Invasive Species Compendium

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Datasheet

Ferrisia virgata
(striped mealybug)

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Datasheet

Ferrisia virgata (striped mealybug)

Summary

  • Last modified
  • 20 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Vector of Plant Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Ferrisia virgata
  • Preferred Common Name
  • striped mealybug
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Ferrisia virgata is a highly polyphagous mealybug. It reproduces quite rapidly in tropical conditions, but it tolerates subtropical and to some extent temperate conditions too. It has been reported on host-plants belonging to over 203 gen...

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Pictures

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PictureTitleCaptionCopyright
Adult females oval, greyish-yellow, 4-4.5 mm long, with two longitudinal, submedian, interrupted dark stripes on the dorsum showing through the waxy secretion. The dorsum also bears numerous straight, glassy threads of wax.
TitleAdult F. virgata on rambutan
CaptionAdult females oval, greyish-yellow, 4-4.5 mm long, with two longitudinal, submedian, interrupted dark stripes on the dorsum showing through the waxy secretion. The dorsum also bears numerous straight, glassy threads of wax.
CopyrightCrown Copyright
Adult females oval, greyish-yellow, 4-4.5 mm long, with two longitudinal, submedian, interrupted dark stripes on the dorsum showing through the waxy secretion. The dorsum also bears numerous straight, glassy threads of wax.
Adult F. virgata on rambutanAdult females oval, greyish-yellow, 4-4.5 mm long, with two longitudinal, submedian, interrupted dark stripes on the dorsum showing through the waxy secretion. The dorsum also bears numerous straight, glassy threads of wax.Crown Copyright
Adult female with ovisac.
TitleAdult
CaptionAdult female with ovisac.
Copyright©Georg Goergen/IITA Insect Museum, Cotonou, Benin
Adult female with ovisac.
AdultAdult female with ovisac.©Georg Goergen/IITA Insect Museum, Cotonou, Benin

Identity

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Preferred Scientific Name

  • Ferrisia virgata (Cockerell, 1893)

Preferred Common Name

  • striped mealybug

Other Scientific Names

  • Dactylopius ceriferus Newstead, 1894
  • Dactylopius magnolicida King, 1902
  • Dactylopius segregatus Cockerell, 1893
  • Dactylopius setosus Hempel, 1900
  • Dactylopius talini Green, 1896
  • Dactylopius virgatus Cockerell, 1893
  • Dactylopius virgatus farinosus Cockerell, 1893
  • Dactylopius virgatus humilis Cockerell, 1893
  • Dactylopius virgatus madagascariensis Newstead, 1908
  • Ferrisia neovirgata Khalid and Shafee, 1988
  • Ferrisia virgata (Cockerell) Fullaway, 1923
  • Ferrisiana setosus (Hempel) Ali, 1970
  • Ferrisiana virgata (Cockerell) Takahashi, 1929
  • Heliococcus malvastrus McDaniel, 1962
  • Pseudoccus virgatus farinosus (Cockerell) Cockerell, 1902
  • Pseudococcus bicaudatus Keuchenius, 1915
  • Pseudococcus magnolicida (King) Cockerell, 1902
  • Pseudococcus marchali Vayssière, 1912
  • Pseudococcus segregatus (Cockerell) Fernald, 1903
  • Pseudococcus virgatus (Cockerell) Kircaldy, 1902
  • Pseudococcus virgatus humilis (Cockerell) Fernald, 1903
  • Pseudococcus virgatus madagascariensis (Newstead) Lindinger, 1913

International Common Names

  • English: guava mealybug; spotted mealybug; tailed coffee mealybug; tailed mealybug; white-tailed mealybug
  • Spanish: cochinilla embandada
  • French: cochenille rayée

Local Common Names

  • Egypt: white mealybug
  • Germany: weisse Lamtoro-laus
  • Netherlands: lamtoroluis; witte lamtoro-luis
  • Philippines: grey mealybug
  • South Africa: gestreepte witluis

EPPO code

  • FERRVI (Ferrisia virgata)
  • PSECVI (Pseudococcus virgatus)

