Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Planococcus citri
(citrus mealybug)

Toolbox

Datasheet

Planococcus citri (citrus mealybug)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Vector of Plant Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Planococcus citri
  • Preferred Common Name
  • citrus mealybug
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Planococcus citri is a highly polyphagous, adaptable mealybug that can feed on many host plants in a variety of conditions, and can reproduce rapidly. It has been reported on over 200 host-plant species belongi...

Don't need the entire report?

Generate a print friendly version containing only the sections you need.

Generate report

Pictures

Top of page
PictureTitleCaptionCopyright
Planococcus citri (citrus mealybug); adult females on citrus. Florida, USA.
TitleAdult females
CaptionPlanococcus citri (citrus mealybug); adult females on citrus. Florida, USA.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adult females on citrus. Florida, USA.
Adult femalesPlanococcus citri (citrus mealybug); adult females on citrus. Florida, USA.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); close view of adult female mealybug, on citrus. Florida, USA.
TitleAdult female
CaptionPlanococcus citri (citrus mealybug); close view of adult female mealybug, on citrus. Florida, USA.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); close view of adult female mealybug, on citrus. Florida, USA.
Adult femalePlanococcus citri (citrus mealybug); close view of adult female mealybug, on citrus. Florida, USA.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adult male mealybug, on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. October2007.
TitleAdult male
CaptionPlanococcus citri (citrus mealybug); adult male mealybug, on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. October2007.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adult male mealybug, on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. October2007.
Adult malePlanococcus citri (citrus mealybug); adult male mealybug, on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. October2007.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); eggs are deposited as white cottony masses called ovisacs. Here they are on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. Florida, USA. October2007.
TitleEggs
CaptionPlanococcus citri (citrus mealybug); eggs are deposited as white cottony masses called ovisacs. Here they are on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. Florida, USA. October2007.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); eggs are deposited as white cottony masses called ovisacs. Here they are on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. Florida, USA. October2007.
EggsPlanococcus citri (citrus mealybug); eggs are deposited as white cottony masses called ovisacs. Here they are on Croton (Codiaeum variegatum, Euphorbiaceae). Florida, USA. Florida, USA. October2007.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); nymphs can also be found on fruit. Florida, USA.
TitleNymphs
CaptionPlanococcus citri (citrus mealybug); nymphs can also be found on fruit. Florida, USA.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); nymphs can also be found on fruit. Florida, USA.
NymphsPlanococcus citri (citrus mealybug); nymphs can also be found on fruit. Florida, USA.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adults and nymphs on Schefflera sp. (Araliaceae). Florida, USA. May 2013.
TitleAdults and nymphs
CaptionPlanococcus citri (citrus mealybug); adults and nymphs on Schefflera sp. (Araliaceae). Florida, USA. May 2013.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adults and nymphs on Schefflera sp. (Araliaceae). Florida, USA. May 2013.
Adults and nymphsPlanococcus citri (citrus mealybug); adults and nymphs on Schefflera sp. (Araliaceae). Florida, USA. May 2013.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); females on vine. Florida, USA. March 2008.
TitleFemales
CaptionPlanococcus citri (citrus mealybug); females on vine. Florida, USA. March 2008.
Copyright©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); females on vine. Florida, USA. March 2008.
FemalesPlanococcus citri (citrus mealybug); females on vine. Florida, USA. March 2008.©Lyle J. Buss/Entomology & Nematology Dept./University of Florida - All Rights Reserved
Planococcus citri (citrus mealybug); adults. Laboratory specimens. USA
TitleAdults
CaptionPlanococcus citri (citrus mealybug); adults. Laboratory specimens. USA
Copyright©Jeffrey W. Lotz/Florida Department of Agriculture and Consumer Services/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); adults. Laboratory specimens. USA
AdultsPlanococcus citri (citrus mealybug); adults. Laboratory specimens. USA©Jeffrey W. Lotz/Florida Department of Agriculture and Consumer Services/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); adults, intercepted Georgia Dept. of Agriculture in Coweta County on Croton spp (Euphorbiaceae) plants from Florida, USA.
TitleAdults
CaptionPlanococcus citri (citrus mealybug); adults, intercepted Georgia Dept. of Agriculture in Coweta County on Croton spp (Euphorbiaceae) plants from Florida, USA.
Copyright©Charles Olsen/USDA APHIS PPQ/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); adults, intercepted Georgia Dept. of Agriculture in Coweta County on Croton spp (Euphorbiaceae) plants from Florida, USA.
AdultsPlanococcus citri (citrus mealybug); adults, intercepted Georgia Dept. of Agriculture in Coweta County on Croton spp (Euphorbiaceae) plants from Florida, USA.©Charles Olsen/USDA APHIS PPQ/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp.). Laboratory specimens. USA
TitleInfestation
CaptionPlanococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp.). Laboratory specimens. USA
Copyright©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp.). Laboratory specimens. USA
InfestationPlanococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp.). Laboratory specimens. USA©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); adult on mint (Mentha spp). Intercepted at quarantine from Israel, by CBP Ag Specialist. December 2007.
TitleAdult
CaptionPlanococcus citri (citrus mealybug); adult on mint (Mentha spp). Intercepted at quarantine from Israel, by CBP Ag Specialist. December 2007.
Copyright©Charles Olsen/USDA APHIS PPQ/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); adult on mint (Mentha spp). Intercepted at quarantine from Israel, by CBP Ag Specialist. December 2007.
Adult Planococcus citri (citrus mealybug); adult on mint (Mentha spp). Intercepted at quarantine from Israel, by CBP Ag Specialist. December 2007.©Charles Olsen/USDA APHIS PPQ/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); adult female. Slide mounted specimen. Collected from Fiji in April 1950.
TitleAdult female
CaptionPlanococcus citri (citrus mealybug); adult female. Slide mounted specimen. Collected from Fiji in April 1950.
Copyright©Alessandra Rung/Scale Insects/USDA APHIS ITP/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); adult female. Slide mounted specimen. Collected from Fiji in April 1950.
Adult femalePlanococcus citri (citrus mealybug); adult female. Slide mounted specimen. Collected from Fiji in April 1950.©Alessandra Rung/Scale Insects/USDA APHIS ITP/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); infested and damaged marigold plant (Tagetes spp). USA.
TitleInfestation
CaptionPlanococcus citri (citrus mealybug); infested and damaged marigold plant (Tagetes spp). USA.
Copyright©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); infested and damaged marigold plant (Tagetes spp). USA.
InfestationPlanococcus citri (citrus mealybug); infested and damaged marigold plant (Tagetes spp). USA.©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp). USA.
TitleInfestation
CaptionPlanococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp). USA.
Copyright©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp). USA.
InfestationPlanococcus citri (citrus mealybug); infestation, on marigold (Tagetes spp). USA.©Chazz Hesselein/Alabama Cooperative Extension System/Bugwood.org - CC BY 3.0 US
Planococcus citri (citrus mealybug); natural enemy. The beetle (ladybird, ladybeetle) Cryptolaemus montrouzieri (mealybug destroyer), predating a citrus mealybug.
TitleNatural enemy
CaptionPlanococcus citri (citrus mealybug); natural enemy. The beetle (ladybird, ladybeetle) Cryptolaemus montrouzieri (mealybug destroyer), predating a citrus mealybug.
Copyright©Sonya Broughton/Department of Agriculture & Food Western Australia/Bugwood.org - CC BY-NC 3.0 US
Planococcus citri (citrus mealybug); natural enemy. The beetle (ladybird, ladybeetle) Cryptolaemus montrouzieri (mealybug destroyer), predating a citrus mealybug.
Natural enemyPlanococcus citri (citrus mealybug); natural enemy. The beetle (ladybird, ladybeetle) Cryptolaemus montrouzieri (mealybug destroyer), predating a citrus mealybug.©Sonya Broughton/Department of Agriculture & Food Western Australia/Bugwood.org - CC BY-NC 3.0 US

Identity

Top of page

Preferred Scientific Name

  • Planococcus citri (Risso, 1813)

Preferred Common Name

  • citrus mealybug

Other Scientific Names

  • Coccus citri (Risso)
  • Coccus tuliparum Bouché, 1844
  • Dactylopius brevispinus Targioni Tozzetti, 1881
  • Dactylopius citri (Risso)
  • Dactylopius destructor Comstock, 1881
  • Dactylopius tuliparum (Bouché)
  • Dorthesia citri (Risso, 1813)
  • Dorthezia citri Risso
  • Lecanium phyllococcus Ashmead, 1879
  • Phenacoccus spiriferus Hempel, 1900
  • Planococcoides cubanensis Ezzat and McConnell, 1956
  • Planococcus citricus Ezzat and McConnell, 1956
  • Planococcus cucurbitae Ezzat and McConnell, 1956
  • Pseudococcus citri (Risso)
  • Pseudococcus citri var. phenacocciformis Brain, 1915
  • Pseudococcus citricoleorum Marchal, 1908

International Common Names

  • English: common mealybug
  • Spanish: algodon del naranjo; cochinilla harinosa de los cítricos; cotonet
  • French: cochenille blanche de l'oranger; cochenille blanche des agrumes; cochenille de l'oranger; cochenille du cafeier; cochenille farineuse de la vigne; cochenille farineuse des agrumes

Local Common Names

  • Argentina: cochinilla harinosa de los citrus
  • Brazil: cochonilha branca
  • Chile: chanchito blanco de los citrus
  • Denmark: uldlus, kortfrynset
  • Finland: villakilpikirva
  • Germany: Gewächshausschmierlaus; Weisse Kaffeewurzellaus; Zitrus Schmierlaus
  • Israel: haknima hakimchit she haadar
  • Italy: cocciniglia bianca farinosa della vite; cocciniglia cotonosa degli agrumi; cocciniglia cotonosa della vite
  • Japan: mikan-no-konakaigaramusi
  • Mexico: cotonet del naranja; escama algodonosa; piojo harinoso de los citricos
  • Netherlands: citrus wolluis; witte citrus-luis
  • Norway: ull-lus, kortfrynset
  • Portugal/Madeira: cochonilha-algodao
  • South Africa: sitrus wolluis
  • Sweden: ullus, vanlig
  • Turkey: turuncgil unlu biti

EPPO code

  • PSECCI (Planococcus citri)

Summary of Invasiveness

Top of page

Planococcus citri is a highly polyphagous, adaptable mealybug that can feed on many host plants in a variety of conditions, and can reproduce rapidly. It has been reported on over 200 host-plant species belonging to 191 genera and 82 families, and can seriously damage many crops, particularly citrus and glasshouse tomatoes. It is known to transmit some plant virus diseases like Cacaoa swollen shoot virus. The mealybug is of Old World origin, but its polyphagy has facilitated its spread about the world by human transport of infested plants over many years, and it is now established in in all the temperate and tropical zoogeographic regions, and lives under glass in higher latitudes. Its small size and cryptic habits makes it difficult to detect and identify at plant quarantine inspection. The increase in international trade in fresh plant material in recent years is facilitating its continued spread.

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

Top of page

Planococcus citri was first described as Dorthesia citri by Risso in 1913 from specimens on Citrus in southern France. The species probably originated from China (Bartlett, 1978). It was placed in the genus Pseudococcus Westwood by Brain in 1915 and remained there until it was transferred to the genus Planococcus by Ferris (1950). Synonyms and generic combinations are listed by Ferris (1950), Ezzat and McConnell (1956), De Lotto (1964), McKenzie (1967), Cox (1983, 1989) and García et al. (2016).

The species exhibits a wide range of morphological variation which probably represents a complex of different ecological, biological and geographical races (Ferris, 1950; De Lotto, 1964; Padi, 1990, 1994) or possibly cryptic species (Rung et al., 2009). Cox (1983) demonstrated that P. citri is morphologically highly variable under different environmental conditions and on different host plants. She concluded that specimens identified as Planococcoides cubanensis and Planococcus cucurbitae by Ezzat and McConnell (1956) were aberrant specimens of P. citri. Pseudococcus citri var. coleorum Marchal described from Coleus in 1908 was also synonymized with P. citri by Borchsenius in 1949. Cox (1989) and Padi (1994, 1997a) suggested that earlier records of P. citri on cocoa in West Africa most probably included Planococcus minor (Maskell), which is morphologically very similar.

As on cocoa in West Africa (Entwistle, 1972), an aerial form and a root form of P. citri have been recorded on coffee in East Africa (Gowdey 1917; Hargreaves, 1925), but the root form was probably a misidentification of Planococcus fungicola, described by Watson and Cox (1990) in an association with the fungus Diacanthodes novoguineensis on coffee roots in East Africa.

Recent cytogenetic characterization of the chromosomes of P. citri and the fluorescence karyotype (D287/170), the location of constitutive heterochromatin, rRNA sites and NORs activity show that in the heterochromatic chromosome set, ribosomal genes are still active (Ferraro et al., 1998). This may be useful in separating P. citri from other morphologically similar species of Planococcus such as P. minor, P. ficus, P. kenyae and P. mali. Similarly, work by Bongiorni et al. (1999) on DNA methylation patterns in P. citri may be a useful taxonomic tool. Molecular analysis of the mitochondrial cytochrome oxidase I gene is useful for separating P. citri from P. minor and P. ficus (Rung et al., 2008).

Description

Top of page

Adult Females

The morphology of adult females was described in detail by Ezzat and McConnell (1956), McKenzie (1967), Entwistle (1972), Cox (1989), Cox and Freeston (1985) and Padi (1990).

The external diagnostic characters include 18 pairs of short, stout wax filaments along margins, of which the anal and two preceding pairs are slightly longer than the rest but less than 20% of body length. Dorsum covered with fine mealy wax with a slightly darker, longitudinal, median stripe from first thoracic to mid-abdominal segments. Body colour beneath wax is usually yellow to peach pink. Antenna 8 segmented. Authoritative identification requires microscopic study of slide-mounted females; Sirisena et al. (2013) provided a method for preparation of slide mounts of adult females.

Body of slide-mounted adult female oval, 1.6-3.2 mm long, 1.2-2.0 mm wide (Cox, 1989). Body margin with 18 pairs of cerarii, each cerarius with two conical setae except for the pre-ocular pair which may have one or three setae each. Legs elongate; hind trochanter + femur 220-350 µm long; hind tibia + tarsus 260-420 µm long. Ratio of hind tibia + tarsus to hind trochanter + femur 1.1-1.3; translucent pores present on hind coxae and tibiae. Circulus quadrate, width 120-200 µm. Cisanal setae shorter than anal ring setae. Anal lobes moderately developed; anal lobe cerarii each situated on a small, moderately sclerotized area; venter of each anal lobe with sclerotized anal lobe bar bearing apical seta and bar seta.

Venter
Multilocular disc pores present around vulva, in single or double rows across posterior edges of abdominal segments III-VII, in single rows across anterior edges of segments V-VII, in marginal groups on abdominal segments IV-VII and sometimes a few pores scattered over median area of the thorax and head, but no more than a total of six pores behind the front coxae. Oral collar tubular ducts of two sizes: smaller ducts in sparse rows across median areas of abdominal segments I-VII; larger ducts in marginal groups of variable size around entire venter including head and thorax, and scattered over median area of thorax.

Dorsum
Multilocular disc pores absent. Tubular ducts without rims, slightly larger than the larger ducts on venter, often adjacent to some cerarii. One or two ducts sometimes present on median areas. Simple pores present of two sizes, smaller pores smaller than the smaller size on the venter, scattered over entire dorsum; larger simple pores, each about twice the size of a trilocular pore, present in small groups along mid-line of thoracic and anterior abdominal segments. Setae short and flagellate, longest seta on abdominal segments VI or VII 30-35 µm long.

Adult Males

Males each have a single pair of wings and no mouthparts. They cannot be authoritatively identified at present. Detailed descriptions of the morphology of the adult male are available in Giliomee (1961), Afifi (1968) and Afifi et al. (1976)

Distribution

Top of page

P. citri occurs almost worldwide, except that it appears to be absent from some South Pacific Islands (Cox, 1989). In northern Europe, North America and southern Australia it mainly occurs in greenhouses (CABI/EPPO, 1999).

Due to the difficulty of separating P. citri from P. minor, some of the following details may be referable to P. minor (Maskell). The mealybug is known on cocoa in virtually all parts of the range of this crop but is not always the dominant mealybug species (Entwistle, 1972). In Sri Lanka and Java, it is much less common than P. lilacinus, whilst in Sierra Leone, Cote d'Ivoire and Ghana, its numbers are vastly exceeded by those of P. njalensis. It is co-dominant with P. kenyae and P. njalensis in Nigeria, possibly due to the lighter shade regime (Entwistle, 1972). It is the dominant species in Brazil, Trinidad, Grenada and Hispaniola [Haiti and Dominican Republic]. In Australia it is reported to be widely established, both in the field and in greenhouses; however, some of these records could be P. minor, as was verified by Williams (1985) for some Queensland collections from passionfruits by Murray (1976; 1978a, b). Le Pelley (1968) reported that P. citri occurs in nearly all coffee-growing countries of the world.

The distribution map includes records based on specimens in the Natural History Museum, London, UK, listed as (NHM, various dates).

