crawler, which has legs. Crawlers can walk up to perhaps 1 m, but can be distributed across much greater distances by wind, flying insects and birds and transport of infested plant material by man.

Natural enemies

Top of page
Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aleurodothrips fasciapennis Predator Adults/Eggs/Nymphs Fiji Cocos nucifera
Anicetus communis Parasite Australia ornamental plants
Aphytis chrysomphali Parasite Adults/Nymphs Fiji Cocos nucifera
Aphytis lingnanensis Parasite Adults/Nymphs India; Karnataka; Kerala Citrus; Dodonaea viscosa
Aphytis melinus Parasite Adults/Nymphs
Azya orbigera Predator Adults/Nymphs Grenada Cocos nucifera
Brumoides suturalis Predator Adults/Nymphs
Chilocorus cacti Predator Adults/Nymphs Dominican Republic Cocos nucifera
Chilocorus circumdatus Predator Adults/Nymphs
Chilocorus dohrni Predator Adults/Eggs/Nymphs
Chilocorus kuwanae Predator Adults/Nymphs
Chilocorus nigrita Predator Adults/Eggs/Nymphs Oman Cocos nucifera
Chilocorus politus Predator Adults/Nymphs Mauritius Cocos nucifera
Chilocorus schioedtei Predator Adults/Eggs/Nymphs
Cladis nitidula Predator Adults/Nymphs St Vincent and the Grenadines Cocos nucifera
Coccidophilus cariba Predator Adults/Nymphs Cayman Islands Cocos nucifera
Comperiella bifasciata Parasite Adults Guam Cocos nucifera
Comperiella unifasciata Parasite Adults Fiji; Mauritius Cocos nucifera
Cryptognatha flaviceps Predator Adults/Nymphs St Lucia Cocos nucifera
Cryptognatha nodiceps Predator Adults/Eggs/Nymphs Angola; Bahamas; Cayman Islands; Dominican Republic; Fiji; Florida; French Polynesia; Grenada; Guam; Hawaii; Jamaica; Pakistan; Puerto Rico; Saipan; Sao Tome and Principe; St Kitts Nevis; St Lucia; St Vincent and the Grenadines; Vanuatu bananas; Cocos nucifera; oil palms; ornamental plants; tobacco
Cryptognatha simillima Predator Adults/Nymphs Puerto Rico; St Lucia Cocos nucifera
Cryptognatha simillima trinitatis Predator Adults/Nymphs Fiji Cocos nucifera
Cryptogonus orbiculus Predator Adults/Nymphs Caroline Islands; Guam Cocos nucifera
Curinus coeruleus Predator Adults/Nymphs St Vincent and the Grenadines Cocos nucifera
Cybocephalus gibbulus Predator Adults/Eggs/Larvae
Egius platycephalus Predator Adults/Nymphs Trinidad Cocos nucifera
Encarsia citrina Parasite Adults/Nymphs Bali; Fiji Cocos nucifera
Encarsia lounsburyi Parasite
Lioscymnus diversipes Predator Adults/Nymphs St Vincent and the Grenadines Cocos nucifera
Lotis neglecta Predator Adults/Nymphs St Vincent and the Grenadines Cocos nucifera
Lotis nigerrima Predator Adults/Nymphs St Vincent and the Grenadines Cocos nucifera
Pentilia castanea Predator Adults/Nymphs Dominican Republic; Puerto Rico Cocos nucifera
Pentilia egena Predator Adults/Nymphs St Lucia Cocos nucifera
Pentilia insidiosa Predator Adults/Nymphs Fiji Cocos nucifera
Pharoscymnus c-luteum Predator Adults/Nymphs Oman Cocos nucifera
Pharoscymnus horni Predator
Pharoscymnus taoi Predator Adults/Nymphs
Pseudoazya trinitatis Predator Adults/Eggs/Nymphs Angola; Bahamas; Fiji; Florida; French Polynesia; Grenada; Jamaica; Pakistan; Puerto Rico; Saipan; St Lucia; Vanuatu Cocos nucifera; ornamental plants
Pseudoscymnus anomalus Predator Adults/Eggs/Nymphs American Samoa; Hawaii; Rota; Saipan; Vanuatu Cocos nucifera
Pseudoscymnus dwipakalpa Predator Adults/Nymphs
Pteroptrix parvipennis Parasite Adults/Nymphs Fiji Cocos nucifera
Rhyzobius lophanthae Predator Adults/Eggs/Larvae/Nymphs/Pupae Guam; Republic of Georgia; St Vincent and the Grenadines; Trinidad Cocos nucifera
Rhyzobius pulchellus Predator Adults/Eggs/Nymphs Fiji; Grenada; Montserrat; St Lucia Cocos nucifera
Rhyzobius satelles Predator Adults/Eggs/Nymphs American Samoa; Caroline Islands; Saipan; Vanuatu; Wallis and Futuna Cocos nucifera
Rodolia rubea Predator Adults/Nymphs
Scymnus cadapani Predator Adults/Eggs/Larvae Philippines
Scymnus gabrieli Predator Adults/Nymphs Philippines
Scymnus luteus Predator Adults/Eggs/Larvae
Scymnus severini Predator Adults/Nymphs
Signiphora borinquensis Parasite
Spaniopterus crucifer Parasite Fiji; Mauritius Cocos nucifera
Sukunahikona prapawan Predator Adults/Nymphs
Telsimia nitida Predator Adults/Eggs/Larvae American Samoa; Ponape; St Vincent and the Grenadines Cocos nucifera
Telsimia sanchezi Predator Adults/Eggs/Larvae Philippines Cocos nucifera
Thomsonisca pakistanensis Parasite
Zagloba aeneipennis Predator Adults/Nymphs Fiji; Puerto Rico Cocos nucifera
Zaomma lambinus Parasite Adults Mauritius Cocos nucifera