Summary of Invasiveness

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Ferrisia virgata is a highly polyphagous mealybug. It reproduces quite rapidly in tropical conditions, but it tolerates subtropical and to some extent temperate conditions too. It has been reported on host-plants belonging to over 203 genera in 77 families, and can damage many crops, particularly tropical fruit, nut and spice crops and field crops like soybean and tomato. It is known to transmit plant badnavirus diseases of cocoa and black pepper. It is of Neotropical origin and spread around the world in only about 10 years after being first described from Jamaica. Its polyphagy has facilitated its spread by human transport of infested plants, and it is now established in all the subtropical and tropical zoogeographic regions. Its small size and cryptic habits make it difficult to detect and identify at plant quarantine inspection. The increase in international trade in fresh plant material in recent years is likely to facilitate its continued spread.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hemiptera
  •                         Suborder: Sternorrhyncha
  •                             Unknown: Coccoidea
  •                                 Family: Pseudococcidae
  •                                     Genus: Ferrisia
  •                                         Species: Ferrisia virgata

Notes on Taxonomy and Nomenclature

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The genus Ferrisia was described to accommodate F. virgata, a widespread tropical species. In 1929, Takahashi claimed (erroneously) that the name Ferrisia was a homonym, and replaced it with Ferrisiana; however, the name Ferrisia has since been recognized as valid.

It has long been known that specimens normally identified as Ferrisia virgata were either biparental or parthenogenetic (Williams, 1996). Nur (1977), using electrophoretic techniques, indicated that there was one uniparental species and at least two biparental species, while Miller and Kostarab (1979) mentioned seven species. The type species of F. virgata is biparental and Williams (1985) illustrated the uniparental form, showing morphological differences between the two forms. The uniparental form was later described as F. consobrina (Williams and Watson, 1988) and subsequently synonymized under F. malvastra by Williams (1996). Descriptions and illustrations of F. virgata prior to 1980 often contain a combination of the features of F. virgata and F. malvastra.

Furthermore, Kaydan and Gullan (2012) revised the genus Ferrisia and recognized 18 species, eight of them new to science.

Early records of F. virgata from all parts of its distribution therefore need to be verified due to confusion with F. malvastra (Ben-Dov, 1994) and the new species described by Kaydan and Gullan (2012).

Description

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In life, adult females are oval, up to 5 mm long, greyish-yellow, with two longitudinal, submedian, interrupted dark stripes on the dorsum showing through the waxy secretion -- hence the common name 'striped mealybug'. The dorsum also bears numerous straight, glassy threads of wax up to 4.0-4.5 mm long. Several members of the genus Ferrisia have this appearance in life, so authoritative identification requires expert study of stained, slide-mounted adult females using the key in Kaydan and Gullan (2012), or nucleotide sequence data.

Slide-mounted adult female Ferrisia species are easy to recognise by the presence of only one pair of cerarii, situated on the anal lobes, and the presence of enlarged tubular ducts, each with the orifice surrounded by a flat, circular, sclerotized area associated with one or more short setae. Kaydan and Gullan (2012) provided a thorough revision of the genus Ferrisia and a morphological key to world species, including detailed morphological description, illustration and discussion of F. virgata. F. virgata is very difficult to separate from some of the other species, particularly F. dasylirii. It has both anterior and posterior pairs of ostioles; ventral oral-collar tubular ducts of at least 2 sizes; smaller ducts present singly or in segmental clusters on body margin, only on last 2–3 abdominal segments; minute discoidal pores in sclerotised area of enlarged dorsal tubular ducts and larger ventral oral-collar tubular ducts rarely if ever touching rim of duct opening (or only very rarely on ventral ducts); discoidal pores associated with sclerotised area around orifices of dorsal enlarged tubular ducts on anterior abdomen normally not touching outer margin of sclerotised area and very rarely projecting from that margin; dorsal enlarged tubular ducts numbering 69-101; abdominal segment VI with 11-28 multilocular disc pores, usually with more than 15 in a double row; each anal lobe cerarius with 3 (occasionally 2) enlarged conical setae; and hind coxa with translucent pores (Kaydan and Gullan, 2012).