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaPresentBen-Dov, 1994
AzerbaijanPresentCABI/EPPO, 1999
BangladeshPresentUllah and Parveen, 1993; CABI/EPPO, 1999
British Indian Ocean TerritoryPresentCABI/EPPO, 1999
Brunei DarussalamPresentWaterhouse, 1993; CABI/EPPO, 1999
CambodiaPresentCABI/EPPO, 1999
ChinaPresentTang et al., 1992; CABI/EPPO, 1999
-AnhuiRestricted distributionWeng and Huang, 1988; CABI/EPPO, 1999
-FujianWidespreadCABI/EPPO, 1999
-GuangdongWidespreadCABI/EPPO, 1999
-GuangxiWidespreadCABI/EPPO, 1999
-GuizhouWidespreadCABI/EPPO, 1999
-HainanRestricted distributionCABI/EPPO, 1999
-Hong KongPresentCABI/EPPO, 1999
-HubeiWidespreadCABI/EPPO, 1999
-HunanWidespreadCABI/EPPO, 1999
-JiangsuWidespreadCABI/EPPO, 1999
-JiangxiRestricted distributionCABI/EPPO, 1999
-SichuanWidespreadCABI/EPPO, 1999
-YunnanWidespreadCABI/EPPO, 1999
-ZhejiangWidespreadCABI/EPPO, 1999
Georgia (Republic of)PresentCABI/EPPO, 1999
IndiaWidespreadKrishnamoorthy and Mani, 1994; Mani, 1994; CABI/EPPO, 1999
-GujaratPresentWilliams, 2004
-KarnatakaPresentReddy and Bhat, 1993a; Krishnamoorthy, 1990; CABI/EPPO, 1999
-KeralaPresentPrakasan, 1987; CABI/EPPO, 1999
-MaharashtraPresentCABI/EPPO, 1999
-MeghalayaPresentRao et al., 2001
-SikkimPresentCABI/EPPO, 1999
-Tamil NaduPresentCABI/EPPO, 1999
-TripuraPresentCABI/EPPO, 1999
-Uttar PradeshPresentCABI/EPPO, 1999
-West BengalPresentCABI/EPPO, 1999
IndonesiaPresentWaterhouse, 1993; CABI/EPPO, 1999
-Irian JayaAbsent, unreliable recordCABI/EPPO, 1999
-JavaPresentCABI/EPPO, 1999
-SulawesiPresentLe Pelley, 1968; CABI/EPPO, 1999
-SumatraPresentKenten and Woods, 1976; CABI/EPPO, 1999
IranPresentCABI/EPPO, 1999
IraqPresentCABI/EPPO, 1999
IsraelPresentBlumberg et al., 1995; CABI/EPPO, 1999
JapanPresentYukawa, 1978; CABI/EPPO, 1999
-Bonin IslandPresentKawai, 1987
-HonshuPresentCABI/EPPO, 1999
-KyushuPresentCABI/EPPO, 1999
-Ryukyu ArchipelagoPresentCABI/EPPO, 1999
JordanPresentCABI/EPPO, 1999
Korea, Republic ofPresentPaik, 1972; CABI/EPPO, 1999
LaosPresentCABI/EPPO, 1999
LebanonPresentCABI/EPPO, 1999
MalaysiaPresentWaterhouse, 1993; CABI/EPPO, 1999
-Peninsular MalaysiaPresentCABI/EPPO, 1999
-SabahPresentLim et al., 1992; CABI/EPPO, 1999
-SarawakPresentCABI/EPPO, 1999
MaldivesPresentNHM, 1976; CABI/EPPO, 1999
MyanmarPresentWaterhouse, 1993; CABI/EPPO, 1999
OmanPresentKinawy et al., 2008
PakistanPresentCABI/EPPO, 1999
PhilippinesPresentWaterhouse, 1993; CABI/EPPO, 1999; Williams, 2004
Saudi ArabiaPresentCABI/EPPO, 1999
Sri LankaPresentCABI/EPPO, 1999
SyriaPresentCABI/EPPO, 1999
TaiwanWidespreadTsay, 1991; Su and Li, 1993
TajikistanPresentCABI/EPPO, 1999
ThailandPresentWaterhouse, 1993; CABI/EPPO, 1999
TurkeyPresentSoylu and Urel, 1977; Uygun et al., 1987; CABI/EPPO, 1999
TurkmenistanPresentBen-Dov, 1994; CABI/EPPO, 1999
United Arab EmiratesAbsent, unreliable recordAbd El Rahim et al., 1977; CABI/EPPO, 1999
UzbekistanPresentCABI/EPPO, 1999
VietnamWidespreadLe Pelley, 1968; Waterhouse, 1993; CABI/EPPO, 1999
YemenPresentCABI/EPPO, 1999

Africa

AldabraPresentCABI/EPPO, 1999
AlgeriaPresentCABI/EPPO, 1999
AngolaPresentCABI/EPPO, 1999
BeninPresentCABI/EPPO, 1999
Burkina FasoPresentCABI/EPPO, 1999
BurundiPresentCABI/EPPO, 1999
CameroonPresentCABI/EPPO, 1999
Cape VerdePresentNHM, 1986; CABI/EPPO, 1999
ChadPresentCABI/EPPO, 1999
ComorosPresentCABI/EPPO, 1999
CongoPresentMatile-Ferrero, 1978; CABI/EPPO, 1999
Congo Democratic RepublicPresentNHM, 1929; CABI/EPPO, 1999
Côte d'IvoireWidespreadEntwistle, 1972; CABI/EPPO, 1999
EgyptPresentGaaboub et al., 1979; CABI/EPPO, 1999
EritreaPresentCABI/EPPO, 1999
EthiopiaPresentCABI/EPPO, 1999
GabonPresentCABI/EPPO, 1999
GhanaPresentStrickland, 1951a; Strickland, 1951b; Padi, 1990; CABI/EPPO, 1999
GuineaPresentCABI/EPPO, 1999
KenyaWidespreadDe Lotto, 1964; Baum, 1970; CABI/EPPO, 1999
LiberiaPresentCABI/EPPO, 1999
LibyaPresentBitaw and Saad, 1990; CABI/EPPO, 1999
MadagascarPresentNHM, 1989; CABI/EPPO, 1999
MalawiPresentDe Lotto, 1964; CABI/EPPO, 1999
MaliPresentCABI/EPPO, 1999
MauritiusPresentCABI/EPPO, 1999
MoroccoPresentCABI/EPPO, 1999
MozambiquePresentLe Pelley, 1968; CABI/EPPO, 1999
NigerPresentCABI/EPPO, 1999
NigeriaWidespreadOnazi, 1969; Entwistle, 1972; CABI/EPPO, 1999
RéunionPresentCABI/EPPO, 1999
Rodriguez IslandPresentCABI/EPPO, 1999; CABI/EPPO, 1999
RwandaPresentCABI/EPPO, 1999
Saint HelenaPresentCABI/EPPO, 1999
Sao Tome and PrincipePresentCABI/EPPO, 1999
SenegalPresentCABI/EPPO, 1999
SeychellesPresentCABI/EPPO, 1999
Sierra LeonePresentCABI/EPPO, 1999
South AfricaWidespreadDe Lotto, 1975; Georgala et al., 1975; CABI/EPPO, 1999
Spain
-Canary IslandsPresentTsalev, 1970; CABI/EPPO, 1999
SudanPresentCABI/EPPO, 1999
SwazilandPresentNHM, 1987; CABI/EPPO, 1999
TanzaniaPresentDe Lotto, 1964; CABI/EPPO, 1999
-ZanzibarPresentWilliams and Matile-Ferrero, 2005
TogoPresentMancion and Aliber, 1936; Dufour, 1991; CABI/EPPO, 1999
TunisiaPresentCABI/EPPO, 1999; Halima-Kamel et al., 2014
UgandaPresentDe Lotto, 1964; CABI/EPPO, 1999
ZambiaPresentNHM, 1967; CABI/EPPO, 1999
ZimbabwePresentPyle, 1979; CABI/EPPO, 1999

North America

BermudaPresentCABI/EPPO, 1999
MexicoPresentRico-Grey and Thien, 1989; CABI/EPPO, 1999; Arriola et al., 2016
USARestricted distributionCABI/EPPO, 1999
-AlabamaPresentCABI/EPPO, 1999
-ArizonaPresentCABI/EPPO, 1999
-ArkansasPresentCABI/EPPO, 1999
-CaliforniaPresentHickman, 1981; CABI/EPPO, 1999
-FloridaPresentPena and Duncan, 1990; CABI/EPPO, 1999
-HawaiiPresentHansen et al., 1992a; Hansen et al., 1992b; Zimmerman, 1948; Hata and Hara, 1992; CABI/EPPO, 1999
-IowaPresentBen-Dov, 1994
-KansasPresentCABI/EPPO, 1999
-LouisianaPresentCABI/EPPO, 1999
-MarylandPresentCABI/EPPO, 1999
-MassachusettsPresentCABI/EPPO, 1999
-MissouriPresentCABI/EPPO, 1999
-New MexicoPresentCABI/EPPO, 1999
-OhioPresentLindquist, 1979; Lindquist, 1981; CABI/EPPO, 1999
-South CarolinaPresentCABI/EPPO, 1999
-TennesseePresentCABI/EPPO, 1999
-TexasPresentMeyerdirk et al., 1978; CABI/EPPO, 1999
-VirginiaPresentCABI/EPPO, 1999

Central America and Caribbean

Antigua and BarbudaPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
BahamasPresentCABI/EPPO, 1999
BarbadosPresentCABI/EPPO, 1999
British Virgin IslandsPresentCABI/EPPO, 1999
Cayman IslandsPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
Costa RicaPresentCABI/EPPO, 1999
CubaPresentMestre et al., 1991; Angeles Martínez et al., 1992; CABI/EPPO, 1999
DominicaPresentCABI/EPPO, 1999
Dominican RepublicPresentCABI/EPPO, 1999
El SalvadorPresentCABI/EPPO, 1999
GrenadaPresentCABI/EPPO, 1999
GuadeloupePresentAPPPC, 1987
GuatemalaPresentLe Pelley, 1968; CABI/EPPO, 1999
HaitiAbsent, unreliable recordCABI/EPPO, 1999
HondurasPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
JamaicaPresentCABI/EPPO, 1999
MartiniquePresentMatile-Ferrero and Étienne, 2006
MontserratPresentCABI/EPPO, 1999
Puerto RicoPresentLe Pelley, 1968; CABI/EPPO, 1999
SabaPresentLim et al., 1992
Saint LuciaPresentCABI/EPPO, 1999
Saint Vincent and the GrenadinesPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
Trinidad and TobagoPresentMcComie, 1987; CABI/EPPO, 1999
United States Virgin IslandsPresentBeatty, 1944Saint Croix

South America

ArgentinaPresentBen-Dov, 1994; CABI/EPPO, 1999; Granara and de Willink Claps, 2003
BrazilPresentCABI/EPPO, 1999
-BahiaPresentCABI/EPPO, 1999
-Espirito SantoPresentCABI/EPPO, 1999; Culik et al., 2009
-Minas GeraisPresentCABI/EPPO, 1999
-ParanaPresentFoldi and Kozár, 2006
-Rio de JaneiroPresentCABI/EPPO, 1999
-Rio Grande do SulPresentCABI/EPPO, 1999
-Santa CatarinaPresentCABI/EPPO, 1999
-Sao PauloPresentCABI/EPPO, 1999
ChilePresentCABI/EPPO, 1999; González, 2011
ColombiaPresentCABI/EPPO, 1999
EcuadorPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
-Galapagos IslandsPresentWilliams and Granara de Willink, 1992; CABI/EPPO, 1999
GuyanaPresentCABI/EPPO, 1999
ParaguayPresentCABI/EPPO, 1999
PeruPresentLe Pelley, 1968; CABI/EPPO, 1999
SurinamePresentCABI/EPPO, 1999
UruguayPresentCABI/EPPO, 1999
VenezuelaPresentCABI/EPPO, 1999

Europe

AustriaPresentCABI/EPPO, 1999
BelgiumPresentRonse, 1990; CABI/EPPO, 1999
BulgariaPresentTsalev, 1970; CABI/EPPO, 1999
CroatiaPresentMilek et al., 2008
CyprusPresentKrambias and Kontzonis, 1980; CABI/EPPO, 1999
Czech RepublicPresentBen-Dov, 1994
Czechoslovakia (former)PresentCABI/EPPO, 1999
Former USSR-in-EuropePresentKhalilov, 1972; Kurdyukov and Alan, 1973
FrancePresentPanis, 1981; CABI/EPPO, 1999
-CorsicaPresentCABI/EPPO, 1999
GermanyPresentSengonca et al., 1995
GreecePresentCabitza et al., 1994; CABI/EPPO, 1999
-CretePresentCABI/EPPO, 1999; CABI/EPPO, 1999
HungaryPresentDarvas and Szabo, 1987; CABI/EPPO, 1999
ItalyPresentMineo, 1992; Barbagallo et al., 1993; CABI/EPPO, 1999
-SardiniaPresentCABI/EPPO, 1999; CABI/EPPO, 1999
-SicilyPresentCABI/EPPO, 1999
MaltaWidespreadCABI/EPPO, 1999
NetherlandsPresentVos et al., 1993; Kozár et al., 1996; CABI/EPPO, 1999
PolandPresentCABI/EPPO, 1999
PortugalPresentFernandes, 1990; Pedroso et al., 1991; CABI/EPPO, 1999
-AzoresPresentCABI/EPPO, 1999
-MadeiraPresentCABI/EPPO, 1999
RomaniaAbsent, unreliable recordCABI/EPPO, 1999
Russian FederationPresentNiyazov, 1969; Bazarov, 1988; CABI/EPPO, 1999
-Southern RussiaPresentCABI/EPPO, 1999
San MarinoPresentWilliams and Granara de Willink, 1992
SloveniaPresent
SpainPresentRipolles and Garcia, 1989; CABI/EPPO, 1999
-Balearic IslandsPresentCABI/EPPO, 1999
SwitzerlandPresentKozár et al., 1994
UKPresentCABI/EPPO, 1999; Malumphy and Badmin, 2012Under glass
-England and WalesPresentCABI/EPPO, 1999; Malumphy and Badmin, 2012Under glass
-ScotlandPresentCABI/EPPO, 1999Under glass
UkrainePresentBen-Dov, 1994; CABI/EPPO, 1999

Oceania

AustraliaPresentCABI/EPPO, 1999
-Australian Northern TerritoryPresentWilliams, 1985; CABI/EPPO, 1999
-New South WalesPresentWilliams, 1985; CABI/EPPO, 1999Also in Australian Capital Territory
-QueenslandPresentWilliams, 1985; Smith et al., 1988; CABI/EPPO, 1999
-South AustraliaPresentWilliams, 1985; CABI/EPPO, 1999
-TasmaniaPresentWilliams, 1985; CABI/EPPO, 1999
-VictoriaPresentCABI/EPPO, 1999
-Western AustraliaPresentWilliams, 1985; CABI/EPPO, 1999
Caroline IslandsPresentBeardsley jr, 1966
Cook IslandsWidespreadWilliams and Watson, 1988; CABI/EPPO, 1999
FijiAbsent, unreliable recordWilliams and Watson, 1988; CABI/EPPO, 1999
French PolynesiaPresentCABI/EPPO, 1999
GuamPresentCABI/EPPO, 1999
KiribatiAbsent, unreliable recordCABI/EPPO, 1999
Marshall IslandsPresentCABI/EPPO, 1999
Micronesia, Federated states ofPresentBeardsley jr, 1966; CABI/EPPO, 1999Chuuk, Kosrae, Pohnpei, Yap
New CaledoniaAbsent, unreliable recordCABI/EPPO, 1999
New ZealandPresentNHM, 1979; CABI/EPPO, 1999
NiuePresentWilliams and Watson, 1988; CABI/EPPO, 1999
Northern Mariana IslandsAbsent, unreliable recordCABI/EPPO, 1999
PalauPresentCABI/EPPO, 1999
Papua New GuineaPresentAPPPC, 1987; Williams and Watson, 1988; CABI/EPPO, 1999
SamoaPresentWilliams and Watson, 1988; CABI/EPPO, 1999
Solomon IslandsAbsent, unreliable recordGollifer et al., 1977; CABI/EPPO, 1999
TongaPresentWilliams and Watson, 1988; CABI/EPPO, 1999
Wallis and Futuna IslandsAbsent, unreliable recordCABI/EPPO, 1999

History of Introduction and Spread

Top of page

P.  citri is of Old World origin (Cox, 1989), but was accidentally spread by human trade in fresh plants from an early date, before such things were recorded. It is least widely established in the Austro-Oriental, South Pacific, Malagasian and northern Neotropical Regions. Due to the difficulty of separating P. citri from several very similar species, some geographical records may be based on misidentifications. 

Risk of Introduction

Top of page

P. citri is a vector of Cacao swollen shoot virus: to prevent virus transmission, it is necessary to effect quarantine measures on cocoa material, particularly seedlings which may harbour virus-infected mealybugs. The mealybug has been introduced to many countries on ornamental plants such as Coleus and on food plants such as potatoes.

P. citri was able to survive on flowers of Alpinia purpurata even after they were subjected to hot air treatment followed by hot water immersion. Mealybug survival increased by 0.4% after 2 h in hot air plus hot water treatment, and by 40% after 4 h hot air plus hot water treatment (Hara et al., 1997).

Habitat List

Top of page
CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Principal habitat Harmful (pest or invasive)
Cultivated / agricultural land Principal habitat Natural
Protected agriculture (e.g. glasshouse production) Principal habitat Harmful (pest or invasive)
Protected agriculture (e.g. glasshouse production) Principal habitat Natural
Managed forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Managed forests, plantations and orchards Principal habitat Natural
Managed grasslands (grazing systems) Secondary/tolerated habitat Natural
Disturbed areas Secondary/tolerated habitat Natural
Rail / roadsides Secondary/tolerated habitat Natural
Urban / peri-urban areas Secondary/tolerated habitat Harmful (pest or invasive)
Urban / peri-urban areas Secondary/tolerated habitat Natural
Buildings Secondary/tolerated habitat Harmful (pest or invasive)
Buildings Secondary/tolerated habitat Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Secondary/tolerated habitat Natural
Natural grasslands Secondary/tolerated habitat Natural
Riverbanks Secondary/tolerated habitat Natural
Wetlands Secondary/tolerated habitat Natural
Scrub / shrublands Secondary/tolerated habitat Natural

Hosts/Species Affected

Top of page

P. citri is polyphagous and occurs on a wide range of flowering plants, having been recorded on over 200 host species belonging to 191 genera in 82 families (García et al., 2016).

In the tropics, it occurs mainly on the aerial parts of crops such as cocoa, bananas, tobacco and coffee and on wild trees such as Ceiba pentandra and Leucaena (Strickland, 1951a,b; Le Pelley, 1968; Entwistle, 1972). In Ghana, P. citri has been recorded on about 54 host plants, including cocoa, cola, pineapples, Musa paradisiaca and others within the families Bombacaceae, Euphorbiaceae, Fabaceae, Moraceae, Rubiaceae, Solanaceae, Sterculiaceae, Tiliaceae and Urticaceae (Strickland, 1951a; Padi et al., 1999). Hargreaves (1937) cited Anisophyllea laurina as an alternative food plant in Sierra Leone.


In the South Pacific region, P. citri has been recorded on 20 host plants, including Brassica, Ceiba, Citrus, cocoa, Cyrtosperma, Cucurbita, Gardenia, Inocarpus, Ipomoea, Leucaena, Morinda, Ocimum, Psidium, Pueraria and Solanum spp. (Williams and Watson, 1988). Its host plants in Australia include pumpkins in New South Wales; Clerodendrum, Coleus, Croton and Erythrina species in hothouses, and Ceratonia, Siliqua and Veronica species in the open in Adelaide (Brooks, 1957); and pineapples (Carter, 1942), Vitis vinifera and passionfruits in Queensland (Williams, 1973; Murray, 1978b).

In India, P. citri occurs on mandarin orange (Amitava Konar, 1998) and has been recorded for the first time on soyabean (Jadhav et al., 1996).

In temperate regions, P. citri mainly occurs on greenhouse plants such as Coleus, ferns and gardenias, but also occurs outdoors under summer conditions on Citrus, grapes, figs, taro, date palms and potatoes (Bivins and Deal, 1973; Gibson and Turner, 1977). It mainly attacks Citrus but not grapes in the Mediterranean region (Cox, 1989) and California. In the former Soviet Union, it occurs on over 20 species of plants, notably Citrus, figs and pomegranates (Niyazov, 1969). In Turkmenistan, pomegranates are most liable to heavy infestation. It occurs on Areca sp. and a wide range of greenhouse ornamental plants in Korea (Paik, 1972) and Bulgaria (Tsalev, 1970).

In Texas, USA, P. citri has been recorded on the milk vine Cynanchum unifarium [Cynanchum racemosum var. unifarium] (French and Reeve, 1978). Host plants in India include Macadamia ternifolia (Wysoki, 1977).