Notes on Natural Enemies

Top of page Predators play a significant part in limiting A. destructor populations. The most common are the coccinellid beetles Chilocorus spp., Azya trinitatis, Cryptognatha nodiceps, Rhyzobius lophanthae and Pentilia castanea. Parasites of local significance include Comperiella, Aphytis and Encarsia. A number of parasitoids and predators of A. destructor are described by Rosen (1990).

The parasitoids of A. destructor in India are described by Tandon and Srivastava (1980) and those in Pakistan by Rafiq Ahmad and Ghani (1972). Natural enemies of A. destructor have been described from Sri Lanka (Sinnathamby, 1980), China (Zhou et al., 1993) and Taiwan (Wu and Tao, 1976).

Gordon (1978) describes coccinellid predators from the West Indies. Mariau and Julia (1977) report that predation by coccinellids was usually sufficient to maintain scale populations below the economic level in coconuts in Ivory Coast. The coccinellid Cryptognatha nodiceps is a particularly effective predator (Rosen, 1990).

Natural enemies of other scale insects may adapt to feeding on A. destructor as it colonizes new areas. However, continuous unchecked attacks of A. destructor were recorded on coconuts in Fiji, Mauritius and New Hebrides, until natural enemies were introduced.

Narendra and Rao (1974) describe an entomogenous fungus attacking A. destructor in India. Evans and Prior (1990) list fungal pathogens of A. destructor.

Impact

Top of page A. destructor is potentially the most destructive pest species on coconut, wherever it occurs in the world (Chua and Wood, 1990); the undersurface of the leaves is mainly attacked, but frond stalks, flower clusters and young fruit can also be affected. In extreme cases, the leaves dry up, entire fronds drop off, the crown dies and the whole crop is lost. Neglected coconut plantations are particularly susceptible to damage by A. destructor. A. destructor is also an important economic pest of mango in Asia, Africa, the Philippines, India and Brazil; and of banana in Asia, the Pacific Islands, West Indies, Africa, Madagascar and South America. It attacks the leaves and fruits of oil palms, reducing the quality of the fruits (Chua and Wood, 1990). The species is also a pest of bananas worldwide (Chua and Wood, 1990). However, natural controls appear to keep A. destructor in check in most regions, and few major outbreaks have been recorded in recent years.