The adult male has long antennae, six well developed legs, one pair of simple wings, no mouthparts, and a pair of long white wax filaments at the posterior end. The morphology of Ferrisia males has not been studied in detail so the species cannot be identified using male morphology.

Distribution

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The genus Ferrisia is apparently of New World origin (Williams, 1996). F. virgata is by far the most widespread species in the genus (Kaydan and Gullan, 2012). Its area of origin is not known; it was described from Jamaica but it was a pest there at the time, so it may have been an introduction from elsewhere (Kaydan and Gullan, 2012). According to the literature, it has spread to all zoogeographical regions, mainly in the tropics, but often extends well into the temperate regions -- see Commonwealth Institute of Entomology (1966) and García et al. (2016). It is likely that the most northern populations are found only in greenhouses. Early records of F. virgata from all parts of its distribution need to be verified due to confusion with F. malvastra (Ben-Dov, 1994) and the new species described by Kaydan and Gullan (2012).

Distribution Table

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

Last updated: 25 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AngolaPresent
BeninPresent
Cabo VerdePresent
CameroonPresent
ComorosPresent
Congo, Democratic Republic of thePresent, Widespread
Congo, Republic of thePresent
Côte d'IvoirePresent
EgyptPresent
EritreaPresent
EthiopiaPresent
GhanaPresent
GuineaPresent
Guinea-BissauPresent
KenyaPresent
MadagascarPresent
MalawiPresent
MaliPresent
MauritiusPresent
-RodriguesPresent
MozambiquePresent
NigeriaPresent
RéunionPresent
São Tomé and PríncipePresent
SenegalPresent
SeychellesPresent
-Aldabra IslandsPresent
Sierra LeonePresent
SomaliaPresent
South AfricaPresent
SudanPresent
TanzaniaPresent
-Zanzibar IslandPresent
TogoPresent
TunisiaPresent2016InvasiveKhezama City; Original citation: Ben Halima et al. (2018)
UgandaPresent
ZambiaPresent
ZimbabwePresent

Asia

BangladeshPresent
British Indian Ocean TerritoryPresent
BruneiPresent
CambodiaPresent
ChinaPresent
-GuangdongPresent
-HunanPresent
Cocos IslandsPresent
Hong KongPresent
IndiaPresent
-Andaman and Nicobar IslandsPresent
-Andhra PradeshPresent
-AssamPresent
-BiharPresent
-GoaPresent
-GujaratPresent
-KarnatakaPresent
-KeralaPresent
-Madhya PradeshPresent
-MaharashtraPresent
-OdishaPresent
-PunjabPresent
-RajasthanPresent
-Tamil NaduPresent
-TripuraPresent
-Uttar PradeshPresent
-West BengalPresent
IndonesiaPresent
-Irian JayaPresent
-JavaPresent
-Lesser Sunda IslandsPresent
-SulawesiPresent
-SumatraPresent
IranPresent
IsraelPresent
JapanPresent
LaosPresent
MalaysiaPresent
-Peninsular MalaysiaPresent
-SabahPresent
-SarawakPresent
MaldivesPresent
MyanmarPresent
PakistanPresent
PhilippinesPresent
Saudi ArabiaPresent
SingaporePresent
Sri LankaPresent
TaiwanPresent
ThailandPresent
United Arab EmiratesPresent
VietnamPresent
YemenPresent

Europe

FrancePresent

North America

Antigua and BarbudaPresentBarbuda; Original citation: Williams and Granara de Willink (1992)
BahamasPresent
BarbadosPresent
BelizePresent
BermudaPresent
Cayman IslandsPresent
Costa RicaPresent
CubaPresent
DominicaPresent
GuadeloupePresentOriginal citation: Matile-Ferrero and Étienne (2006)
GuatemalaPresent
HaitiPresent
HondurasPresent
JamaicaPresent
MartiniquePresent
MexicoPresent
NicaraguaPresent
PanamaPresent
Puerto RicoPresent
Saint Kitts and NevisPresentIn both Saint Kitts and Nevis; Original citation: Williams and Granara de Willink (1992)
Trinidad and TobagoPresentOn both islands: Trinidad and Tobago
U.S. Virgin IslandsPresent
United StatesPresentPresent based on regional distribution.
-AlabamaPresent
-CaliforniaPresent
-District of ColumbiaPresent
-FloridaPresent
-HawaiiPresent
-LouisianaPresent
-MarylandPresent
-MassachusettsPresent
-MississippiPresent
-New JerseyPresent
-New MexicoPresent
-New YorkPresent
-OhioPresent
-PennsylvaniaPresent
-TexasPresent
-VirginiaPresent