Host Plants and Other Plants Affected

Top of page
Plant nameFamilyContext
Alpinia purpurata (red ginger)ZingiberaceaeUnknown
Ananas comosus (pineapple)BromeliaceaeOther
Anisophyllea laurinaAnisophylleaceaeWild host
AnnonaAnnonaceaeOther
Annona muricata (soursop)AnnonaceaeOther
Annona squamosa (sugar apple)AnnonaceaeOther
Arachis hypogaea (groundnut)FabaceaeOther
ArecaArecaceaeOther
BrassicaBrassicaceaeOther
Cajanus cajan (pigeon pea)FabaceaeOther
Carica papaya (pawpaw)CaricaceaeOther
Ceiba pentandra (kapok)BombacaceaeOther
CeratoniaFabaceaeOther
CitrusRutaceaeMain
Clerodendrum (Fragrant clerodendron)LamiaceaeOther
Cocos nucifera (coconut)ArecaceaeOther
Codiaeum variegatum (croton)EuphorbiaceaeOther
Coffea (coffee)RubiaceaeOther
ColaSterculiaceaeOther
ColeusLamiaceaeOther
Crossandra undulifoliaAcanthaceaeOther
CrotonEuphorbiaceaeOther
Cucurbita (pumpkin)CucurbitaceaeOther
Cynanchum racemosum var. unifarium (talayote)AsclepiadaceaeWild host
Cynodon dactylon (Bermuda grass)PoaceaeOther
Cyrtosperma merkusii (giant swamp taro)AraceaeOther
Dioscorea (yam)DioscoreaceaeOther
Diospyros kaki (persimmon)EbenaceaeOther
ErythrinaFabaceaeOther
EugeniaMyrtaceaeOther
FicusMoraceaeOther
GardeniaRubiaceaeOther
Glycine max (soyabean)FabaceaeOther
Gossypium hirsutum (Bourbon cotton)MalvaceaeOther
InocarpusFabaceaeOther
Ipomoea batatas (sweet potato)ConvolvulaceaeOther
Laurus nobilis (sweet bay)LauraceaeOther
LeucaenaFabaceaeOther
MacadamiaProteaceaeOther
Macadamia ternifolia (Queensland nut)ProteaceaeOther
Malus domestica (apple)RosaceaeOther
Mangifera indica (mango)AnacardiaceaeOther
Manihot esculenta (cassava)EuphorbiaceaeOther
MorindaMain
Musa (banana)MusaceaeOther
Musa x paradisiaca (plantain)MusaceaeOther
Nephelium lappaceum (rambutan)SapindaceaeOther
Nicotiana tabacum (tobacco)SolanaceaeOther
Ocimum basilicum (basil)LamiaceaeOther
Olea (olive)OleaceaeOther
Passiflora (passionflower)PassifloraceaeOther
Passiflora edulis (passionfruit)PassifloraceaeOther
Persea americana (avocado)LauraceaeOther
Phaseolus lunatus (lima bean)FabaceaeOther
Phoenix dactylifera (date-palm)ArecaceaeOther
Psidium guajava (guava)MyrtaceaeOther
PuerariaFabaceaeOther
Punica granatum (pomegranate)PunicaceaeOther
Pyrus communis (European pear)RosaceaeOther
Saccharum officinarum (sugarcane)PoaceaeOther
SiliquaOther
Solanum (nightshade)SolanaceaeOther
Solanum lycopersicum (tomato)SolanaceaeOther
Solanum tuberosum (potato)SolanaceaeOther
Theobroma cacao (cocoa)MalvaceaeOther
Vanilla africanaOrchidaceaeOther
Veronica (Speedwell)ScrophulariaceaeOther
Vitis vinifera (grapevine)VitaceaeOther
Xanthium strumarium (common cocklebur)AsteraceaeOther
Ziziphus mauritiana (jujube)RhamnaceaeMain

Growth Stages

Top of page Flowering stage, Fruiting stage, Post-harvest, Seedling stage, Vegetative growing stage

Symptoms

Top of page

P. citri feeding leads to general wilting due to sap depletion. On cocoa, flower stalks, buds and young pods are attacked (Entwistle, 1972). In Taiwan, infested immature coffee berries become deformed and drop to the ground (Moriyama, 1941).

P. citri infestation also causes indirect physical damage because sugary honeydew excreted by the mealybugs fouls plant surfaces, giving rise to sooty moulds (Gausman and Hart, 1974) that block light and air from the leaves, inhibiting photosynthesis.

Citrus mealybug is the second most important vector of several strains of Cacao swollen shoot virus;  symptoms include leaf chlorosis, root necrosis, root and stem swellings and dieback (Posnette, 1941; Cotterell, 1943).

List of Symptoms/Signs

Top of page
SignLife StagesType
Fruit / abnormal shape
Fruit / external feeding
Fruit / honeydew or sooty mould
Fruit / premature drop
Growing point / external feeding
Growing point / honeydew or sooty mould
Inflorescence / external feeding
Inflorescence / wilt
Leaves / abnormal colours
Leaves / honeydew or sooty mould
Leaves / wilting
Roots / external feeding
Roots / stubby roots
Stems / external feeding
Stems / honeydew or sooty mould

Biology and Ecology

Top of page

Life Cycle

The life history of P. citri has been described by James (1932, 1937), Myres (1932), Betrem (1936), Risbec (1937), Entwistle (1958) and Tang et al. (1992). The species reproduces sexually and is oviparous; egg production starts about 9-14 days after fertilization. The adult female lays amber-yellow eggs in a fluffy posterior ovisac that is about equal in length to the body. The number of eggs varies from 150-200 (Côte d'Ivoire) and 20-250 (Ghana) to about 300 (on cocoa in Trinidad) and up to 500 (on Citrus in California). The incubation period varies from 2 to 10 days (Le Pelley, 1968).


Females pass through three nymphal instars before reaching adulthood. The nymphs differ from the adult females by having a much thinner wax covering and fewer antennal segments, and are much more mobile. Male nymphs are similar to the females for the first two instars but at the third instar they pass through non-feeding prepupal and then a quiescent pupal stage before becoming adults. Betrem (1936) found that males were rare and believed parthenogenesis might occur. Other workers (James, 1937; Entwistle, 1972; Padi, 1997b) have found that males and females are produced in approximately equal numbers: the specific sex ratio established from 11413 progeny from 43 females was 101.6 ±1.5 males per 100 females (James, 1937) and 82 males to 106 females (Myres, 1932).

Joint duration of the female nymphal instars are variously quoted as 16 days in Trinidad and 32-38 days in Côte d'Ivoire and Ghana (Entwistle, 1972). The adult female continues to grow and lays her first eggs when 9-10 days old. The period from hatching to egg laying is approximately 5 weeks (Entwistle, 1972). The adult male emerges 3 days after pupation and the period from egg laying to emergence of the adult male is 16-17 days in Trinidad. Females tend to be more active in the 12-hour period following each moult and, in adults, shortly before oviposition (Kirkpatrick, 1953).


Experiments indicated that the duration of the different developmental stages of P. citri reared on potato tubers varied considerably according to food plant quality and the temperature. Arai (1996) concluded that the higher developmental threshold temperature must be around 30°C. Lower developmental threshold temperatures and thermal constants for P. citri were 7.7°C and 401 DD during the nymphal stage and 8.0°C and 378 DD during the preovipositional period.

Males are attracted to females by a sex pheromone, dextro-cis-planococcyl acetate, produced by the female (Gravitz and Wilson, 1968; Rotundo and Tremblay, 1974, 1976b, 1980a; Benassy et al., 1976; Panis, 1979). Monitoring experiments using the synthetic pheromone showed that the male P. citri is an early morning flier with a flight range in the field of up to 183 m (assisted by wind) compared to a maximum of 2 cm direct flight to the virgin female (Moreno et al., 1984).

In California, USA, there are four to five overlapping generations per year (Bartlett, 1978), but Gray (1954) found almost double this number. In southern California, overwintering is predominantly in the egg stage, but some of all stages are usually present (Myres, 1932; McKenzie, 1967). In Ghana, there appear to be six to ten generations per year on cocoa and field studies have shown that there is no period in the year when colonies cannot be found, although they are more common in the dry than in the wet season (Entwistle, 1972). In Peru, P. citri overwinters in the nymphal or adult stage and has four generations in the year (Khalilov, 1972). In Morocco on Citrus cv. Maroc Late, Abdelkhalek et al. (1998) found that P. citri had six to eight overlapping generations annually and high population levels occurred during June-December.

In Italy, Onorato et al. (1998) used pheromone trap catches to develop a method to determine the relationship between temperature and infestations of P. citri on citrus fruits.

Population Dynamics

In Cuba, populations were most abundant from November with a peak in February. Angeles Martínez et al. (1991) showed that the insects move on a plant in response to soil temperature below 25°C and air humidity below 85% RH, when they migrated preferentially to the roots, but moved to the aerial parts when these parameters increased.

Spatial distribution studies on grapevines in Israel showed that P. citri was more abundant during October and that its within-tree distribution was the result of intrinsic behaviour (eg. thigmotaxis and negative phototaxis) during crawler dispersal (Nestel et al., 1995) and that there was no effect of the host on phenology distribution. However, this record might be referable to P. ficus.

On cocoa, Campbell (1975) studied changes in canopy populations of P. citri in relation to leaf flushing, whilst Bigger (1975a) conducted studies on the effects of parasitoids on annual population changes and their inter-relations with other insects.

In Italy, Viggiani (1975b) observed that P. citri infestation on citrus fruits rose from 0.5% in mid-July to 29.5% in November. In Spain, Limon de la Oliva et al. (1972) also conducted studies on seasonal development of P. citri. Le Pelley (1968) related the seasonal trends in P. citri populations with seasonal changes in degree of protein metabolism whilst Betrem (1936) considered humidity to be the most important ecological factor influencing P. citri through its influence on fungi parasitic on the insect.

In Java, P. citri increases markedly in long dry spells (Bernard, 1926), particularly in the dry monsoon (Betrem, 1936). Infestation in the field begins to increase when the average relative humidity at noon falls below 70%, which is normally at the beginning of the dry season, and reaches its peak 3 to 4 months later. Quillis Perez (1935) in Spain also found that its development is retarded at high humidity.

Associations

P. citri is a vector of Cacoa swollen shoot virus and has been implicated with the transmission of Grapevine leafroll-associated virus 3 (GLRaV-3) (Cabaleiro and Segura, 1997). It has also been reported, for the first time, as a vector of Banana streak virus and Cucumber mosaic virus infecting banana cultivars (Musa spp.) in Taiwan (Su HongJi et al., 1997).

P. citri was identified as a vector of the Dioscorea bacilliform virus (DaBV) on Dioscorea alata from Barbados and West Africa, and on other Dioscorea spp. from West African, Caribbean, Asian and South American countries (Phillips et al., 1999). The mealybug also transmits a previously undescribed badnavirus, Schefflera ringspot virus (SRV) from infected schefflera and aralia to healthy schefflera seedlings, but not to Fatsia japonica or Hedera helix. The virus occurs on Brassaia actinophylla [Schefflera actinophylla], Schefflera arboricola and a number of Araliaceae, among others, in Australia, Barbados, Cuba, Mauritius, Honduras, Taiwan, Thailand and the USA (Lockhart et al., 1996).

ELISA studies on the acquisition and transmission of Grapevine leafroll-associated virus 3 (GLRaV-3) showed that P. citri could retain the virus for up to 24 hours but lost the capacity for effective transmission to vines within 1 hour after transfer. In newly infected vines, the virus remained latent or undetectable by ELISA for at least 13 months.

Silva and Mexia (1999) observed a significant positive correlation between Cryptoblabes gnidiella and P. citri on sweet orange (Citrus sinensis) groves in the Algarve, Portugal, supporting the hypothesis of several authors that an infestation of P. citri is necessary for C. gnidiella attack in citrus. Even low levels of infestation by C. gnidiella can cause serious damage by fruit drop and, consequently, greatly reduce the production of sweet orange, mainly in cv. Navel.

Strickland (1951a,b) recorded 20 species of ants attending P. citri on cocoa in Ghana. Among these were ten species of Crematogaster, including C.luctans and C.kneri, nine species of Pheidole and one Camponotus species. Nevertheless, he observed that P. citri was less strictly associated with ants than was Planococcoides njalensis, the commonest mealybug species on cocoa in Ghana.

In Brazil, P. citri is attended by the ant Wasmannia auropunctata (Delabie, 1988; Delabie and Cazorla, 1991). In Trinidad, there were more P. citri attended by O. auropunctatus on shaded than on unshaded plots, and more on trees which received nitrogenous fertilizer than those which did not (Fennah, 1959). Infestation levels were also negatively correlated with the depth of gravel below the soil surface. It was also established that P. citri development was not prevented by lack of carbohydrate even under deep shade but that nitrogenous components of food are of great importance. In addition to providing a good feeding site, the sepals of grapefruits are believed to provide good protection for P. citri against its parasites (Berlinger et al., 1978).

Campbell (1994) observed that P. citri and P. njalensis on cocoa in Ghana were attended mainly by Pheidole megacephala, although P. citri is generally less frequently attended by ants (Entwistle, 1972). Studies on two cocoa progenies, T85/799 X T17/359 and E1:C43/291 x T63/967 (Series 11B hybrid) by Bigger (1975b, c) in Ghana indicated that for a given canopy size, P. citri was 2.3 times more prevalent in trees of the Series 11 hybrid. It was also concluded that the apparent preference for the Series 11 hybrid was more likely to be due to a physiological difference between the two progenies than to purely environmental differences. Firempong (1984), also in Ghana, tested seven cocoa progenies both in the laboratory and in the field. He concluded that two progenies, 85D/176 x M7/537 and T12/116 x T62/977 were the most resistant to mealybug [Planococcoides njalensis and P. citri] attack.

In Java, the habit of P. citri is influenced by conditions at different altitudes: above 2000 feet the insect's primary food plant is the shade tree Leucaena glauca [L. leucocephala], but below this coffee is the chief host (De Fluiter, 1937). Below 300 feet of altitude it is probably too warm for infestations to occur on coffee.

Climate

Top of page
ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

Top of page
Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
36 36

Air Temperature

Top of page
Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) 7.7

Natural enemies

Top of page
Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aenasius abengouroui Parasite Adults/Nymphs
Allotropa citri Parasite Bermuda; Chile; USA Croton; fruit trees; polyphagous
Allotropa kamburovi Parasite
Allotropa mecrida Parasite
Anagrus greeni Parasite
Anagyrus amoenus Parasite
Anagyrus beneficiens Parasite Adults/Nymphs
Anagyrus bohemani Parasite
Anagyrus pseudococci Parasite Nymphs Chile; USA; Texas Citrus; fruit trees
Anagyrus sawadai Parasite
Anagyrus sp. nr. kivuensis Parasite California pears
Arhopoideus pretiosus Parasite Italy Citrus
Aspergillus flavipes Pathogen
Aspergillus flavus Pathogen Adults/Nymphs
Blepyrus saccharicola Parasite California
Brumoides lineatus Predator Adults/Nymphs
Brumoides suturalis Predator Adults/Nymphs
Cephalosporium Pathogen
Chilocorus renipustulatus Predator Adults/Nymphs
Chrysoperla carnea Predator Adults/Nymphs
Chrysoperla zastrowi Predator Adults/Nymphs
Chrysoplatycerus splendens Parasite USA; Texas Citrus
Cladosporium oxysporum Pathogen Adults/Nymphs
Clausenia josefi Parasite
Coccidoxenoides peregrinus Parasite Adults/Nymphs Bermuda; California; Chile; Procida; USA; Texas Citrus; fruit trees; polyphagous
Coccinella transversalis Predator Adults/Nymphs
Coccodiplosis coffeae Predator Adults/Nymphs
Coccodiplosis smithi Predator Adults/Nymphs
Coccophagus gurneyi Parasite Adults/Nymphs Italy Citrus
Cryptolaemus affinis Parasite
Cryptolaemus montrouzieri Predator Adults/Nymphs Australia; Queensland; Bahamas; California; Chile; Cyprus; France; Greece; Greece; Crete; Hawaii; Israel; Italy; Italy; Sardinia; Karnataka; Montserrat; Peru; Saudi Arabia; Sicily; South Africa; Spain; St Helena; Taiwan Citrus; fruit trees; grapefruits; orchards; polyphagous
Cybocephalus micans Predator Adults/Nymphs
Diadiplosis hirticornis Predator Adults/Nymphs
Dicrodiplosis manihoti Predator
Diomus pumilio Predator Adults/Nymphs
Diomus rubidus Predator Adults/Nymphs Israel Citrus
Domomyza perspicax Predator Adults/Nymphs
Entomophthora Pathogen
Euseius stipulatus Predator Adults/Nymphs
Exochomus flaviventris Predator Adults/Nymphs
Exochomus metallicus Predator Adults/Nymphs California
Holcencyrtus myrmicoides Parasite California
Hyperaspis egregia Predator
Hyperaspis raynevali Predator Adults/Nymphs
Leptomastidea abnormis Parasite Nymphs Australia; Queensland; Belgium; Bermuda; California; Chile; Costa Rica; Hawaii; Israel; South Africa; USA; Texas; USSR Citrus; Ficus carica; fruit trees; polyphagous
Leptomastidea sp. nr. abnormis Parasite California
Leptomastix bifasciatus Parasite Adults/Nymphs
Leptomastix dactylopii Parasite Adults/Nymphs Australia; Australia; Queensland; Belgium; California; Chile; Cyprus; France; India; India; Karnataka; Israel; Italy; Italy; Sardinia; Sicily; South Africa; Spain; Texas; USA; Texas; USSR Annona; Citrus; Ficus carica; fruit trees; oranges; orchards; polyphagous
Leptomastix nigrocoxalis Parasite
Leptomastix trilongifasciatus Parasite
Leucopis alticeps Predator
Mallada basalis Predator Adults/Nymphs
Mallada boninensis Predator Adults/Nymphs
Neozygites fresenii Pathogen Adults/Nymphs
Nephus bipunctatus Predator Adults/Nymphs California
Nephus georgei Predator
Nephus reunioni
Oligochrysa lutea Predator Adults/Nymphs
Phytoseiulus persimilis Predator Adults/Nymphs China; Zhejiang Citrus
Prochiloneurus pulchellus Parasite
Pseudaphycus maculipennis Parasite
Pseudaphycus perdignus Parasite Bermuda; California; Chile Citrus; fruit trees; polyphagous
Pseudoscymnus pallidicollis Predator Adults/Nymphs
Scymnus apiciflavus Predator Adults/Nymphs
Scymnus coccivora Predator Adults/Nymphs
Scymnus includens Predator Adults/Nymphs
Scymnus roepkei Predator Adults/Nymphs
Spalgis epeus
Sympherobius amicus Predator Adults/Nymphs USSR Citrus
Sympherobius barberi Predator Adults/Nymphs
Timberlakia gilva Parasite
Triommata coccidivora Predator Adults/Nymphs
Tropidophryne melvillei Parasite Adults/Nymphs
Typhlodromus phialatus Predator Adults/Nymphs

Notes on Natural Enemies

Top of page

The natural enemies recorded for P. citri are numerous (Donald, 1956; Le Pelley, 1968; Niyazov, 1976; Barbier and Raimbault, 1985; Uygun et al., 1987; Mani and Krishnamoorthy, 1990c; Dufour, 1991; Reyd et al., 1991; Islam and Jahan, 1993; Su & Li, 1993; Vos et al., 1993; Blumberg et al., 1995; García et al., 2016; Noyes, 2016). On coffee, Le Pelley (1968) lists seven primary parasitoids from Indonesia, Malaysia, South America and south India. He also reports eight coccinnelid predators from Java, Australia, India, New Guinea and Uganda. Natural enemies identified on cocoa in Trinidad, Brazil and West Africa are few (Donald, 1956; Ackonor et al., 1993; Ackonor, 1997, 2000). In Ghana, Bigger (1976) recorded a mean level of parasitism of 25.6% (range, 8.3-59.4%), taking into account nymphs that dropped to the ground to pupate, and concluded that the level previously recorded by Donald (1956) (mean of 3.4%; range, 0.5-15.4%) was an underestimation. Ackonor (1997, 2000) and Ackonor et al. (1993) recorded 14 hymenopterous parasitoid species including five species of Encyrtidae, Aenasius abengouri, Leptomastix dactylopii (introduced in 1949), Anagyrus beneficians, Tropidophryne melvellei and Anagyrus amoenus and three predatory species, Coccodiplosis coffeae, Hyperaspis egregia and Scymnus (Pullus) sp. Parasitism levels recorded for the three most abundant parasitoid species were Anagyrus beneficians (3.1±3.1%), Aenasius abengouroui (4.6±2.6) and Leptomastix dactylopii (4.0±2.4%). Infestation by the predatory species were 16.0±5.8% for C. coffeae, 6.9±4.3% for Scymnus (Pullus) sp. and 6.3±4.8% for H. egragia.