Before the introduction of successful biological control in 1955, copra production in Principe fell from 1400 to 500 tons per year owing to an invasion of A. destructor (Rosen, 1990a). After a heavy attack by A. destructor on coconuts in Côte d'Ivoire, yield was reduced by at least 25% over the next 2-3 years, although some heavily infested trees were able to catch up production in the 2 years after elimination of the infestation (Mariau and Julia, 1977).

A. destructor is a cosmetic pest on a wide range of fruits, causing blemishes and other marks that reduce quality. On mango, A. destructor prefers grafted varieties; its economic impact is caused by feeding on tender shoots in nursery plants and because it adversely affects fruit setting in older plants. On oil palm, A. destructor is found feeding on leaves and fruit. It occasionally causes severe damage to guava in India (Hayes, 1970).

This species is highly polyphagous and therefore can easily be re-introduced, even if it is successfully controlled on the primary host crop.

Similarities to Other Species/Conditions

Top of page Aspidiotus excisus looks very similar to A. destructor in life but can be distinguished when slide-mounted adult females are examined microscopically; the median lobes on the pygidium of A. excisus are recessed into the margin, whereas in A. destructor they are not recessed (Williams and Watson, 1988).

Prevention and Control

Top of page

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Introduction

A. destructor is highly polyphagous and therefore can easily be re-introduced, even if it is successfully controlled on the primary host crop.

Host-Plant Resistance

Differences in size and mortality rates were observed on different coconut cultivars by Tabibullah and Gabriel (1973).

Chemical Control

Chemical control may be difficult owing to the height of the trees and may not be commercially viable in some cases owing to the cost. Chemical control may be necessary in the dry season. Malathion has been used successfully. However, insecticide sprays may also kill natural enemies and affect biological control.

Jalaluddin and Mohanasundaram (1991) describe effective chemical treatment for A. destructor in coconut nurseries in India. Zhou et al. (1993) describe the effective chemical control of the larval stages of A. destructor in China. Mariau and Julia (1977) describe the effective chemical control of A. destructor on young coconuts on the Ivory Coast.

Cultural Control

During the early stages of an outbreak of A. destructor on coconut, cutting and burning the affected fronds may be effective.

Phytosanitary Measures

Dharmaraju and Laird (1984) describe the transport of A. destructor around Oceania, mainly through human agency. They emphasize the importance of rigid quarantine procedures.

Biological Control

The most successful biological control of A. destructor has been achieved using predators rather than parasitoids (Rosen, 1990).

An early example of biological control was the introduction of Cryptognatha nodiceps from Trinidad into Fiji in 1928, which stemmed devastating losses in the coconut/copra industry. During 1955, C. nodiceps was introduced into Principe and again almost complete control was obtained and massive economic losses in the copra industry were eliminated (Rosen, 1990). C. nodiceps has since been introduced to a number of oceanic islands; however, it was less successful in the New Hebrides, where Rhyzobius pulchellus, introduced from New Caledonia, was more effective (Rosen, 1990).

Biological control programmes have been described in the Federated States of Micronesia (Suta and Esguerra, 1993), American Samoa (Tauili' ili and Vargo, 1993) and New Hebrides (Chazeau, 1981).

The predatory coccinellid, Chilocorus nigritus, was successfully introduced into Oman from India during 1985 as a biocontrol agent against A. destructor (Kinawy, 1991). Sadakathulla (1993) has developed a technique for the mass production of Chilocorus nigritus.

Rhyzobius lophanthae effectively controlled A. destructor in tea plantations in the Republic of Georgia (Gaprindashvili, 1975).

In Taiwan, the natural enemies of A. destructor were so effective in areas with a rich flora that no chemical control measures were required (Wu and Tao, 1976).

References

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APPPC, 1987. Insect pests of economic significance affecting major crops of the countries in Asia and the Pacific region. Technical Document No. 135. Bangkok, Thailand: Regional Office for Asia and the Pacific region (RAPA)

Beardsley JW, 1966. Insects of Micronesia, Homoptera: Coccoidea. Insects of Micronesia, 6:377-562

Beardsley JW, 1969. Aspidiotus destructor Signoret, an armored scale pest new to the Hawaiian Islands. Proceedings of the Hawaiian Entomological Society, 20(3):505-508

Bennett FD, Alam MM, 1985. An annotated check-list of the insects and allied terrestrial arthropods of Barbados. Bridgetown, Barbados; Caribbean Agricultural Research & Development Institute, vi + 81 pp

Buyckx EJE, 1962. Précis des maladies et des insectes nuisibles recontrés sur les plantes cultivées au Congo, au Rwanda et au Burundi. Brussels, Belgium: Institut National pour l'Etude Agronomique du Congo (INEAC)

Charlin CR, 1973. Coccoidea of Easter Island (Homoptera). Revista Chilena de Entomologia, 7:111-114

Chazeau J, 1981. The biological control of the transparent coconut scale Temnaspidiotus destructor (Signoret) in the New Hebrides (Homoptera, Diaspididae). Cahiers ORSTOM, Serie Biologie, No. 44:11-22

Chua TH, Wood BJ, 1990. Other Tropical Fruit Trees and Shrubs. In: Rosen D, ed. Armoured Scale Insects, their Biology, Natural Enemies and Control. Vol. B. Amsterdam, Netherlands: Elsevier, 543-552

CIE, 1966. Aspidiotus destructor. [Distribution map]. Distribution Maps of Plant Pests, June. Wallingford, UK: CAB International, Map 218

Claps LE, Wolff VRS, González RH, 2001. Catálogo de las Diaspididae (Hemiptera: Coccoidea) exóticas de la Argentina, Brasil y Chile. Revista de la Socieded Entomológica Argentina, 60:9-34

CSIRO, 2001. World Wide Web page at http://www.ento.csiro.au/aicn

Danzig EM, Pellizzari G, 1998. Diaspididae. In: Kozßr F, ed. Catalogue of Palaearctic Coccoidea. Hungarian Academy of Sciences. Budapest, Hungary: Akaprint Nyomdaipari Kft., 172-370

Davidson JA, Miller DR, 1990. Ornamental plants. In: Rosen D, ed. Armoured Scale Insects, their Biology, Natural Enemies and Control. Vol. 4B. Amsterdam, Netherlands: Elsevier, 603-632

Dharmaraju E, 1984. Transport and the spread of crop pests in tropical Polynesia. Commerce and the spread of pests and disease vectors., 257-272; [2 fig.]

Evans HC, Prior C, 1990. Pathogens. In: Rosen D, ed. World Crop Pests. 4B. Armoured Scale Insects: their biology, natural enemies and control. Amsterdam, Netherlands: Elsevier, 3-17

Fernandes IM, 1974. Study of some Coccids from S. Tome. Garcia de Orta, Serie Zoologia, 3(1):1-3

Foldi I, 1988. New contribution to the study of scale insects from Brazilian Amazonia (Homoptera: Coccoidea). Annales de la Societe Entomologique de France, 24(1):77-87

Gaprindashvili NK, 1975. Biological control of the main pests on tea plantations in the Georgian SSR. VIII International Plant Protection Congress, Moscow, 1975. Vol. III. Papers at sessions V... VI... and VII... Moscow. USSR, 29-33

Germain JF, Matile-Ferrero D, 2005. Scale insects from greenhouses in France: an illustrated survey. III - Diaspididae. (Les cochenilles sous serres en France: inventaire illustré. III - Les Diaspididae.) Phytoma, No.583:32-35

Goberdhan LC, 1962. Scale insects of the coconut palm with special reference to Aspidiotus destructor. Journal of the Agricultural Society of Trinidad and Tobago, 62:49-70

Gordon RD, 1978. West Indian Coccinellidae II (Coleoptera): some scale predators with keys to genera and species. Coleopterists Bulletin, 32(3):205-218

Hayes WB, 1970. Fruit growing in India. Kitabistan, Allahabad

Heu RA ed., 2002. Distribution and host records of agricultural pests and other organisms in Hawaii. USA: State of Hawaii Department of Agriculture.