In Trinidad, the only primary parasitoid is the successful Leptomastix dactylopii, which is, however, heavily restrained by a complex of parasitoids (Kirkpatrick, 1953). The most frequent predator in Trinidad was a Scymnus species. In the Markham Valley in Papua New Guinea the coccinelid Cryptolaemus affinis is the main predator (Entwistle, 1972). Nephus bisignatus [N. georgei] has been recorded for the first time as a predator of P. citri on Thuja orientalis and Pistacia lentiscus in Greece (Kontodimas, 1997).

Information on the biology and establishment of several parasitoids incuding Anagyrus pseudococci, Anagyrus sawadai, Anagyrus sp., Coccidoxenoides peregrinus, Leptomastidea abnormis and Leptomastix dactylopii in Australia, India and elsewhere is provided by van Baaren and Nénon (1996a, b), Krishnamoorthy and Mani (1996), Smith et al. (1996), Cadée et al. (1997), Islam et al. (1997), Islam and Copland (1997), Mani and Krishnamoorthy (1997), Rivero-Lynch and Godfray (1997), Su TsongHong et al. (1997), Yang and Sadof (1997) and Ceballo et al. (1998). Information on the predators Cryptolaemus montrouzieri, Dicrodiplosis manihoti and Spalgis epeus is provided by Rahiman and Vijayalakshmi (1998), Abbas (1999), Baskaran et al. (1999), Garcia and O'Neil (2000) and Voigt (2000).

Oviposition responses of Cryptolaemus montrouzieri on P. citri in the laboratory showed that the females delayed oviposition and withheld mature eggs in their lateral oviducts in the absence of wax filaments produced by the prey (Merlin et al., 1996). Thus, contact chemical cues perceived by females when probing the wax filaments with their mouthparts were the signals inducing the search for oviposition sites. The second step was under the control of the ovipositor by which females located confined sites to lay eggs. This behaviour could have a considerable impact on the prey exploitation strategy of this important biocontrol agent and might help to explain its apparent ineffectiveness in situations of low prey density.

Very few pathogenic fungi have been recorded attacking P. citri: Entomophthora fresenii [Neozygites fresenii] in Java (De Fluiter, 1939), Aspergillus flavipes in Cuba, Cephalosporium sp. (Rojter et al., 1966) and Entomophthora sp. from ricefields in India (Samal et al., 1978). Attempts to use A. flavipes for the control of P. citri in Cuba, however, showed that it could not be recommended because of its mycotoxins which are harmful to man and animals (Martínez and Bravo, 1989).

Means of Movement and Dispersal

Top of page

Natural Dispersal

Local dispersal of P. citri is mainly by the crawling of the first instars through thigmokinesis and negative photokinesis (Nestel et al., 1995); but all the female developmental stages can walk, tending to be more active shortly before moulting and just before oviposition (Kirkpatrick, 1953); they will move to avoid unfavourable conditions (Angeles Martínez et al., 1991). Additionally, wind may pick up and carry crawlers longer distances (Gerson, 2016).

Vector Transmission

Human transport of infested plants is a common way in which P. citri gets transported over long distances, in-country or internationally. Humans and farm machinery working in infested fields can accidentally carry crawlers on them to other sites (Gerson, 2016). Plant fanciers often share or trade material, which can help to introduce or disperse mealybugs. In unfavourable conditions, attendant ants sometimes pick up mealybugs and carry them to new feeding sites (Way and Khoo, 1991; Anon., 2016). Sometimes mealybug crawlers walk onto the feet of birds perching on infested trees, and get carried to new plants.

Accidental Introduction

Import of infested plants is a common way for mealybugs to be carried between countries accidentally, such as trade in nursery stock, and import of unusual plants to botanical gardens and food plant material to zoos. There is also considerable trade in planting material like ornamental bamboos and orchids, bought online and sent via mail or courier, sometimes with no customs declaration on the package.

Pathway Causes

Top of page
CauseNotesLong DistanceLocalReferences
Botanical gardens and zoosAccidental introduction on imported plants Yes
Crop productionAccidental introduction on imported plants, transport on used farm machinery Yes Yes Gerson, 2016
Cut flower tradeAccidental introduction on imported plant material Yes Yes
Escape from confinement or garden escape Yes
HorticultureAccidental introduction on imported plants, transport on workers, tools and machinery Yes Yes Gerson, 2016
Landscape improvementAccidental introduction on imported plants Yes Yes Gerson, 2016
Nursery tradeAccidental introduction on imported plants, transport on workers, tools and machinery Yes Yes Gerson, 2016

Pathway Vectors

Top of page
VectorNotesLong DistanceLocalReferences
ConsumablesAccidental introduction on imported plant material Yes
MailIn mailed plant material Yes
Plants or parts of plants Yes Yes
Wind Yes Gerson, 2016

Plant Trade

Top of page
Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Flowers/Inflorescences/Cones/Calyx adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Fruits (inc. pods) adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants nymphs Yes Yes Pest or symptoms usually visible to the naked eye
Leaves adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Roots adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Stems (above ground)/Shoots/Trunks/Branches adults; eggs; larvae; nymphs; pupae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Seedlings/Micropropagated plants
True seeds (inc. grain)
Wood

Impact Summary

Top of page
CategoryImpact
Economic/livelihood Negative

Economic Impact

Top of page

Citrus, Fruit and Vegetables


P. citri is reported as a serious pest, causing damage to various crops such as Citrus, grapes and mangoes, although crop loss is usually difficult to assess. It is the most injurious of the mealybugs on Citrus in the Mediterranean region (Panis, 1977). In South Africa, it is one of the six most important pests of deciduous fruits (Myburgh et al., 1973). P. citri was reported to cause more than 75% damage to bunches of grapes (variety Black Champa) at Hessaraghatta, India (Mani and Thontadarya, 1987). Damage to leaves, shoots and main stems of grapes lead eventually to plant death in Trans-Caucasus, North Caucasus, Krasnodar region, Crimea, Central Asia and Dagestan in the former USSR (Abdullagatov, 1978). The appearance of cork scars was directly related to the level of P. citri infestation in fruits of Citrus variety 'Sweetie' (Citrus paradisi x C. maxima) in orchards in Israel (Gross et al., 1999). The majority of injuries became evident during the first and second generations of P. citri. Only medium to high population densities of P. citri caused a significant reduction in production, both in terms of fruit weight and fruit size, in orange groves in Portugal (Silva et al., 1997); low numbers of P. citri caused fruit discoloration, fruit splitting and chlorotic spots.

In Australia, Citrus sp., Solanum tuberosum, Nerium oleander, Croton sp., Coleus sp. and Dracaena sp. in New South Wales were severely infested (Williams, 1985). Brimblecombe (1962) also reported it on Citrus from several counties in the coastal areas of Queensland as far north as Townsville where it was an economic pest, and on Annona squamosa, but more particularly on bunches of grapes where there were heavy infestations. Williams (1973) recorded it on Macadamia integrifolia from Berwaa where it attacked fruit racemes and young terminal growths. A field survey in Nigeria showed that infestation of stored potato tubers (Solanum tuberosum) originated from the field and that affected seed potatoes stored over considerable periods shrivelled and died (Onazi, 1969). In the USA, P. citri is a serious pest of greenhouse crops (Bivins and Deal, 1973). P. citri also transmits bacilliform particles of a virus disease on edible aroids, including Colocasia esculenta and Xanthosoma sagittifolium in the Solomon Islands and the South Pacific (Carpenter et al., 1976; Gollifer et al., 1977). 

Most of the early Pacific reports of P. citri causing severe outbreaks (Barrett, 1966; Szent-Ivany and Stevens, 1966) should refer to Planococcus minor, with which it was (and sometimes still is) confused (Williams, 1982; Williams and Watson, 1988).

Coffee

In the tropics, P. citri is more usually a pest of coffee than of citrus (De Lotto, 1964). In Java it is an important pest of coffee, often occurring with Ferrisiavirgata, and together they "give a lot of trouble" (Ultee, 1924). In Taiwan, it infests berries of coffee and the immature berries become deformed or drop to the ground (Moriyama, 1941). It is sometimes a severe pest in Guatemala (Barrie, 1958) and in Puerto Rico and other islands, it is an "appreciable" pest (Wolcott, 1951). Although severe damage to the roots of Coffea arabica and C. liberica has been reported in Guatemala, Puerto Rico, Vietnam (DuPasquier, 1930) and in India (Anstead, 1917), confusion exists over species identity.

Cocoa

Numerous reports of physical damage to cocoa caused by P. citri all refer to the West Indies. In Grenada, for instance, some trees form abnormally large cushions, bearing high numbers of flowers but producing few pods (Ballou, 1921). Damage caused to the tips of lateral growths on cocoa in Papua New Guinea is thought to allow entry of secondary fungi (Szent-Ivany, 1961). However, Cox and Freeston (1985) showed that records of Planococcus on cocoa in the Oriental and Neotropical Regions were P. minor and not P. citri, although most published records are under the latter name.

In West Africa, P. citri is a vector of many cocoa viruses and is the second most important vector of the Cacao swollen shoot virus disease in Ghana (Strickland, 1947, 1951a, b; Donald, 1955; Sutherland, 1953; Entwistle, 1972; Bigger, 1981; Dufour, 1991). In North Sumatra, Indonesia, it is a vector of a disease which causes leaf symptoms ranging from clearing of the major veins to a more complex mosaic pattern that is similar to symptoms of Cacao swollen shoot virus (Kenten and Woods, 1976).

Yam

In Nigeria, P. citri, together with the scale insect Aspidiella hartii and unspecified beetles, has been associated with fungal infections (Fusarium spp., Penicillium spp., Aspergillus spp., Curvularia spp., Epicoccum spp. and Helminthosporium spp.) on stored yam tubers (Morse et al., 2000).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Tolerant of shade
  • Capable of securing and ingesting a wide range of food
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Host damage
  • Negatively impacts agriculture
  • Negatively impacts livelihoods
  • Reduced amenity values
  • Negatively impacts animal/plant collections
  • Damages animal/plant products
  • Negatively impacts trade/international relations
Impact mechanisms
  • Competition - monopolizing resources
  • Pest and disease transmission
  • Fouling
  • Herbivory/grazing/browsing
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally illegally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

Detection and Inspection

Top of page

Although generally cryptic in nature, P. citri can be easily detected on fruits and inflorescences. On cocoa, it can be readily detected on the surface of pods, where it usually forms large colonies. Colonies in terminal buds, bases of leaf petioles, points of attachment of chupons [suckers], fruits and pods, and the bark of trees can be detected using a hand lens or, in the case of terminal buds, by teasing them apart and inspecting them under a dissecting microscope. On Citrus, the area underneath the calyx and the peduncle of the fruit provides a good hiding place (Meyerdirk et al., 1981). P. citri can also be detected in the field by the presence of ants and sooty moulds that develop on excreted honeydew (Gausman, 1974) and by wilt of plant parts such as leaves, inflorescences and fruits or berries.

Similarities to Other Species/Conditions

Top of page

P. citri is morphologically very similar to P. ficus, P. halli, P. kenyae, P. lilacinus, P. mali, and P. minor: all these species were initially either confused with, or misidentified as P. citri. The different species can now confidently be distinguished from each other by both external (Le Pelley, 1943; Entwistle, 1972) and microscopic characters (Cox, 1989). Thus, the mealybug found on vines throughout the Mediterranean basin is now believed to be P. ficus and not P. citri (Cox, 1989).

Planococcus kenyae, the common coffee mealybug in East Africa, differs from P. citri by having fewer ventral multilocular disc pores that usually occur in only one row in posterior edges of abdominal segments V-VIII and that do not extend to the margin of the segments. Moreover, P. kenyae lays naked eggs, not enclosed in an ovisac as occurs with P. citri. Planococcus lilacinus also differs from P. citri in not producing an ovisac, and in having body contents in young adults pale violet to maroon, becoming brownish-red when fully mature. In P. citri and P. kenyae the body contents are yellowish.
 

Separation of P. citri from P. ficus, P. halli and P. mali using morphology is difficult even when slide-mounted adult females are studied at high magnification; Cox (1989) provided a key. Morphological separation of P. citri from P. minor is exceptionally difficult, even using the discriminant function base don scores of six characters, provided by Cox (1989); according to Cox, the cut-off score is 35, with specimens scoring higher being P. citri and lower being P. minor. However, some populations of P. citri can have lower scores so it is necessary to score multiple specimens, and a determination of P. minor only be made if the scores are around 20 or lower.


Molecular sequencing of the mitochondrial cytochrome oxidase gene provides a separation of P. citri from P. minor (Rung et al., 2008; 2009).

Males of P. citri and P. ficus can also be separated: P. citri has 1-2 median pronotal pores whilst P. ficus has 3-6 (Afifi et al., 1976).

Prevention and Control

Top of page

Introduction

In several countries, P. citri has been successfully controlled with chemicals and natural enemies or a combination of the two. Attempts have also been made to control the pest using semiochemicals, cultural methods and resistant plant material.

Cultural Control and Sanitary Measures

Keeping fruit trees pruned so that they do not touch each other may help slow spread of P. citri, and cleaning farm equipment and other objects immediately after use in the field can help prevent its transport between trees and orchards. 

In the highlands of Java, Indonesia, the shade tree Leucaena glauca is the main food plant of P. citri at altitudes above 600 m. Measures that proved successful for the control of P. citri were mainly directed against infestation of this tree and consisted of removing the flower clusters or, when necessary, pruning all foliage and flowers. It is also claimed that P. citri can be controlled by increasing the shade in plantations and that this was undesirable for Robusta coffee but suitable for Arabica at high altitudes (Le Pelley, 1968). Good results were obtained by providing three covers, one above the other, of Leucaena, Erythrina and Albizia, or with Leucaena and Albizia only. It is further suggested that because the insect infests mainly the flowers and pods of L. glauca, other shade trees that seldom flower, such as L. pulverulenta [L. leucocephala], or a sterile hybrid of L. glauca and L. glabrosa should be planted (De Fluiter, 1939).

In Texas, USA, P. citri infestation on Citrus was controlled indirectly by controlling the milk vine Cynanchum unifarium [Cynanchum racemosum var. unifarium(milk vine), a climbing plant that grows within the citrus tree canopy (French and Reeve, 1978).

Biological Control

P. citri is often attended by ants for the sugary honeydew it excretes. Ants from multiple genera are involved, including species of Crematogaster, Pheidole, Camponotus and Wasmanniaauropunctata, with different species active in different countries and conditions, on various crops (Way and Khoo, 1991). Ants defend the mealybugs from predators and parasitoids, and can render biological control agents ineffective. Ant control is therefore an essential component in the biological control of P. citri.

P. citri has natural enemies in 22 genera belonging to 7 families (García et al., 2016). Several natural enemies, both parasitoids and predators, have been studied and evaluated for the biological control of P. citri (Viggiani, 1974; Berlinger et al., 1979, 1985; Barbier and Raimbault, 1985; Mani and Krishnamoorthy, 1990a,b,c; Reddy et al., 1991, 1992, 1997, Reyd et al., 1991; Islam and Jahan, 1992, 1993; Kanika-Kiamfu et al., 1993; Khandakar and Jahan, 1993; Su & Li, 1993, Vos et al., 1993; van Baaren et al., 1994; Yigit et al., 1994; Blumberg et al., 1995; Mani, 1995). The food plants of the pest may affect the activity of natural enemies (Copland et al., 1993). Natural enemies that have been successfully used include the parasitoids Leptomastix dactylopii, of South American origin, and Anagyrus pseudococci; and the predators Cryptolaemus montrouzieri and Exochomus flavipes (Panis and Brun, 1971; Panis, 1977; Narasimham, 1987; Pillai, 1987; Smith et al., 1988; Viggiani, 1988; Nagarkatti et al., 1992; Reddy et al., 1992, Barbagallo et al., 1993; Reddy and Bhat, 1993a; Mani, 1994).

P. citri was successfully controlled with the introduced predator C. montrouzieri and the parasitoid L. dactylopii in Italy (Panis, 1977), Sicily (Liotta et al., 1976), France (Panis and Brun, 1971), India (Pillai, 1987; Reddy et al., 1992; Reddy and Bhat, 1993b; Mani, 1994), former USSR (Kurdyukov and Alan, 1973), Queensland in Australia (Smith et al., 1988), Italy (Fronteddu et al., 1996) and Morocco (Abdelkhalek et al., 1998).

In Peru, the encyrtid parasitoid Pauridia peregrina [Coccidoexnoides peregrinus], which was introduced accidentally from Texas (USA) and first discovered in Peru in 1963, gave remarkably good control of P. citri on Citrus (Salazar, 1972).

On Citrus in the former Soviet Union (Niyazov, 1969), P. citri was kept under control by the indigenous parasitoid Anagyrus pseudococci, which destroyed up to 75% of the mealybug population in areas that were not treated with insecticides. In Turkmenistan and Georgia, however, the effects of Leptomastix abnormis [Leptomastidea abnormis] and Leptomastix dactylopii introduced from the USA in 1960 were much reduced by the presence of hyperparasitoids such as Chartocerus subaeneus, which was also responsible for 18-20% parasitism of another parasitoid, Allotropa mecrida, which in turn was responsible for up to 20% parasitism of P. citri. The larvae of predatory Leucopis alticeps and Crysopa carnea [Chrysoperla carnea] are reported as virtually destroying all stages of P. citri in the former USSR, and introduction of Clausenia josefi from the Mediterranean was discussed (Niyazov, 1969).

In Athens (Greece), introduced C. montrouzieri failed to establish on Citrus, whereas in Antibes (France) it was found to be effective at temperatures above 20°C but proved ineffective at lower temperatures or in the presence of attendant ants (Panis and Brun, 1971). In Israel, laboratory and field experiments showed that the efficacy of predators was reduced when P. citri fed on the alkaloid-containing legumes Erythrina corallodendrum and Spartium junceum, compared to non-toxic plants (Mendel et al., 1992).

Studies conducted in Italy on hypersensitivity and allergic responses in a group of workers employed in breeding insects for biological pest control showed a lesser degree of allergy to P. citri than to other insects (Cipolla et al., 1997).