IPPC, 2005. IPP Report No. TV-1/2. Rome, Italy: FAO

Jalaluddin M, Mohanasundaram M, 1989. Control of the coconut scale Aspidiotus destructor Sign. in the nursery. Entomon, 14(3-4):203-206

Jalaluddin SM, Mohanasundaram M, 1992. Detailed study on the morphological features of coconut scale Aspidiotus destructor Sign. Indian Coconut Journal (Cochin), 22(11):10-13; 3 ref

Kawai S, 1980. Scale Insects of Japan in Colours. Tokyo, Japan: National Agriculture Education Association, 455 pp

Kinawy MM, 1991. Biological control of the coconut scale insect (Aspidiotus destructor Sign. Homoptera: Diaspididae) in the southern region of Oman (Dhofar). Tropical Pest Management, 37(4):387-389

Kondo T, 2001. Las cochinillas de Colombia (Hemiptera: Coccoidea). Biota Colombiana, 2(1):31-48

Longo S, Marotta S, Pellizzari G, Russo A, Tranfaglia A, 1995. An annotated list of the scale insects (Homoptera: Coccidea) of Italy. Israel Journal of Entomology, 29:113-130

Mariau D, Julia JF, 1977. New research on the coconut scale Aspidiotus destructor (Sign.). Oleagineux, 32(5):217-224

Miller DR, 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington, 98(1):68-86; 33 ref

Murakami Y, 1970. A review of biology and ecology of diaspine scales in Japan (Homoptera: Coccoidea). MUSHI, 43(7):65-114

Nakahara S, 1982. Checklist of the Armored Scales (Homoptera: Diapididae) of the Conterminous United States. Washington, USA: USDA, Animal and Plant Health Inspection Service, Plant Protection and Quarantine, 110 pp

Narendra DV, Rao VG, 1974. A new entomogenous species of Phoma. Mycopathologia & Mycologia Applicata, 54(1):135-140

Rafiq Ahmad, Ghani MA, 1972. Studies on Aspidiotus destructor Sign.(Hem.:Diaspididae) and its parasites, Aphytis melinus DeBach (Hym.:Aphelinidae) and Pakencyrtus pakistanensis Ahmad. (Hym.: Encyrtidae) in Pakistan. Technical Bulletin of the Commonwealth Institute of Biological Control, No. 15:51-57

Rai BK, 1977. Damage to coconut palms by Azteca sp. (Hymenoptera: Formicidae) and insecticidal control with bait, in Guyana. Bulletin of Entomological Research, 67(1):175-183

Rosen D, 1990. World Crop Pests. 4B. Armoured Scale Insects: their biology, natural enemies and control. Amsterdam, Netherlands: Elsevier Science Publishers, 688 pp

Sadakathulla S, 1993. Technique of mass production of the predatory coccinellid, Chilocorus nigritus (Fabricius) on coconut scale, Aspidiotus destructor Sign. Indian Coconut Journal (Cochin), 23(9):12-13

Shah AH, Jhala RC, Patel CB, 1988. Bioefficacy of aldicarb and BPMC against mango scales and its residues on/in mango fruits. Gujarat Agricultural University Research Journal, 13(2):19-22

Sinnathamby SV, 1980. Developments in the control of coconut scale, Aspidiotus destructor Sign. in Sri Lanka. Ceylon Coconut Quarterly, 28(3-4):81-88

Suta AR, Esguerra NM, 1993. Recent history of biological control in the freely associated states of Micronesia. Micronesica, No. 4 suppl:61-64

Tabibullah M, Gabriel BF, 1973. Biological study of Aspidiotus destructor Signoret in different coconut varieties and other host plants. Philippine Entomologist, 2(6):409-426

Takagi S, 1969. Diaspididae of Taiwan based on material collected in connection with the Japan - U.S. co-operative science programme, 1965 (Homoptera: Coccoidea). Part I. Insecta Matsumurana, Series Entomology, 32:1-110

Tandon PL, Srivastava RP, 1980. New records of parasites and predators of important insect pests of mango. Entomon, 5(3):243-244

Tang SJ, Qin HZ, 1991. Study on Temnaspidiotus destructor (Signoret). Journal of Shanghai Agricultural College, 9(3):190-196

Tao C, 1999. List of Coccoidea (Homoptera) of China. Taichung, Taiwan: Taiwan Agricultural Research Institute, Wufeng, 1-176

Tauili' ili P, Vargo AM, 1993. History of biological control in American Samoa. Biological Control of Exotic Pests in the Pacific. Proceedings of a Plenary Session and Symposium, XIX International Congress of Entomology, Beijing, June 1992. Micronesica, 4 Supplement: 57-60

Taylor THC, 1935. The campaign against Aspidiotus destructor Sign., in Fiji. Bulletin of Entomological Research, 26:1-102

Velasquez FJ, 1971. Some Philippine armored scale insects of the tribe Aspidiotini (Diaspididae, Homoptera). Philippine Entomologist, 2(2):89-154

Waterhouse DF, 1993. The Major Arthropod Pests and Weeds of Agriculture in Southeast Asia. ACIAR Monograph No. 21. Canberra, Australia: Australian Centre for International Agricultural Research, 141 pp

Watson GW, Ooi PAC, Girling DJ, 1995. Insects on plants in the Maldives and their management. Ascot, UK: International Institute of Biological Control

Williams DJ, Watson GW, 1988. The Scale Insects of the Tropical South Pacific Region. Part 1. The Armoured Scales (Diaspididae). Wallingford, UK: CAB International

Williams JR, Greathead DJ, 1990. Sugar Cane. In: Rosen D, ed. Armoured Scale Insects, their Biology, Natural Enemies and Control. Vol. 4B. Amsterdam, Netherlands: Elsevier, 563-578

Williams JR, Williams DJ, 1988. Homoptera of the Mascarene Islands - an annotated catalogue. Entomology Memoir, Department of Agriculture and Water Supply, Republic of South Africa, No. 72, 98 pp

Wong CY, Chen SP, Chou LP, 1999. Guidebook to scale insects of Taiwan. (In Chinese.) Taichung, Taiwan: Taiwan Agricultural Research Institute, 1-98

Wu KC, Tao CCC, 1976. Natural enemies of the transparent scale and control of the leaf bud beetle attacking coconut palm in Taiwan. Journal of Agricultural Research of China, 25(2):141-155

Zhou CA, Zou JJ, Peng JC, 1993. Bionomics of coconut scale - a main pest insect on Actinidia and its control. Entomological Knowledge, 30(1):18-20

Distribution References

APPPC, 1987. Insect pests of economic significance affecting major crops of the countries in Asia and the Pacific region. In: Technical Document No. 135, Bangkok, Thailand: Regional Office for Asia and the Pacific region (RAPA).

Bennett F D, Alam M M, 1985. An annotated check-list of the insects and allied terrestrial arthropods of Barbados. Bridgetown, Barbados: Caribbean Agricultural Research & Development Institute. vi + 81 pp.

Buyckx EJE, 1962. (Précis des maladies et des insectes nuisibles recontrés sur les plantes cultivées au Congo, au Rwanda et au Burundi)., Brussels, Belgium: Institut National pour l'Etude Agronomique du Congo (INEAC).

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Charlin C R, 1973. Coccoidea of Easter Island (Homoptera). (Coccoidea de la Isla de Pascua (Homoptera).). Revista Chilena de Entomologia. 111-114.

Claps LE, Wolff VRS, González RH, 2001. (Catálogo de las Diaspididae (Hemiptera: Coccoidea) exóticas de la Argentina)., 60 9-34.

CSIRO, 2001. World Wide Web., http://www.ento.csiro.au/aicn

Danzig EM, Pellizzari G, 1998. Diaspididae. In: Catalogue of Palaearctic Coccoidea, [ed. by Kozßr F]. Budapest, Hungary: Hungarian Academy of Sciences, Akaprint Nyomdaipari Kft. 172-370.