Laboratory experiments conducted at Purdue University, West Lafayette, USA, on Coleus, showed that plant size, variegation and other parameters including plant height, leaf number and leaf area had no significant effect on attack rates, searching strategy and selected life history characteristics of C. montrouzieri (Garcia and O'Neil, 2000). On the other hand, increasing prey density on plants significantly increased the rate of attack, which reached a plateau of approximately 3.5-4.0 prey attacked on plants containing 16 prey individuals. There was an inverse relationship between prey density and the estimated area searched. Moreover, C. montrouzieri development was slower, and survival and fourth-instar and adult body weight was lower when provided with fewer prey. However, the sex ratio was unaffected.

Laboratory experiments on the reproduction of C. montrouzieri showed that whilst the ladybirds did not reproduce or lay eggs after consuming Orthezia tillandsiae, a small amount of a feed mixture consisting of the ensign scales O. tillandsiae and P. citri at a ratio of 3:1 was enough to stimulate the ladybirds to lay eggs. A further experiment with the same feed-mixture showed that C. montrouzieri pupated and continued to develop (Voigt, 2000). In India the biology and predation of C. montrouzieri on P. citri and Dactylopius tomentosus was studied in the laboratory at 29.4-32.1°C and 65-71% RH (Baskaran et al., 1999). C. montrouzieri completed its growth successfully with both prey species, but preferred P. citri to D. tomentosus.

See also Waterhouse (1998).

Chemical Control

Few insecticides are known to be very effective against mealybugs; their waxy coating protects them from contact insecticides, their cryptic habits make it difficult to reach them with sprays, and the overlapping generations prevent complete control. Care is necessary when using pesticides, to ensure that the natural enemies are not adversely affected. Spot treatment or integrated pest control are more cost-effective than full pesticidal control.

On coffee in Kenya, P. citri is controlled indirectly by spraying chemicals or applying baits against attendant ants, to encourage natural enemies that would otherwise be killed by them (Le Pelley, 1968). Chlorpyrifos, deltamethrin, ethion and hydramethylnon were used to control ants on coffee in Kenya.

Phosalone, trichlormetafos-3, malathion and dimethoate were effective against P. citri in Peru and the former USSR (Khalilov, 1972; Kurdyukov and Alan, 1973). The most suitable periods for insecticide application on grapevines in the former USSR were during the transition phase and during the emergence of third-instar larvae; spraying could be replaced by the release of the predator Cryptolaemus montrouzieri. ULV application of malathion (Limon de la Oliva and Blasco Pascual, 1973) has also been used.

On grapefruits and oranges in Spain (province of Castellou), spot treatments with fenitrothion, fenitrothion + dimethoate or dimethoate were effective but expensive (Limon de la Oliva et al., 1972).

In Italy, 40 insecticides effective against P. citri were assessed for their effect on the natural enemies, Leptomastidea abnormis and Scymnus includens (Viggiani et al., 1972). Twenty-one of these, including a preparation of Bacillus thuringiensis, a mixed spray of chlorfensulphide [superseded], chlorfenethol [superseded], demeton [superseded], dicofol, liquid mineral oil and pyrethrum, were found to be least harmful to natural enemies.

Other chemicals used elsewhere include fenthion in Italy (Cabitza et al., 1994), dimethoate in Australia (Murray, 1978a) and, indirectly, chlorpyrifos, deltamethrin, ethion, hydramethylnon inacide and used to control attendant ants on coffee in Kenya to encourage survival of parasitoids and predators that would be killed by the ants (Anon., 1990). On persimmon fruits, the application of chlorpyrifos and diazinon around the trunk gave excellent results against mealybug attendant ants and greatly reduced the population of mealybugs (Izhar, 1999).

In Citrus orchards in Israel, the application of chlorpyrifos in early April (before flowering) prevented cork scars on fruits, whereas application in early summer (mid-June) only reduced damage (Gross et al., 1999).

In Maryland, USA, micronized dusts of chlorpyritos dispersed in a greenhouse by release of carbon dioxide proved highly effective against nymphs of P. citri (Smith and Boswell, 1972). In Tokat province (Italy), diazinon gave effective control in two applications made 5 weeks apart in June and July (Aykac and Erguder, 1972). These three insecticides were also found to be effective in Turkey. In Bulgaria, phenthoate was most effective, but was phytotoxic to ferns. Dimethoate also afforded some control (Tsalev, 1970).

In a study of the influence of six selective pesticides on adult longevity, progeny production and prey consumption by the predator Cryptolaemus montrouzieri, no detrimental effects were observed on progeny production of the beetle on plants treated with chlorpyrifos, neem seed kernel extract, dicofol and copper oxychloride. Diflubenzuron had a pronounced effect on the adult females, yielding only 278 progeny compared to 419 by untreated females (Mani et al., 1997). It was suggested that these selective pesticides, with the exception of diflubenzuron, could be considered in integrated pest management programmes.

Pirimiphos-methyl effectively controlled P. citri, scales and beetles on yam tubers in storage and as a result significantly reduced associated fungal infections caused by Fusarium spp., Penicillium spp., Aspergillus spp., Curvularia spp., Epicoccum spp. and Helminthosporium spp. (Morse et al., 2000).

In Cyprus, the development of resistance by P. citri to some insecticides was reported (Cyprus Agricultural Research Institute, 1981).

In quarantine, suspected host plants or plant parts may be dipped in insecticidal solutions such as an insecticidal soap composed of potassium salts of fatty acids with fluvalinate (Hansen et al., 1992a). Alternatively, plants may be subjected to vapour heat treatment as described by Hansen et al. (1992b). Anthurium andreanum, Leucospermum sp. and the flowers and foliage of orchids were, however, very sensitive to the vapour heat treatment.

In Australia, a greater reduction in fruit infected with P. citri in the calyx was achieved with low volume pesticide spraying than with a conventional high volume pesticide sprayer (Cunningham and Harden, 1999). The reduction in insecticide dose rate using lower spray volumes resulted in the pests not being controlled in some of the lower volume treatments.

Growth Regulators

Tests on insect growth regulators showed that kinoprene and epotenonane (R010-3108) gave effective control of P. citri comparable to that achieved with conventional insecticides such as dimethoate and methiodathion (French and Reeve, 1979).

The synthetic juvenile hormone analogues kinoprene (ZR-777) and hydroprene (ZR-512) prolonged the duration of the total life cycle and reduced female fecundity (Gaaboub et al., 1979), but female emergence was much less affected. In Switzerland, Vogel et al. (1977) showed that nymphs of P. citri treated with the juvenile hormone epofenonane moulted several times into intermediate forms between nymph and adult, the number of anal pores being reduced during each supernumerary moult.

In California, the insect growth regulators, ZR-619, ZR-512, ZR-520 and ZR-777 all reduced the number of progeny due to adult mortality as well as sterility induced in the survivors (Staal et al., 1973). In Ohio, insect growth regulators (kinoprene), Safer's soap and Murphy's Oil soap (a non-insecticide preparation) reduced the number of citrus mealybugs (presumably including P. citri) on foliage plants (Lindquist, 1981). The moulting inhibitor buprofenzin inhibited development of first-instar nymphs of P. citri and also (at lower concentrations) resulted in a marked decrease in the egg production of surviving females in laboratory studies in Hungary (Darvas and Szabo, 1987). Vogel et al. (1976) showed that applications of epofenonane could be as effective for the control of P. citri on grapefruits as dimethoate.

The insect growth regulator buprofezin was effective at three different application rates against P. citri on celery plants 26 days after treatment, when large numbers of dead early-instar larvae were observed on buprofezin-treated plants (Bedford et al., 1996). There was an indication that effects on mealybugs were greater, the higher the rate of buprofezin applied. However, low numbers of surviving mealybugs, within even the highest rate, indicated that a repeat application may be required for complete or continued control. No phytotoxic damage or response was observed on any of the celery plants within any of the treatments.

Pheromones

Several synthetic analogues of the sex pheromone dextro-cis-planococcyl acetate have been developed in the former USSR, Sweden, Israel, Japan and Italy (Rotundo and Tremblay, 1974, 1975, 1976a, 1980a, b; Dunkelblum et al., 1986). One analogue has been effectively used to monitor P. citri populations (Panis, 1979); its use for control purposes is quite promising (Rotundo and Tremblay, 1975) because it has proved highly attractive to males under field conditions.

Plant Extracts/Botanical Insecticides

Other unconventional chemical control agents include Citrus oil mixed with azinphos methyl which gave 96% mortality of P. citri in Texas (USA). In Florida, combinations of a hexane extract of the seed of neem (Azadirachta indica) and several of its chromatographic fractions significantly deterred feeding by P. citri and other insects (Jacobson et al., 1978).

In Alexandria (Egypt), application of emulsion sprays containing 3% petroleum distillate and various emulsifiers, all locally produced, gave good control on guavas for up to 5 weeks after treatment in September (El-Sebae and El-Akkawi, 1971).

In a field trial in Karnataka, India, neem oil and Pongamia [?pinnata] oil (both at 4%) were recommended for the control of P. citri on guava, causing 93.2 and 89.4% mortality of the pest, respectively, 10 days after the second spray (applied 10 days after the first) (Hussain et al., 1996).

Simmonds et al. (2000) demonstrated that a crude neem seed extract; a formulation of azadirachtin, a pyrethrum extract, and one of two naphthoquinones isolated from Calceolaria andina (BTG 504 and BTG 505) all influenced the foraging behaviour of C. montrouzieri exposed to leaves and P. citri. Larval and adult foraging behaviour was influenced most by BTG 504 and neem also affected larval behaviour; the predators contacted fewer treated leaves and spent less time on treated than on untreated leaves. The larvae also consumed fewer mealybugs treated with BTG 504 and BTG 505 than untreated mealybugs.

Integrated Pest Management (IPM)

In Florida, USA, a combination of selected chemicals, natural enemies and cultural practices was used to maintain populations of P. citri on Citrus below economic levels (Knapp, 1981).

In greenhouse experiments aimed at developing integrated control methods for P. citri and Nephus reunioni on potted orange trees, the predator C. montrouzieri effectively reduced populations of P. citri at a predator:prey ratio of 1:15 (Hamid et al., 1997). In most cases, no significant differences in pest reductions were detected between C. montrouzieri and insecticide.

On coffee in Kenya, P. citri was controlled indirectly by spraying chemicals such as chlorpyrifos or deltamethrin or applying hydramethylnon as baits against attendant ants in an attempt to encourage natural enemies that would otherwise be killed by the ants (Anon., 1990).

In Crete (Greece) no significant differences were detected between control of P. citri with C. montrouzieri alone and integrated control with C. montrouzieri and chemical treatment (Hamid and Michelakis, 1994). In Italy, however, Viggiani and Tranfaglia (1978) showed in laboratory experiments that the insecticides temephos, methomyl [now banned] and deltamethrin were harmful to L. dactylopii whereas fenbutatin oxide was harmless.

Arthur and Wiendl (1996) showed that irradiation had adverse effects on the development of P. citri. After 50 days of irradiation at different dosages, there were 2087.25 ± 301.34 individuals for the control, and 1348.25 ± 349.77, 288.25 ± 129.62, and 54.25 ± 61.98 individuals for doses of 10, 20 and 30 Gy, respectively. Insects treated with 40 Gy or higher dosages produced no offspring.

Katsoyannos (1996) reviewed progress on integrated pest management, especially biological control, for the main insect pests of citrus including P. citri, in northern Mediterranean countries.

In a field study of an integrated strategy for the control of P. citri, aphids and scale insects on Citrus in eastern Sicily (Italy) involving the control of attendant ants, chlorpyrifos gave the best control and was the least costly (Tumminelli et al., 1997). Insecticidal gum applied to the trunk gave better results than pyrethrum, but had phytotoxic effect on trees less than 10 years old and was also costly to apply. It was concluded that in integrated control programmes, the best time to treat ants and to release beneficial insects was between April and September.

A study in Italy on the biology, damage caused and control of the most important pests of table grapes, including P. citri and P. ficus, produced criteria for reducing the number of pesticide treatments in vineyards and a scheme for the biological control of pests and diseases on grapes (Guario and Laccone, 1996). Also in In Italy, Viggiani (1975a) observed that an infestation level of about 5% was regarded as the threshold for chemical control.

Gaps in Knowledge/Research Needs

Top of page

Molecular studies are needed on samples from many different countries to help establish how many cryptic species are in the Planococcus citri complex, to characterize them and to establish the geographic distribution of each of the cryptic species.

References

Top of page

Abbas MST, 1999. Studies on Dicrodiplosis manihoti Harris (Diptera, Cecidomyiidae), a common predator of mealybugs. Anzeiger fu^umlaut~r Scha^umlaut~dlingskunde, 72(5):133-134; 6 ref.

Abd El Rahim WA; Abdel Salam M; Abdel Wahab A; Kedr H, 1977. Evaluation of some insecticides for the control of pomegranate butterfly and citrus mealybug and their effects on physical and chemical characteristics. Indian Journal of Agricultural Sciences, 44(12):862-865.

Abdelkhalek L; Afellah M; Smaili C, 1998. Biology and biological control of Planococcus citri R. (Hom., Pseudococcidae) on citrus in the Loukos region of Morocco. Mededelingen - Faculteit Landbouwkundige en Toegepaste Biologische Wetenschappen, Universiteit Gent, 63(2b):483-488; 10 ref.

Abdullagatov AZ, 1978. Mealybug - a dangerous grapevine pest. Sadovodstvo, Vinogradarstvo i Vinodelie Moldavii No.6, 38-39.

Ackonor JB, 1997. Natural enemies of cocoa mealybugs. Rep. Cocoa Res. Inst. Ghana, 1995-96:166-169.

Ackonor JB, 2000. Studies on natural enemies of cocoa mealybugs. Rep. Cocoa Res. Inst. Ghana, 1996-97, 135-136.

Ackonor JB; Mordjifa DK; Boateng JK, 1993. Natural enemies of cocoa mealybugs. Rep. Cocoa Res. Inst. Ghana, 1989-90:45-47.

Afifi SA, 1968. Morphology and taxonomy of the adult males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Bulletin of the British Museum (Natural History) Entomology Supplement, 13:1-210.

Afifi SA; Amin AH; Nada; SMA, 1976. Taxonomy of males of two species of Pseudococcidae (Homoptera: Coccoidae). Bulletin de la Societe Entomologique d'Egypte, 60:141-151.

Afifi SA; Amin AH; Nada; SMA, 1976. Taxonomy of males of two species of Pseudococcidae (Homoptera: Coccoidae). Bulletin de la Societe Entomologique d'Egypte, 60:141-151.

Alphen JJM van; Xu CR, 1990. The role of host-plant odours in the attraction of Leptomastix dactylopii and Leptomastidea abnormis, parasitoids of the citrus mealybug, Planococcus citri. Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent, 55(2a):343-353

Amitava Konar, 1998. Seasonal incidence of mealybug species on mandarin orange in Darjeeling, West Bengal. Horticultural Journal, 11(2):105-110; 3 ref.

Angeles Martfnez M de los; Hernández M; Ceballos M, 1992. Ecological relationships of natural enemies of Planococcus citri (Risso) in III Frente, Santiago de Cuba. Revista de Protección Vegetal, 7(1):27-29; 15 ref.

Angeles Martfnez M de los; Mestre N; Fraga N, 1991. Bioecology of Planococcus citri (Risso) (Homoptera: Pseudococcidae). Revista de Protección Vegetal, 6(1):37-42; 3 ref.

Anon., 1990. Integrated control of the coffee mealybugs and scales forming moulds. Kenya Coffee, 55(648):947

Anon., 2016. Mealybugs - outdoors. Help for the home gardener. St Louis, Missouri, USA: Missouri Botanic Garden. http://www.missouribotanicalgarden.org/gardens-gardening/your-garden/help-for-the-home-gardener/advice-tips-resources/pests-and-problems/insects/mealybugs/mealybugs-outdoors.aspx

Anstead DR, 1917. Coffee borer in Indo-China. Planters' Chronicle, 12(48):608.

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).

Arai T, 1996. Temperature-dependent development rate of three mealybug species, Pseudococcus citriculus Green, Planococcus citri (Risso), and Planococcus kraunhiae (Kuwana) (Homoptera: Pseudococcidae) on Citrus. Japanese Journal of Applied Entomology and Zoology, 40(1):25-34; [En captions and tables]; 15 ref.

Arriola Padilla VJ; Estrada Martínez E; Romero Nápoles J; González Hernández H; Pérez Miranda R, 2016. Scale insects (Hemiptera: Coccomorpha) on ornamental plants in greenhouses from the central zone of the Morelos state, Mexico. (Insectos escama (Hemiptera: Coccomorpha) en plantas ornamentales en viveros de la zona centro del estado de Morelos, México.) Interciencia, 41(8):552-560. http://www.interciencia.org/v41_08/indexe.html

Arthur V; Wiendl FM, 1996. Irradiation of Planococcus citri (Risso) (Homoptera: Pseudococcidae) with gamma radiation from cobalt-60 to determine the disinfestation dose. Anais da Sociedade Entomolo^acute~gica do Brasil, 25(2):345-346; 3 ref.

Aykac MK; Erguder TM, 1972. A study of control measures against Planococcus citri Risso in the vineyards of Tokat province. Tarim Bakanligi Zirai Mucadele ve Zirai Karantina Gene Mudurlugu Arastirma Subesi: Plant Protection Research Annual.: Zirai Mucadele Arastirma Yilligi, 43:171-172.

Baaren J van; Nenon JP, 1994. Factors involved in host discrimination by Epidinocarsis lopezi and Leptomastix dactylopii (Hym., Encyrtidae). Journal of Applied Entomology, 118(1):76-83

Baaren J van; Nénon JP, 1996. Host location and discrimination mediated through olfactory stimuli in two species of Encyrtidae. Entomologia Experimentalis et Applicata, 81(1):61-69; 37 ref.

Baaren J van; Nénon JP, 1996. Intraspecific larval competition in two solitary parasitoids, Apoanagyrus (Epidinocarsis) lopezi and Leptomastix dactylopii. Entomologia Experimentalis et Applicata, 81(3):325-333; 3 pp. of ref.

Ballou HA, 1921. Report on a visit to Grenada. Agriculture News, Bridgetown, 20(503):250-251.

Barbagallo S; Longo S; Rapisarda C; Siscaro G, 1993. Status of the biological control against citrus whiteflies and scale insects in Italy. Bulletin OILB/SROP, 16(7):7-15

Barbier R; Raimbault JP, 1985. Formation of spherulp around the egg-shell in Leptomastix dactylopii (Hymenoptera, Encyrtidae). Bulletin de la Societe Zoologique de France, 110(3):339-346

Barrett JH, 1966. Insect pests of Coffea arabica in the New Guinea Highlands a general survey with notes on bionomics and control. Papua New Gum. agric. J, 18(3):83-100 pp.

Barrie JW, 1958. Coffee in the Highlands. Papua New Guinea Agricultural Journal, 11(1):1-29.

Bartlett BR, 1978. Introduced parasites and predators of arthropod pests and weeds. Section on Pseudococcidae. Agricultural Handbook of US Department of Agriculture, 480:150-153.

Baskaran RKM; Lakshmi LG; Uthamasamy S, 1999. Comparative biology and predatory potential of Australian ladybird beetle (Cryptolaemus montrouzieri) on Planococcus citri and Dactylopius tomentosus. Indian Journal of Agricultural Sciences, 69(8):605-606; 13 ref.

Baum H, 1970. Investigations on the root mealybug complex of Planococcus citri (Risso) on Coffea arabica in Kenya/East Africa. Zeitschrift fur Angewandte Entomologie, 65(3):295-304.