Fernandes I M, 1974. Study of some Coccids from S. Tome. (Estudo de algumas cochonilhas de S. Tome.). Garcia de Orta, Serie Zoologia. 3 (1), 1-3.

Foldi I, 1988. New contribution to the study of scale insects from Brazilian Amazonia (Homoptera: Coccoidea). (Nouvelle contribution a l'étude des cochenilles de l'Amazonie Brésilienne (Homoptera: Coccoidea).). Annales de la Société Entomologique de France. 24 (1), 77-87.

Germain J F, Matile-Ferrero D, 2005. Scale insects from greenhouses in France: an illustrated survey. III - Diaspididae. (Les cochenilles sous serres en France: inventaire illustré. III - Les Diaspididae.). Phytoma. 32-35.

IPPC, 2005. IPP Report No. TV-1/2., Rome, Italy: FAO.

Kawai S, 1980. Scale Insects of Japan in Colours., Tokyo, Japan: National Agriculture Education Association. 455 pp.

Kinawy M M, 1991. Biological control of the coconut scale insect (Aspidiotus destructor Sign. Homoptera: Diaspididae) in the southern region of Oman (Dhofar). Tropical Pest Management. 37 (4), 387-389.

Kondo T, 2001. The scale insects of Colombia (Hemiptera: Coccoidea). (Las Cochinillas de Colombia (Hemiptera: Coccoidea).). Biota Colombiana. 2 (1), 31-48.

Longo S, Marotta S, Pellizzari G, Russo A, Tranfaglia A, 1995. An annotated list of the scale insects (Homoptera: Coccidea) of Italy. In: Israel Journal of Entomology [Proceedings of the VII International Symposium of Scale Insect Studies, held in Bet Dagan, Israel, June 12-17 1994.], 29 [ed. by Ascher K R S, Ben-Dov Y]. 113-130.

Miller D R, 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington. 98 (1), 68-86.

Nakahara S, 1982. Checklist of the Armored Scales (Homoptera: Diapididae) of the Conterminous United States., Washington, USA: USDA, Animal and Plant Health Inspection Service, Plant Protection and Quarantine. 110 pp.

NHM, 1930. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1940. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1957. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1958. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1967. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1968. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1969. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1985. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

NHM, 1992. Specimen record from the collection in the Natural History Museum (London, UK)., London, UK: Natural History Museum (London).

Suta A R, Esguerra N M, 1993. Recent history of biological control in the freely associated states of Micronesia. In: Micronesica [Biological Control of Exotic Pests in the Pacific. Proceedings of a Plenary Session and Symposium, XIX International Congress of Entomology, Beijing, June 1992.], 61-64.

Takagi S, 1969. Diaspididae of Taiwan based on material collected in connection with the Japan-U.S. Co-operative Science Programme, 1965 (Homoptera: Coccoidea) Part I. Insecta Matsumurana, Entomology. 32 (1), 1-110.

Tao C, 1999. List of Coccoidea (Homoptera) of China., Taichung, Taiwan: Taiwan Agricultural Research Institute, Wufeng. 1-176.

UK, CAB International, 1966. Aspidiotus destructor. [Distribution map]. In: Distribution Maps of Plant Pests, Wallingford, UK: CAB International. Map 218.

Velasquez F J, 1971. Some Philippine armored scale insects of the tribe Aspidiotini (Diaspididae, Homoptera). Philippine Entomologist. 2 (2), 89-154.

Waterhouse D F, 1993. The major arthropod pests and weeds of agriculture in Southeast Asia. Canberra, Australia: ACIAR. v + 141 pp.

Watson G W, Ooi P A C, Girling D J, 1995. Insects on plants in the Maldives and their management. Ascot, UK: International Institute of Biological Control (IIBC). 124 pp.

Wong CY, Chen SP, Chou LP, 1999. Guidebook to scale insects of Taiwan., Taichung, Taiwan: Taiwan Agricultural Research Institute. 1-98.

Zhou C A, Zou J J, Peng J C, 1993. Bionomics of coconut scale - a main pest insect on Actinidia and its control. Entomological Knowledge. 30 (1), 18-20.

Distribution Maps

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