Bazarov BB, 1988. The grape mealybug. Zashchita Rastenii (Moskva), No. 8:29-30

Beardsley JW, 1966. Insects of Micronesia. Homoptera: Coccoidea. Insects of Micronesia, 6(7):377-562 pp.

Beatty HA, 1944. The insects of St. Croix, V.I. Journal of the Agricultural University of the Virgin Islands, 28(3/4):114-172.

Bedford ID; Kelly A; Markham PG, 1996. The effects of buprofezin against the citrus mealybug Planococcus citri. Brighton Crop Protection Conference: Pests & Diseases - 1996: Volume 3: Proceedings of an International Conference, Brighton, UK, 18-21 November 1996, 1065-1070; 7 ref.

Benassy C; Deportes L; Onillon J-C; Panis A, 1976. Orientation towards integrated control in citrus culture in south-east France. (Orientation vers la lutte integree en agrumiculture dans le sud-est de la France.) Pepinieristes Horticulteurs Maraichers, No.167:41-48.

Ben-Dov Y, 1994. A systematic catalogue of the mealybugs of the world (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with data on geographical distribution, host plants, biology and economic importance. Andover, UK; Intercept Limited, 686 pp.

Berlinger MJ; Dahan R; Podoler H; Segre L, 1985. Preliminary survey of insects attacking jojoba in southern Israel and of their natural enemies. Proceedings of the sixth international conference on jojoba and its uses, 21-26 October 1984, Israel Tucson, AZ 85719, USA; University of Arizona Press, 219-223

Berlinger MJ; Gol' berg AM, Chapman RF ed. , Bernays EA, 1978. The effect of the fruit sepals on the citrus mealybug population and on its parasite. Proceedings of the 4th International Symposium - Insect and Host Plant held at Fulmer Grange, Slough, England, 4-9 June 1978. Entomologia Experimentalis et Applicata, 24:238-243.

Berlinger MJ; Tzahor J; Gol' berg AM, 1979. Contribution to the phenology of Chilocorus bipustulatus L. (Coccinellidae) in citrus groves and the control of Planococcus citri (Pseudococcidae) in Israel. [XXXI International Symposium on Crop Protection.]: XXXI Internationaal Symposium over Fytofarmacie en Fytiatrie. Mededelingen van de Faculteit Landbouwwetenschappen Rijksuniversiteit Gent, 44:49-54.

Bernard, 1926. Verslag over het Jaar 1925. Meded, Proefstn. Mid. Java, 42. Salatiga.

Betrem JG, 1936. Gegevens omtrent de biologie van de dompolanluis en de lamtorolois', arch. Koffiecult. Ned. Indie, 10:43-84.

Bigger M, 1975. Recent work on the mealybug vectors on cocoa swollen shoot virus. In: Kumar R, ed. Proceedings of the 4th Conference of West African Cocoa Entomologists, Zoology Department, University of Ghana, Legon, Ghana, 9th-13th December 1974. Zoology Department, University of Ghana. Legon Ghana, 62-66

Bigger M, 1975. Susceptibility of two cocoa progenies to attack by insect species. I. Proportion of trees infested. Experimental Agriculture, 11(3):187-192

Bigger M, 1975. Susceptibility of two cocoa progenies to attack by insect species. II. Effects of canopy size on numbers of mealybugs. Experimental Agriculture, 11(3)193-199.

Bigger M, 1976. Virus outbreak survey. In: Report of the Cocoa Research Institute, Ghana, 1973-74. pp. 101-107.

Bigger M, 1981. The relative abundance of the mealybug vectors (Hemiptera: Coccidae and Pseudococcidae) of cocoa swollen shoot disease in Ghana. Bulletin of Entomological Research, 71(3):435-448

Bitaw AA; Saad AA, 1990. Survey of natural enemies of date palm pests in Libya. Arab Journal of Plant Protection, 8(1):12-15

Bivins JL; Deal AS, 1973. Systemic insecticides for control of Citrus mealybug in gardenias. California Agriculture, 27(8):5-6

Blumberg D; Klein M; Mendel Z, 1995. Response by encapsulation of four mealybug species (Homoptera: Pseudococcidae) to parasitization by Anagyrus pseudococci. Phytoparasitica, 23(2):157-163

Bongiorni S; Cintio O; Prantera G, 1999. The relationship between DNA methylation and chromosome imprinting in the coccid Planococcus citri. Genetics, 151(4):1471-1478; 32 ref.

Brimblecombe AR, 1962. Studies of the Coccoidea 12. Species occurring on deciduous fruit and nut trees in Queensland. Queensland Journal of Agricultural Science, 19:219-229.

Brooks HM, 1957. The Coccidae (Homoptera) naturalised in South Australia: an annotated list. Transactions of the Royal Society of South Australia, 80:81-90.

Cabaleiro C; Segura S, 1997. Field transmission of grapevine leafroll associated virus 3 (GLRaV-3) by the mealybug Planococcus citri. Plant Disease, 81(3):283-287; 38 ref.

CABI/EPPO, 1999. Planococcus citri. [Distribution map]. Distribution Maps of Plant Pests, June (2nd revision). Wallingford, UK: CAB International, Map 43.

CABI/EPPO, 1999. Planococcus citri. [Distribution map]. Distribution Maps of Plant Pests, June (2nd revision). Wallingford, UK: CAB International, Map 43.

Cabitza F; Cubeddu M; Maurichi S; Ballore S; Pala M; Lovicu G, 1994. Application of the integrated control technique to the control of the Mediterranean fruit fly on clementines in Sardinia. Informatore Agrario, 50(24):45-48

Cadée N; Alphen JJM van, 1997. Host selection and sex allocation in Leptomastidea abnormis, a parasitoid of the citrus mealybug Planococcus citri. Entomologia Experimentalis et Applicata, 83(3):277-284; 23 ref.

Campbell CAM, 1975. The distribution of mealybug vectors of CSSV within trees. In: Kumar R, ed. Proceedings of the 4th Conference of West African Cocoa Entomologists, Zoology Department, University of Ghana, Legon, Ghana, 9th-13th December 1974. Zoology Department, University of Ghana. Legon Ghana, 67-71

Campbell CAM, 1994. Homoptera associated with the ants Crematogaster clariventris, Pheidole megacephala and Tetramorium aculeatum (Hymenoptera: Formicidae) on cocoa in Ghana. Bulletin of Entomological Research, 84(3):313-318

Carpenter JM; Kassanis B; Jones P; Dabek AJ; Plumb RT, 1976. Properties of viruses and virus diseases. Rothamsted Experimental Station Report for 1975. Part 1, 243-246.

Carter W, 1942. Geographical distribution of mealybug wilt with some other insect pest of pineapple. Journal of Economic Entomology, 35:10-15.

Ceballo FA; Papacek D; Walter GH, 1998. Survey of mealybugs and their parasitoids in south-east Queensland citrus. Australian Journal of Entomology, 37(3):275-280; 29 ref.

Cipolla C; Lugo G; Sassi C; Belisario A; Nucci MC; Palermo A; Pescarelli MA; Nobile M; Raffi GB, 1997. Hypersensitivity and allergic diseases in a group of workers employed in breeding insects for biological pest control. Medicina del Lavoro, 88(3):220-225; 6 ref.

Copland MJW; Perera HAS; Heidari M, 1993. Influence of host plant on the biocontrol of glasshouse mealybug. Bulletin OILB/SROP Route de Marseilles - BP 91, Netherlands; International Organization for Biological Control of Noxious Animals and Plants (OIBC/OILB), West Palaearctic Regional Section (WPRS/SROP), 16(8):44-47

Cotterell GS, 1943. Entomology section, Report Cent. Cacao Research Station, Tafo, Gold Coast, 1938, 42:51-54.

Cox JM, 1983. An experimental study of morphological variation in mealybugs (Homoptera: Coccoidea: Pseudococcidae). Systematic Entomology, 8(4):361-382

Cox JM, 1989. The mealybug genus Planococcus (Homoptera: Pseudococcidae). Bulletin of the British Museum (Natural History), Entomology, 58(1):1-78

Cox JM; Freeston AC, 1985. Identification of mealybugs of the genus Planococcus (Homoptera: Pseudococcidae) occurring on cacao throughout the world. Journal of Natural History, 19(4):719-728

Culik MP; Ventura JA; Martins Ddos S, 2009. Scale insects (Hemiptera: Coccidae) of pineapple in the State of Espírito Santo, Brazil. Acta Horticulturae [Proceedings of the Sixth International Pineapple Symposium, Joao Pessoa, Brazil, 18-23 November 2007.], No.822:215-218. http://www.actahort.org

Cunningham GP; Harden J, 1999. Sprayers to reduce spray volumes in mature citrus trees. Crop Protection, 18(4):275-281; 13 ref.

Cyprus Agricultural Research Institute, 1981. Annual report for 1980. Annual report for 1980, 87 pp.

Darvas B; Szabo L, 1987. Effect of moulting inhibitors (buprofezin and diflubenzuron) on Acyrtosiphon pisum (Hom., Aphididae), Planococcus citri (Hom., Pseudococcidae) and Quadraspidotus perniciosus (Hom., Diaspididae). Novenyvedelem, 23(8):343-351

De Fluiter HJ, 1937. Over den invloed van het voedsel op insecten, Versl. 16e Vergad. Ver. Proefstn. Person., Djember., Oct. 1936, 153-70.

De Fluiter HJ, 1939. Het witte luis-vraagstuk bij de Koffie. Bergcultures, 13:760-765.

De Lotto G, 1964. Observation on African mealybugs (Homoptera: Pseudococcidae). Bulletin of the British Museum (Natural History) Entomology, 143:341-397.

De Lotto G, 1975. Notes on the vine mealybug (Homoptera: Coccoidea: Pseudococcidae). Journal of the Entomological Society of Southern Africa, 38(2):125-130.

Delabie JHC, 1988. Occurrence of Wasmannia auropunctata (Roger, 1863) (Hymenoptera, Formicidae, Myrmicinp) in cocoa plantations in Bahia, Brazil. Revista Theobroma, 18(1):29-37

Delabie JHC; Cazorla IM, 1991. Damage caused by Planococcus citri Risso (Hemiptera: Pseudococcidae) in the production of cocoa. Agrotropica, 3(1):53-57

Donald RG, 1955. Mealybug studies. Annual Report of West African Cocoa Research Institute, 1954-1955, 101-104.

Donald RG, 1956. The natural enemies of some Pseudococcidae in the Gold Coast. Journal of West African Science Association, 2(1):48-60.

Dufour B, 1991. The role and importance of different insect species in the ecology of CSSV (cacao swollen shoot virus) in Togo. Cafe, Cacao, The, 35(3):197-204

Dunkelblum E; Ben Dov Y; Goldschmidt Z; Wolk JL; Somekh L; Dov Y Ben, 1986. Synthesis and field bioassay of the sex pheromone and some analogues of the citrus mealybug, Planococcus citri (Risso). In: Proceedings of the Fifth International Symposium on Scale Insect Studies, Portici, Italy, 24-28 June, 1986. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Italy, 43: Supplement, 149-154.

DuPasquier R, 1930. Recherches sur les Hymenopteres parasites du Borer (Xylotrechus quadripes Chev.),. Proc. 4th Pac. Sci. Cong. Java 1929, iv, Agric. Papers:519-24, Batavia.

El-Sebae AH; El-Akkawi MM, 1971. Studies on the chemical structure and insecticidal activity. IV. Efficiency of locally formulated spray mineral oils. Alexandria Journal of Agricultural Research, 19(1):131-138

Entwistle PF, 1958. Mealybugs: II. Biology with special reference to cocoa. The Ghana Cocoa farmer, 2(2):64-69.

Entwistle PF, 1972. Pests of coffee. London, UK: Longman Group Limited.

Ezzat YM; McConnell; HS, 1956. A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the University of Maryland Agricultural Experimental Station, A-84:1-108.

Fennah RG, 1959. Nutritional factors associated with the development of mealybugs on cocoa. Annual Report of Cocoa Research Institute, Trinidad, 1957-1958. Trinidad: Cocoa Research Institute, 19-28.

Fernandes IM, 1990. Contribution to knowledge of the Coccoidea (Homoptera) of Portugal. I - Annotated list of the coccoids of the garden of the Zoological Centre. Garcia de Orta. Serie de Zoologia, 17(1-2):59-63

Ferraro M; Epifani C; Bongiorni S; Nardone AM; Parodi-Delfino S; Prantera G, 1998. Cytogenetic characterization of the genome of mealybug Planococcus citri (Homoptera, Coccoidea). Caryologia, 51(1):37-49; 18 ref.

Ferris GF, 1950. Atlas of the Scale Insects of North America. Series V. The Pseudococeidae (Part I). Stanford Univ. Pr., California; London, G. Cumberlege, Oxford Univ. Pr., vii+ 278 pp.

Firempong S, 1984. Laboratory and field evaluation of cocoa progenies for resistance to mealybug vectors (Hemiptera: Pseudococcidae) of swollen shoot virus. Bulletin of Entomological Research, 74(1):97-102

Foldi I; Kozár F, 2006. New species of Cataenococcus and Puto from Brazil and Venezuela, with data on others species (Hemiptera, Coccoidea). Nouvelle Revue d'Entomologie, 22(4):305-312.

French JV; Reeve JR, 1978. Relationship of vines to management of other pests on Texas citrus. Journal of the Rio Grande Valley Horticultural Society, 32:67-70.

French JV; Reeve RJ, 1979. Insect growth regulators and conventional insecticides for suppression of citrus mealybug. Southwestern Entomologist, 4(3):238-243

Fronteddu F; Canu D; D'Amico R; Delpiano N; Fancello F; Nanni G, 1996. Applications of integrated control methods in citrus cultivation: biotechnical control against Ceratitis capitata and biological control of Planococcus citri. Informatore Fitopatologico, 46(11):34-39; 10 ref.

Gaaboub IA; Rawash IA; Abdel-Lattif MA; El-Minshawy A; Abdel-Rahman AM, 1979. Biological studies and effect of treatment with two juvenile hormone mimics on the developmental stages of Icerya purchasi (Mask) and Planococcus citri (Risso). Mededelingen van de Faculteit Landbouwwetenschappen Rijksuniversiteit Gent, 44:185-203

Garcia JF; O'Neil RJ, 2000. Effect of Coleus size and variegation on attack rates, searching strategy, and selected life history characteristics of Cryptolaemus montrouzieri (Coleoptera: Coccinellidae). Biological Control, 18(3):225-234; 41 ref.

García M; Denno B; Miller DR; Miller GL; Ben-Dov Y; Hardy NB, 2016. ScaleNet: a literature-based model of scale insect biology and systematics. http://scalenet.info

Gausman HW; Hart WG, 1974. Reflectance of four levels of sooty-mold deposits produced from the honedew of three insect species. Journal of the Rio Grande Valley Horticultural Society, 28:131-136.

Georgala MB; Buitendag CH; Hofmeyr JH; Kellerman C, 1975. Recommendations for the control of major citrus pests and diseases on bearing trees during the 1975/76 season. Part I. Citrus and Sub Tropical Fruit Journal, 500(9):11-13, 15-22.

Gerson U, 2016. Planococcus citri. Pest plants of the Middle East. Jerusalem, Israel: The Hebrew University of Jerusalem.

Gibson RW; Turner RH, 1977. Insect-trapping hairs on potato plants. PANS, 23(3):272-277

Giliomee JH, 1961. Morphological and taxonomic studies on the males of three species of the genus Pseudococcus (Hemiptera: Coccoidea). Annale Universiteit van Stellenbosch, 36:239-296.

Gollifer DE; Jackson GVH; Dabek AJ; Plumb RT; May YY, 1977. The occurrence and transmission of viruses of edible aroids in the Solomon Islands and the Southwest Pacific. PANS, 23(2):171-177

González RH, 2011. Pseudococcids of fruit production in Chile (Hemiptera: Pseudococcidae). (Pseudocóccidos de importancia frutícola en Chile (Hemiptera: Pseudococcidae).) In: Extension, Department of the Faculty of Ciencias, Serial Publications in Agriculture. Santiago, Chile: University of Chile, 186 pp.

Gowdey CC, 1917. A list of Uganda Coccidae, their food plants and natural enemies. Bulletin of Entomological Research, 8:187-189.

Granara de Willink MC; Claps LE, 2003. Scale insects (Hemiptera: Coccoidea) present in ornamentals in Argentina. (Cochinillas (Hemiptera: Coccoidea) presentes en plantas ornamentales de la Argentina.) Neotropical Entomology, 32(4):625-637.

Gravitz N; Wilson C, 1968. A sex pheromone from the citrus mealybug. Journal of Economic Entomology, 61:1458-1459.

Gray HE, 1954. The development of the citrus mealybug. Journal of Economic Entomology, 47:174-176.

Gross S; Dreishpoun Y; Blachinski D; Shmueli S; Steinberg S; Mendel Z, 1999. Cork scars on fruits of the citrus variety 'Sweetie' as related to infestation by the citrus mealybug. Alon Hanotea, 53(11):463-468.

Guario A; Laccone G, 1996. The defence of table grapes from pests. Informatore Agrario Supplemento, 52(50):31-40.

Halima-Kamel MB; Germain JF; Mdellel L; Abdelaoui K, 2014. Phenacoccus madeirensis (Hemiptera: Pseudococcidae): a new species of mealybug in Tunisia. Bulletin OEPP/EPPO Bulletin, 44(2):176-178. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2338

Hamid HA; Michelakis S, 1994. The importance of Cryptolaemus montrouzieri Mulsant (Col., Coccinellidae) in the control of the citrus mealybug Planococcus citri (Hom., Coccoidea) under specific conditions. Journal of Applied Entomology, 118(1):17-22

Hamid HA; Michelakis SE; Vacante V, 1997. The use of Cryptolaemus montrouzieri (Mulsant) for the control of Planococcus citri (Risso) in Crete, Greece. Proceedings of the meeting held at Florence, Italy, 29 August 1996. Bulletin OILB-SROP, 20(7):7-12.

Hansen JD; Hara AH; Tenbrink VL, 1992. Insecticidal dips for disinfesting commercial tropical cut flowers and foliage. Tropical Pest Management, 38(3):245-249

Hansen JD; Hara AH; Tenbrink VL, 1992. Vapor heat: a potential treatment to disinfest tropical cut flowers and foliage. HortScience, 27(2):139-143

Hara AH; Hata TY; Hu BKS; Tsang MMC, 1997. Hot-air induced thermotolerance of red ginger flowers and mealybugs to postharvest hot-water immersion. Postharvest Biology and Technology, 12(1):101-108; 12 ref.

Hargreaves E, 1937. Entomological work. Report of the Department of Agriculture, Sierra Leone, 1931, 18-20.

Hargreaves H, 1925. Report of the Government Entomologist. In: Report of the Department of Agriculture, Uganda, 1924. Uganda: Department of Agriculture, 21-28.

Hata TY; Hara AH, 1992. Evaluation of insecticides against pests of red ginger in Hawaii. Tropical Pest Management, 38(3):234-236

Hickman GW, 1981. Mealybug control on greenhouse foliage plants. Flower and Nursery Report. 1981, Spring, 3-4.

Hussain MA; Puttaswamy; Viraktamath CA, 1996. Management of citrus mealybug, Planococcus citri Risso on guava using botanical oils. Insect Environment, 2(3):73-74.

Islam KS; Copland MJW, 1997. Host preference and progeny sex ratio in a solitary koinobiont mealybug endoparasitoid, Anagyrus pseudococci (Girault), in response to its host stage. Biocontrol Science and Technology, 7(3):449-456; 23 ref.

Islam KS; Jahan M, 1992. Influence of different types of food on the reproductive performance of Anagyrus pseudococci (Hymenoptera: Encyrtidae). Bangladesh Journal of Entomology, 2:17-22; 15 ref.

Islam KS; Jahan M, 1993. Oviposition and development of the mealybug parasitoid Anagyrus pseudococci (Girault) at different constant temperatures. Pakistan Journal of Scientific and Industrial Research, 36(8):322-324

Islam KS; Perera HAS; Copland MJW, 1997. The effects of parasitism by an encyrtid parasitoid, Anagyrus pseudococci on the survival, reproduction and physiological changes of the mealybug, Planococcus citri. Entomologia Experimentalis et Applicata, 84(1):77-83; 27 ref.

Izhar J, 1999. Preliminary experiment to control ants as a transferring factor of millibug (Pseudococcus citriculus, Planococcus citri) in persimmon. Alon Hanotea, 53(12):490-492; 2 ref.

Jacobson M; Reed DK; Crystal MM; Moreno DS; Soderstrom EL; Chapman RF; Bernays EA, 1978. Chemistry and biological activity of insect feeding deterrents from certain weed and crop plants. Proceedings of the 4th International Symposium - Insect and Host Plant - held at Fulmer Grange, Slough, England, 4-9 June 1978. Entomologia Experimentalis et Applicata, 24:448-457.

Jadhav RG; Madane NP; Kathamale DK, 1996. Record of soybean as a new host in India for citrus mealybug. Insect Environment, 2(3):90; 1 ref.

James HC, 1932. Coffee mealybug research. Department of Agriculture Kenya Bulletin, 18:18.

James HC, 1937. Sex ratios and the status of the male Pseudococcinae. Bulletin of Entomological Research, 28:429-461.

Kanika-Kiamfu J; Iperti G; Brun J, 1993. Study of food consumption of Exochomus flaviventris (Col. Coccinellidae), predator of Phenacoccus manihoti (Hom.: Pseudococcidae). Entomophaga, 38(3):291-298; 8 ref.

Katsoyannos P, 1996. Integrated insect pest management for citrus in Northern Mediterranean countries. Benaki Phytopathologique Institute Report, Athens; Greece: 110 pp.

Kawai S, 1987. The coccid fauna of the Ogasawara (Bonin) Islands. Development, Technique and Cooperation in Agriculture: A Collection of Papers to Commemorate the 30th Anniversary. Tokyo, Japan: Tokyo University of Agriculture.

Kawai S; Matsubara Y; Umesawa K, 1971. A preliminary revision of the Coccoidea-fauna of the Ogasawara (Bonin) Islands (Homoptera : Coccoidea) [51 species]. Applied Entomology and Zoology, 6(1):11-26.

Kenten RH; Woods RD, 1976. A virus of the cocoa swollen shoot group infecting cocoa in North Sumatra. PANS, 22(4):488-490

Khalilov BB, 1972. Pests of grape vines in Azerbaidzhan. Zashchita Rastenii, 17(2):36-40

Khandakar Shariful Islam; Jahan M, 1993. Influence of honeydew of citrus mealybug (Planococcus citri) on searching behaviour of its parasitoid, Anagyrus pseudococci. Indian Journal of Agricultural Sciences, 63(11):743-746

Kinawy MM; Al-Waili HM; Almandhari AM, 2008. Review of the successful classical biological control programs in Sultanate of Oman. Egyptian Journal of Biological Pest Control [Proceedings of the 2nd Arab Conference of Applied Biological Pest Control, Cairo, Egypt, 7-10 April 2008.], 18(1):1-10. http://www.esbcp.org

Kirkpatrick TW, 1953. Notes on minor insect pests of cocoa in Trinidad. Report on Cocoa Research, 1952, 62-71.

Knapp JL, 1981. Integrated pest management in Florida citrus. Miscellaneous Publications of the Entomological Society of America, 12:91-98

Kontodimas DC, 1997. First record of the predatory insect Nephus bisignatus (Boheman) (Coleoptera: Coccinellidae) in Greece. Annales de l'Institut Phytopathologique Benaki, 18(1):61-63; 10 ref.

Kozár F; Fowjhan MA; Zarrabi M, 1996. Check-list of Coccoidea and Aleyrodoidea (Homoptera) of Afghanistan and Iran, with additional data to the scale insects of fruit trees in Iran. Acta Phytopathologica et Entomologica Hungarica, 31:61-74.

Kozár F; Guignard E; Bachmann F; Mani E; Hippe C, 1994. The scale insect and whitefly species of Switzerland (Homoptera: Coccoidea and Aleyrodoidea). Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 67(1-2):151-161; 24 ref.

Krambias A; Kontzonis A, 1980. Establishment of Leptomastix dactylopii (How.) in Cyprus. Fruits, 35(12):783-785

Krishnamoorthy A, 1990. Evaluation of permanent establishment of Leptomastix dactylopii How. against Planococcus citri (Risso) in Citrus orchards in India. Fruits (Paris), 45(1):29-32

Krishnamoorthy A; Mani M, 1989. Records of green lacewings preying on mealybugs in India. Current Science, 58(3):155-156

Krishnamoorthy A; Mani M, 1994. Manage mealybug in citrus orchards. Indian Horticulture, 39(1):12-13

Krishnamoorthy A; Mani M, 1996. Record of hyperparasitoids on exotic parasitoid Leptomastix dactylopii How. parasitizing citrus mealybug Planococcus citri (Risso) in India. Entomon, 21(1):111-112; 5 ref.

Kurdyukov VV; Alan MN, 1973. Damage caused by citrus mealy bug, Pseudococcus citri [on grapevines] and its chemical control. Byulleten' Vsesoyuznogo Nauchno Issledovatel'skogo Instituta Zashchity Rastenii, No. 26, 26-30.

Le Pelley RH, 1943. An Oriental mealybug (Pseudococcus lilacinus Ckll.) (Hemiptera) and its insect enemies. Transactions of the Royal Entomological Society of London, 93(1):73-93.

Lim GT, Ooi PAC (ed. ), Lim GS (ed.), Teng PS, 1992. Recent development of cocoa insect pests management in Sabah Malaysia. Proceedings of the 3rd International Conference on Plant Protection in the Tropics, No. 4:36-53.

Limon de la Oliva F; Blasco Pascual J, 1973. Preliminaries for the study of natural control and of measures to use against pests of Citrus in the northern part of the Levante region, with a view to the establishment of a programme of integrated control. Boletin Informativo de Plagas, No. 109:69-86

Limon de la Oliva F; Blasco Pascual J; Vicente Lopez S; Verniere Fernandez C, 1972. Control tests against the citrus mealybug on Citrus. Boletin Informativo de Plagas, No.99:57-65

Lindquist RK, 1979. Insecticide evaluations for spider mite, aphid, and mealybug control on selected foliage plants in 1978. Ohio Florists' Association Bulletin, 596:5-7, 9.

Lindquist RK, 1981. Controlling the citrus mealybug on greenhouse foliage plants. Ohio Florists' Association Bulletin, 622:6-8.

Liotta G; Mineo G; Ragusa S, 1976. On the current state of knowledge concerning certain arthropods injurious to citrus in Sicily. [Sur lnetat actuel des connaissances concernant certains arthropods nuisibles aux agrumes en Sicile.] Bollettino dell'Istituto di Entomologia Agraria e dell'Osservatorio di Fitopatologia di Palermo, 10:29-67

Liotta G; Mineo G; Ragusa S, 1976. On the current state of knowledge concerning certain arthropods injurious to citrus in Sicily. [Sur lnetat actuel des connaissances concernant certains arthropods nuisibles aux agrumes en Sicile.] Bollettino dell'Istituto di Entomologia Agraria e dell'Osservatorio di Fitopatologia di Palermo, 10:29-67

Lockhart BEL; Olszewski NE, Loebenstein G (ed. ), Hammond J (ed.), Gera A (ed.), Derks AFLM (ed.), Zaayen A van, 1996. Schefflera ringspot virus, a widely distributed mealybug-transmitted badnavirus occurring in Schefflera and Aralia. Proceedings of the 9th international symposium on virus diseases of ornamental plants, Herzliya, Israel, 17-22 March 1996. Acta Horticulturae, No. 432:196-202.

Malumphy C; Badmin JS, 2012. Scale insects and whiteflies (Hemiptera: Coccoidea and Aleyrodoidea) of Watsonian Kent; with a discussion on the impact of naturalised non-native species. British Journal of Entomology and Natural History, 25:15-49.

Mancion J; Aliber H, 1936. La production du cafe au Togo et quelques insectes depredateurs du cafeier. Agron. Colon., 224:33-43.

Mani M, 1994. Effectiveness of the exotic encyrtid parasitoid, Leptomastix dactylopii How. in the control of mealybug, Planococcus citri (Risso) in guava orchards. Journal of Entomological Research, 18(4):351-355

Mani M, 1995. Comparative development, progeny production and sex ratio of the exotic parasitoid Leptomastix dactylopii Howard (Hym., Encyrtidae) on Planococcus lilacinus and P. citri (Homop., Pseudococcidae). Entomon, 20(1):23-26

Mani M; Krishnamoorthy A, 1990. Natural suppression of mealybugs in guava orchards. Entomon, 15(3-4):245-247

Mani M; Krishnamoorthy A, 1990. Outbreak of mealybugs and record for their natural enemies on pomegranate. Journal of Biological Control, 4(1):61-62

Mani M; Krishnamoorthy A, 1990. Predation of Mallada boninensis on Ferrisia virgata, Planococcus citri and P. lilacinus. Journal of Biological Control, 4(2):122-123

Mani M; Krishnamoorthy A, 1997. Suppression of Planococcus citri (Risso) on sapota. Pest Management in Horticultural Ecosystems, 3(1):45-47; 8 ref.

Mani M; Lakshmi VJ; Krishnamoorthy A, 1997. Side effects of some pesticides on the adult longevity, progeny production and prey consumption of Cryptolaemus montrouzieri Mulsant (Coccinellidae, Coleoptera). Indian Journal of Plant Protection, 25(1):48-51; 17 ref.

Mani M; Thontadarya TS, 1987. Record of mealybug species on grapevine in Karnataka. Current Science, India, 56(22):1192

Martínez AM; Bravo N, 1989. Aspergillus flavus Dirk., natural control of different species of mealybugs. Revista de Protección Vegetal, 4(1):83-84; 3 ref.

Matile-Ferrero D, 1978. Cassava mealybug in the People's Republic of Congo. In: Nwanze KF, Leuschner K, ed. Proceedings of the International Workshop on the cassava mealybug Phenacoccus manihoti Mat.-Ferr. (Pseudococcidae) held at INERA-M'vuazi, Bas-Zaire, Zaire, June 26-29, 1977. International Institute of Tropical Agriculture. Ibadania Niger, 29-46

Matile-Ferrero D; Étienne J, 2006. Mealybugs of the French Antilles and some other Caribbean islands. (Cochenilles des Antilles françaises et de quelques autres îles Caraïbes (Hemiptera, Coccoidea).) Revue Française d'Entomologie, 28(4):161-190.

McComie LD, 1987. The soursop (Annona muricata L.) in Trinidad: its importance, pests and problems associated with pest control. Journal of the Agricultural Society of Trinidad and Tobago, No. 87:42-55

McKenzie HL, 1967. Mealybugs of California, with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). viii+526 pp.

Mendel Z; Blumberg D; Wysoki M, 1992. Biological control of four homopterans in Israeli horticulture: achievements and problems. Phytoparasitica, 20(suppl.):45-49

Mendel Z; Blumberg D; Zehavi A; Weissenberg M, 1992. Some polyphagous Homoptera gain protection from their natural enemies by feeding on the toxic plants Spartium junceum and Erythrina corallodendrum (Leguminosae). Chemoecology, 3(3-4):118-124

Merlin J; Lemaitre O; GrTgoire JC, 1996. Oviposition in Cryptolaemus montrouzieri stimulated by wax filaments of its prey. Entomologia Experimentalis et Applicata, 79(2):141-146; 26 ref.

Mestre N; Angeles Martfnez Mde los, 1991. Tegumental ultrastructure of Planococcus citri (Risso) (Homoptera: Pseudococcidae). Revista de Proteccio^acute~n Vegetal, 6(1):32-36; 4 ref.

Meyerdirk DE; Chandler LD; Summy KR; Hart WG, 1981. Spatial distribution of citrus mealybug on grapefruit trees. Journal of Economic Entomology, 74(6):662-664

Meyerdirk DE; Hart WG; Dean HA, 1978. Two newly established primary parasites, Leptomastix dactylopii Howard and Anagyrus sp., found attacking Planococcus citri (Risso) in Texas. Southwestern Entomologist, 3(4):295-298

Milek TM; Šimala M; Krcmar S, 2008. Species of the genus Planococcus Ferris, 1950 (Hemiptera: Coccoidea: Pseudococcidae) with special regard to Planococcus vovae (Nasonov, 1908) as a species newly recorded in Croatia. Natura Croatica, 17(3):157-168. http://hrcak.srce.hr/index.php?show=casopis&id_casopis=51

Mineo G, 1992. Current pests of citriculture in Italy. Istituto di Entomologia Agraria, Universita di Palermo, Italy. Rivista di Frutticoltura e di Ortofloricoltura, 54(2):50-53.

Moreno DS; Fargerlund J; Ewaet WH, 1984. Citrus mealybug Planococcus citri (Homoptera: Pseudococcidae): behaviour of males to sex pheromone in laboratory and field. Annals of the Entomological Society of America, 77(1):32-38.

Moriyama T, 1941. Insects injurious to coffee berries (in Japanese). Formosan Agricultural Review, Taihoku, 37:14-20.

Morse S; Acholo M; McNamara N; Oliver R, 2000. Control of storage insects as a means of limiting yam tuber fungal rots. Journal of Stored Products Research, 36(1):37-45.

Murray DAH, 1976. Insect pests on passion fruit. Queensland Agricultural Journal, 102(2):145-151

Murray DAH, 1978. Effect of fruit fly sprays on the abundance of the citrus mealybug, Planococcus citri (Risso), and its predator, Cryptolpmus montrouzieri Mulsant, on passion-fruit in south-eastern Queensland. Queensland Journal of Agricultural and Animal Sciences, 35(2):143-147

Murray DAH, 1978. Population studies of the citrus mealybug, Planococcus citri (Risso), and its natural enemies on passion-fruit in south-eastern Queensland. Queensland Journal of Agricultural and Animal Sciences, 35(2):139-142

Myburgh AC; Whitehead VB; Daiber CC, 1973. Pests of deciduous fruit, grapes and miscellaneous other horticultural crops in South Africa. Entomology Memoir. Department of Agricultural Technical Services, Republic of South Africa. Pretoria., 27:38 pp.

Myres LE, 1932. Two economic greenhouse mealybugs of Mississippi. Journal of Economic Entomology, 25:891-896.

Nagarkatti S; Singh SP; Jayanth KP; Bhummannavar BS, 1992. Introduction and establishment of Leptomastix dactylopii How. against Planococcus spp. in India. Indian Journal of Plant Protection, 20(1):102-104

Narasimham AU, 1987. Scale insects and mealybugs on coffee, tea and cardamom and their natural enemies. Journal of Coffee Research, 17(1):7-13

Nestel D; Cohen H; Saphir N; Klein M; Mendel Z, 1995. Spatial distribution of scale insects: comparative study using Taylor's power law. Environmental Entomology, 24(3):506-512

Niyazov OD, 1969. The parasites and predators of grape mealybug. Zashchita Rastenii, 14(11):38-40

Niyazov OD, 1976. Revision of the list of parasites of the vine mealybug. In: Tashlieva AO, ed. Ecology and economic importance of the insects of Turkmenia.: Ekologiya i khozyaistvennoe znachenie nasekomykh Turkmenii. Ylim. Ashkhabad USSR, 117-124

Noyes J, 2016. Universal Chalcidoidea database. London, UK: Natural History Museum. http://www.nhm.ac.uk/our-science/data/chalcidoids/database/

Onazi OC, 1969. The infestation of stored potatoes (Solanum tuberosum) by Planococcus citri (Risso)(Homoptera:Pseudococcidae) on the Jos Plateau, Nigeria. Nigerian Entomologists' Magazine, 2(1):17-18

Onorato M; Casu AP; Gerardi M, 1998. The effect of temperature and development on citrus fruits. Informatore Agrario, 54(32):75-77.

Orlinskii AD; Rzpva LM; Shakhramanov IK, 1989. A promising entomophage. Zashchita Rastenii (Moskva), No. 11:25-26

Padi B, 1990. The application of morphometric analyses and gel electrophoresis to the identification of mealybug vectors (Homoptera: Pseudococcidae) of cocoa swollen shoot virus (CSSV) disease, PhD Thesis. Cardiff, UK: University of Wales, College of Cardiff.

Padi B, 1994. Identification of the mealybug vectors of the cocoa swollen shoot virus (CSSV) disease in Ghana by isoenzyme gel electrophoresis. Proceedings of the 11th International Cocoa research Conference 18th-24th July, 1993. Yamoussoukro, Cote d'Ivoire, 971-983.

Padi B, 1997. Morphological variation in cocoa mealybugs. In: Proc. First International Cocoa Pests and Diseases Seminar 6-10 November, 1995: 218-237.

Padi B, 1997. Parthenogenesis in cocoa mealybugs. Proceedings of Cocoa Pests and Diseases Conference, 6th-10th November, 1995, Accra, Ghana. Tafo, Ghana: Cocoa Research Institute of Ghana, in press.

Padi B; Owusu GK; Kumah NK, 1999. A record of Desplatsia dewevrei (De Wild & Th. Dur) (Tiliales: Tiliaceae) as an alternative and potential breeding host plant for the cocoa mirid Sahlbergella singularis Hagl. Proc. 12th Int. Cocoa Research Conf., November 1996, Salvador, Brazil:31-35.

Paik WH, 1972. Scale insects found in the greenhouses in Korea. Korean Journal of Plant Protection, 11(1):1-4.

Panis A, 1977. Pseudococcids (Homoptera, Coccoidea, Pseudococcidae) within the context of integrated control in citrus groves round the Mediterranean. Boletin del Servicio de Defensa contra Plagas e Inspeccion Fitopatologica, 3:139-145

Panis A, 1979. Mealybugs (Homoptera, Coccoidea: Pseudococcidae) within the framework of integrated control in Mediterranean citrus-growing. Revue de Zoologie Agricole et de Pathologie Vegetale, 78(3):88-96

Panis A, 1981. Effect of ants on the parasitic biocoenosis of the black citrus scale in France (Homoptera, Coccoidea, Coccidae). Fruits, 36(1):47-48

Panis A; Brun J, 1971. Biological control tests against three species of Pseudococcidae (Homoptera, Coccoidea) in greenhouses. Revue de Zoologie Agricole et de Pathologie Vegetale, 70(2):42-47

Pedroso EI; Sequeira OA; Pinto MEG; Simoes V, 1991. Assays of transmission of grapevine virus by pseudococcids. Cie^circumflex~ncia e Te^acute~cnica Vitivini^acute~cola, 10(2):39-46; 19 ref.

Pelley RH le, 1968. Pests of Coffee. London and Harlow, UK: Longmans, Green and Co Ltd.

Pena JE; Duncan R, 1990. Role of arthropods in the transmission of postbloom fruit drop. Proceedings of the Florida State Horticultural Society, 102:249-251

Phillips S; Briddon RW; Brunt AA; Hull R, 1999. The partial characterization of a badnavirus infecting the greater asiatic or water yam (Dioscorea alata). Journal of Phytopathology, 147(5):265-269; 30 ref.

Pillai GB, 1987. Integrated pest management in plantation crops. Journal of Coffee Research, 17(1):150-153

Posnette AF, 1941. Swollen shoot virus disease of cocoa. Tropical Agriculture, 19:87-90.

Prakasan CB, 1987. Biological control of coffee pests. Journal of Coffee Research, 17(1):114-117

Pyle KR, 1979. Integrated control of citrus pests in Zimbabwe Rhodesia. Zimbabwe Rhodesia Agricultural Journal, 76(4):171-179

Quillis Perez M, 1935. Calculo de las fajas isocondicionales y de las lineas de Maximo desarrollo para los insectos, VI Congr. int. Madrid, 2:447-454.

Rahiman PA; Vijayalakshmi CK, 1998. Spalgius epius Westwood (Lepidoptera: Lycaenidae) - a potential predator of coffee mealy bugs. Journal of Entomological Research, 22(2):191-192; 3 ref.

Rao KR; Shylesha AN; Pathak KA, 2001. Spatial dynamics of citrus mealybug Planococcus citri Risso at medium high altitudes of Meghalaya. Indian Journal of Hill Farming, 14(2):48-52.

Reddy KB; Bhat PK, 1993. Effect of endosulfan on the mealybug parasitoid, Leptomastix dactylopii How. Journal of Coffee Research, 23(1):19-23

Reddy KB; Bhat PK, 1993. Effect of seasonal augmentation of Leptomastix dactylopii How. on Planococcus citri (Risso) population. Journal of Coffee Research, 23(1):15-18

Reddy KB; Bhat PK; Naidu R, 1997. Suppression of mealybugs and green scale infesting coffee with natural enemies in Karnataka. Pest Management and Economic Zoology, 5(2):119-121; 7 ref.

Reddy KB; Prakasan CB; Bhat PK; Kumar AC, 1992. Establishment of Leptomastix dactylopii How. (Hym.: Encyrtidae) in Karnataka for control of Planococcus citri (Risso) (Hom.: Pseudococcidae) of coffee. Journal of Coffee Research, 22(1):37-44

Reddy KB; Seetharama HG, 1997. Integrated insect pest management for citrus in Northern Mediterranean countries. Indian Coffee, 61(3): 26-28.

Reddy KB; Sreedharan K; Bhat PK, 1991. Effect of rate of prey, Planococcus citri (Risso) on the fecundity of mealybug predator, Cryptolaemus montrouzieri Mulsant. Journal of Coffee Research, 21(2):149-150

Reyd G; Gery R; Ferran A; Iperti G; Brun J, 1991. Voracity of Hyperaspis raynevali (Col.: Coccinellidae) - a predator of cassava mealybug, Phenacoccus manihoti (Hom.: Pseudococcidae). Entomophaga, 36(2):161-171

Rico-Gray V; Thien LB, 1989. Ant-mealybug interaction decreases reproductive fitness of Schomburgkia tibicinis (Orchidaceae) in Mexico. Journal of Tropical Ecology, 5(1):109-112

Ripolles JL; Garcia A, 1989. Quality control of entomophagous insects reared in captivity: possibility of application in the case of Cryptolaemus montrouzieri Mulsant. Boletin de Sanidad Vegetal, Fuera de Serie, No. 17:399-412

Risbec J, 1937. Observations sur les parasites des plantes cultivees aux Nouvelles-Hebrides, Faune Colon. fr. 6 fasc. 1.

Rivero-Lynch AP; Godfray HCJ, 1997. The dynamics of egg production, oviposition and resorption in a parasitoid wasp. Functional Ecology, 11(2):184-188; 14 ref.

Rojter S; Bonney JK; Legg JT, 1966. Investigations into the use of pathogenic fungus (Cephalosporium sp.) as a means of controlling the mealybug (Pseudococcidae) vectors of swollen shoot virus in Ghana. Ghana Journal of Science, 6:110.

Ronse A, 1990. Integrated pest management in the greenhouses of the national botanic garden of Belgium. Revue de l'Agriculture, 43(3):429-436

Rotundo G; Tremblay E, 1974. Studies on a sexual pheromone or Planococcus citri (Risso) (Homoptera, coccoidea). II. Extraction. Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', Portici, 31:121-131

Rotundo G; Tremblay E, 1975. Studies on a sexual pheromone of Planococcus citri (Risso) (Homoptera Coccoidea). III. Specificity. Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', Portici, 32:125-130

Rotundo G; Tremblay E, 1976. Serum-diagnostic discrimination in the Pseudococcidae (Homoptera: Coccoidea). II. Planococcus citri (Ris.) and Planococcus ficus (Sign.). Bollettino del Laboratorio de Entomologia Agraria 'Filippo Silvestri', Portici, 33:99-107

Rotundo G; Tremblay E, 1976. Simple extraction and bioassay of the female sex pheromone of the Citrus mealybug, Planococcus citri. Annals of Applied Biology, 82(1):165-167

Rotundo G; Tremblay E, 1980. Evaluation of the daily rate of sex pheromone release by the females of two mealybug species (Homoptera Coccoidea Pseudococcidae). Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', Portici, 37:167-170

Rotundo G; Tremblay E, 1980. Serological studies on five species of Pseudococcidae (Homoptera). Systematic Entomology, 5(4):431-435

Rung A; Miller DR; Scheffer SJ, 2009. Polymerase chain reaction-restriction fragment length polymorphism method to distinguish three mealybug groups within the Planococcus citri-P. minor species complex (Hemiptera: Coccoidea: Pseudococcidae). Journal of Economic Entomology, 102(1):8-12. http://docserver.ingentaconnect.com/deliver/connect/esa/00220493/v102n1/s3.pdf?expires=1264388788&id=0000&titleid=10264&checksum=A4C453616639D6A29B5093B70281D069

Rung A; Scheffer SJ; Evans G; Miller D, 2008. Molecular identification of two closely related species of mealybugs of the genus Planococcus (Homoptera: Pseudococcidae). Annals of the Entomological Society of America, 101(3):525-532. http://docserver.ingentaconnect.com/deliver/connect/esa/00138746/v101n3/s6.pdf?expires=1221718848&id=0000&titleid=10263&checksum=8F5E910CE2BFF8899EA7BFAFB6E8E1EC

Salazar TJ, 1972. Contribution to knowledge of Pseudococcidae of Peru. Proceedins of the First Latin-American Congress Latinoamericano de Entomologia, Cusco, Peru. Revista Peruana de Entomologia, 15:277-303.

Samal P; Misra BC; Nayak P, 1978. Entomophthora fumosa Speare, an entomogenous fungus on rice brown planthoppers. Current Science, 47(7):241-242

Seljak G, 2010. .

Sengonca C; Kotikal YK; Schade M, 1995. Olfactory reactions of Cryptolaemus montrouzieri Mulsant (Col., Coccinellidae) and Chrysoperla carnea (Stephens) (Neur., Chrysopidae) in relation to period of starvation. Anzeiger fur Schadlingskunde, Pflanzenschutz, Umweltschutz, 68(1):9-12

Silva EB; Mexia A, 1999. The pest complex Cryptoblabes gnidiella (MilliFre) (Lepidoptera: Pyralidae) and Planococcus citri (Risso) (Homoptera: Pseudococcidae) on sweet orange groves (Citrus sinensis (L.) Osbeck) in Portugal: interspecific association. Boleti^acute~n de Sanidad Vegetal, Plagas, 25(1):89-98; 29 ref.

Silva EB; Mexia A; Vacante V, 1997. The damage caused by Planococcus citri (Risso) on citrus groves. Proceedings of the meeting held at Florence, Italy, 29 August 1996. Bulletin-OILB-SROP, 20(7):26-31.

Simmonds MSJ; Manlove JD; Blaney WM; Khambay BPS, 2000. Effect of botanical insecticides on the foraging and feeding behavior of the coccinellid predator Cryptolaemus montrouzieri. Phytoparasitica, 28(2):99-107; 18 ref.

Sirisena UGAI; Watson GW; Hemachandra KS; Wijayagunasekara HNP, 2013. A modified technique for the preparation of specimens of Sternorrhyncha for taxonomic studies. Tropical Agricultural Research Journal, 24(2):139-149.

Smith D; Freebairn CG; Papacek DF, 1996. The effect of host density and parasitoid inoculum size on the mass production of Leptomastix dactylopii Howard (Hymenoptera: Encyrtidae) and Aphytis lingnanensis Compere (Hymenoptera: Aphelinidae) in Queensland. General and Applied Entomology, 27:57-64; 10 ref.

Smith D; Papacek DF; Murray DAH, 1988. The use of Leptomastix dactylopii Howard (Hymenoptera: Encyrtidae) to control Planococcus citri (Risso) (Hemiptera: Pseudococcidae) in Queensland citrus orchards. Queensland Journal of Agricultural and Animal Sciences, 45(2):157-164

Smith FF; Boswell AL, 1972. Gas-propelled aerosols and micronized dusts for control of insects in aircraft. 8. Toxicity of micronized dusts to some greenhouse insects. Journal of Economic Entomology, 65(5):1466-1468

Soylu OZ; Urel N, 1977. Investigations on the parasites and predators of insects injurious to citrus orchards in the region of South Anatolia. Bitki Koruma Bulteni, 17(2/4):77-112

Staal GB; Nassar S; Martin JW, 1973. Control of the citrus mealybug with insect growth regulators with juvenile hormone activity. Journal of Economic Entomology, 66(4):851-853.

Strickland AH, 1947. Coccids attacking cocoa (Theobroma cacao L.) in West Africa, with descriptions of five new species. Bulletin of Entomological Research, 38:497-523.

Strickland AH, 1951. The entomology of swollen shoot of cocoa. I. The insect species involved, with notes on their biology. Bulletin of Entomological Research, 41:725-748.

Strickland AH, 1951. The entomology of swollen shoot of cocoa. II. The bionomics and ecology of the species involved. Bulletin of Entomological Research, 42:64-103.

Su HongJi; Hung TingHsuan; Wu MengLing, 1997. First report of banana streak virus infecting banana cultivars (Musa spp.) in Taiwan. Plant Disease, 81(5):550; 2 ref.

Su TsongHong; Li ChiTsang, 1993. Factors affecting the sex ratio of Leptomastix dactylopii Howard, a parasitoid of Planococcus citri (Risso). Chinese Journal of Entomology, 13(4):319-329; 24 ref.

Su TsongHong; Su ChungKuang; Su TH; Su CK; Vacante V, 1997. Comparison of Anagyrus sawadadii Ishii and Leptomastix Dactylopii Howard (Hymenoptera: Encyrtidae) parasitizing the citrus mealybug, Planococcus citri (Homoptera: Pseudococcidae) in Taiwan. Proceedings of the meeting held at Florence, Italy, 29 August 1996. Bulletin OILB-SROP, 20(7):32-44.

Sutherland JRC, 1953. Some observations on mealybugs infesting cocoa in Western Region, Nigeria. Proceedings of the West African International Cocoa Research Conference, 12-16 December, 1953, Tafo, Gold Coast, 90-94.

Szent-Ivany JJH, 1961. Insect pests of Theobroma cacao in the Territory of Papua New Guinea. Papua New Guinea Agricultural Journal, 13(4):127-147.

Szent-Ivany JJH; Stevens RM, 1966. Insects of Saccharum spp. in the territory of Papua New Guinea and adjacent islands. Pacific Insects, 2:255-261.

Tang SJ; Qin HZ; Wang JM; Gu P, 1992. Studies on the mealy-bug Planococus citri (Risso). Journal of Shanghai Agricultural College, 10(1):44-52; 6 ref.

Tsalev M, 1970. On some mealybug species, Pseudococcus spp., in Bulgaria. Ovoshcharstvo, 17(12):20-24.

Tsay JG, 1991. The occurrence of Pestalotia rot of bagged guava fruits and screening of fungicides for its control in Taiwan. Plant Protection Bulletin (Taipei), 33(4):384-394

Tumminelli R; Saraceno F; Conti D, 1997. Ants in citrus groves. Informatore Agrario, 53(11):57-60.

Ullah GMR; Parveen A, 1993. Coccoid pests (scale insects and mealybugs) and their host-plants on Chittagong University campus - a checklist. Bangladesh Journal of Zoology, 21(1):181-182

Ultee AG, 1924. Verslag over de werkzaamheden van het Proefstation Malang in 1925. Meded. Proefsin. Malang, 57:52.

Uygun N; Sekeroglu E; Karaca I, 1987. Studies on integrated control in a newly established citrus orchard in Cukurova. Turkiye I. Entomoloji Kongresi Bildirileri, 13-16 Ekim 1987, Ege Universitesi, Bornova, Izmir Bornova/Izmir, Turkey; Ege Universitesi Ataturk Kultur Merkezi, 459-469

Viggiani G, 1974. The Coccoids of Citrus in Italy and problems related to the means of combating them. O.I.L.B. 'Coccoids of Citrus' Study Group meeting in Morocco (26-31 October 1970). Awamia, 37:47-55.

Viggiani G, 1975. Integrated control in citrus orchards: experiment on the control of Planococcus citri. Fruits, 30(4):261-265.

Viggiani G, 1975. Method of estimating populations of Planococcus citri at the citrus orchard level. Fruits, 30(3):177-178.

Viggiani G, 1988. The defence of crops by ecological methods. Biological control of insects. Italia Agricola, 125(1):273-288

Viggiani G; Castronuovo N; Borrelli C, 1972. Secondary effects of 40 pesticides on Leptomastidea abnormis - Grlt. - (Hym. Encyrtidae) and Scymnus includens Kirsch (Col. Coccinellidae), important natural enemies of Planococcus citri (Risso). Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri' Portici, 30:88-103

Viggiani G; Tranfaglia A, 1978. A method for laboratory test of side-effects of pesticides on Leptomastix dactylopii (How.) Hym. Encyrtidae). Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', Portici, 35:8-15

Vogel W; Masner P; Frischknecht ML, 1976. Regulation of development and population growth of mealy bugs treated with epofenonane, a JH active IGR. [Frischknecht, M. L.; Muller, P. J. : The use of insect growth regulators in integrated pest control]. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 49(3-4):245-252.

Vogel W; Masner P; Frischknecht ML; Zang H, 1977. Reversal of metamorphosis in mealy bugs treated with juvenile hormone-active insect growth regulator. Experientia, 33(11):1537-1539

Voigt D, 2000. Infestation of Spanish moss (Tillandsia usneoides L.) by the ensign scale Orthezia tillandsiae Morrison (Homoptera, Coccina: Ortheziidae) and possibilities of their biological control at the Botanical Garden of the Technical University of Dresden. Gesunde Pflanzen, 52(5):148-155; 23 ref.

Vos M; Alphen JJMvan; Kole M, 1993. Sex ratios and mating structure in Leptomastix dactylopii, a parasitoid of the citrus mealybug, Planococcus citri. Mededelingen van de Faculteit Landbouwwetenschappen, Universiteit Gent, 58(2B):561-568; 11 ref.

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.

Waterhouse DF, 1998. Biological Control of Insect Pests: South-east Asian Prospects. Canberra, Australia: Australian Centre for International Agricultural Research (ACIAR), 127 pp.

Watson GW; Cox JM, 1990. Identity of the African coffee root mealybug, with descriptions of two new species of Planococcus (Homoptera: Pseudococcidae). Bulletin of Entomological Research, 80(1):99-105

Way MJ; Khoo KC, 1991. Colony dispersion and nesting habits of the ants, Dolichoderus thoracicus and Oecophylla smaragdina (Hymenoptera: Formicidae), in relation to their success as biological control agents on cocoa. Bulletin of Entomological Research, 81(3):341-350.

Weng WS; Huang YQ, 1988. A preliminary study on a predacious insect, Brumoides lineatus (Weise). Insect Knowledge, 25(2):105-108

Williams DJ, 1973. Scale insects (Homoptera: Coccoidea) on Macadamia. Journal of the Australian Entomological Society, 12(2):81-91

Williams DJ, 1982. The distribution of the mealybug genus Planococcus (Hemiptera: Pseudococcidae) in Melanesia, Polynesia and Kiribati. Bulletin of Entomological Research, 72(3):441-455

Williams DJ, 1985. Australian mealybugs. London, UK; British Museum (Natural History), 431 pp.

Williams DJ, 2004. Mealybugs of southern Asia. Kuala Lumpur, Malaysia: Southdene SDN. BHD, 896 pp.

Williams DJ; Granara de Willink MC, 1992. Mealybugs of Central and South America. Wallingford, UK: CAB International.

Williams DJ; Granara de Willink MC, 1992. Mealybugs of Central and South America. Wallingford, UK: CAB International.

Williams DJ; Matile-Ferrero D, 2005. Mealybugs from Zanzibar and Pemba islands with a discussion of a potential invasive species (Hemiptera, Pseudococcidae). Revue Française d'Entomologie, 27(4):145-152.

Williams DJ; Watson GW, 1988. Scale insects of the tropical South Pacific region. Part 2. Mealybugs (Pseudococcidae). Wallingford, Oxon, UK; CAB International, 260 pp.

Wolcott GN, 1951. The insects of Puerto Rico, University of Puerto Rico Journal of Agriculture, 32:1-975. R. Piedras, PR.

Wysoki M, 1977. Insect pests of macadamia in Israel. Phytoparasitica, 5(3):187-188.

Wysoki M, 1977. Insect pests of macadamia in Israel. Phytoparasitica, 5(3):187-188.

Yang JinSong; Sadof CS, 1997. Variation in the life history of the citrus mealybug parasitoid Leptomastix dactylopii (Hymenoptera: Encyrtidae) on three varieties of Coleus blumei. Environmental Entomology, 26(4):978-982; 23 ref.

Yigit A; Canhilal R; Zaman K, 1994. Cold storage of some natural enemies of citrus mealybug, Planococcus citri (Risso) (Homoptera: Pseudococcidae). Turkiye III. Biyolojik Mucadele Kongresi Bildirileeri, 25-28 Ocak 1994, Ege Universitesi Ziraat Fakultesi, Bitki Koruma Bolumu, Izmir Bornova, Turkey; Ege Universitesi Basimevi, 137-146

Yukawa J, 1978. Some changes in generic names of the Japanese Cecidomyiidi (Diptera, Cecidomyiidae, Cecidomyiinp). Applied Entomology and Zoology, 13(3):222-223

Zimmerman EC, 1948. Insects of Hawaii. Vol. 5. Homoptera: Sternorrhyncha. Honolulu, USA: University of Hawaii Press, 464 pp.

Links to Websites

Top of page
WebsiteURLComment
Scalenethttp://scalenet.info

Contributors

Top of page

06/06/16 Updated by:

Gillian Watson, California Department of Food & Agriculture, Sacramento, California, USA

Distribution Maps

Top of page
You can pan and zoom the map
Save map