Oceania

AustraliaPresent
-Northern TerritoryPresent
-QueenslandPresent
Christmas IslandPresent
Cook IslandsPresent
Federated States of MicronesiaPresentPohnpei, Truk Islands and Yap
FijiPresent
French PolynesiaPresent
KiribatiPresent
Marshall IslandsPresent
New CaledoniaPresent
Northern Mariana IslandsPresent
PalauPresent
Papua New GuineaPresent
SamoaPresent
Solomon IslandsPresent
TongaPresent
TuvaluPresent
United States Minor Outlying Islands
-Wake IslandPresent
VanuatuPresent
Wallis and FutunaPresent

South America

ArgentinaPresent
BoliviaPresent
BrazilPresent
-AmazonasPresent
-Espirito SantoPresent
-ParaibaPresent
-PernambucoPresent
-Rio de JaneiroPresent
-Rio Grande do NortePresent
-Sao PauloPresent
ColombiaPresent
EcuadorPresent
-Galapagos IslandsPresent
GuyanaPresent
ParaguayPresent
PeruPresent
SurinamePresent
VenezuelaPresent

History of Introduction and Spread

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F. virgata was first described from Jamaica (Cockerell, 1893). Within a short period, it had been described from India (by Newstead, 1894), Sri Lanka (by Green, 1896), Madagascar (by Newstead, 1908), West Africa (by Vayssière, 1912) and Indonesia (Java) (by Keuchenius, 1915). Although its morphological affinities indicate that it is of New World origin, in Jamaica it was infesting many plant species in a manner characteristic of a new introduction (Liebhold and Tobin, 2008), so its locality of origin is not known exactly (Kaydan and Gullan, 2012). The speed with which it spread around the world is typical of newly invading insect species (Liebhold and Tobin, 2008). Transport of infested plant material is presumed to have been the main mode of spread.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
India Western Hemisphere early 1890s Crop production (pathway cause); Horticulture (pathway cause) Yes No Newstead (1894)
Jamaica Western Hemisphere early 1890s Crop production (pathway cause); Horticulture (pathway cause) Yes No Cockerell (1893)
Java 1910-1914 Crop production (pathway cause); Horticulture (pathway cause) Yes No Keuchenius (1915)
Madagascar early 1900s Crop production (pathway cause); Horticulture (pathway cause) Yes No Newstead (1908)
Sri Lanka early 1890s Crop production (pathway cause); Horticulture (pathway cause) Yes No Green (1896)
West Africa early 1900s Crop production (pathway cause); Horticulture (pathway cause) Yes No Vayssière (1912)

Risk of Introduction

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All life stages may be carried on consignments of fresh plant material and fruit. F. virgata is often injurious to crop and ornamental plants especially when it is introduced to new geographical areas without its natural enemies (Williams, 1996).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Principal habitat Harmful (pest or invasive)
Terrestrial ManagedCultivated / agricultural land Principal habitat Natural
Terrestrial ManagedProtected agriculture (e.g. glasshouse production) Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Principal habitat Natural
Terrestrial ManagedDisturbed areas Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Secondary/tolerated habitat Natural
Terrestrial ManagedRail / roadsides Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial ManagedRail / roadsides Secondary/tolerated habitat Natural
Terrestrial ManagedUrban / peri-urban areas Principal habitat Harmful (pest or invasive)
Terrestrial ManagedUrban / peri-urban areas Principal habitat Natural
Terrestrial Natural / Semi-naturalNatural forests Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Secondary/tolerated habitat Natural
Terrestrial Natural / Semi-naturalRiverbanks Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalRiverbanks Secondary/tolerated habitat Natural
LittoralCoastal areas Secondary/tolerated habitat Harmful (pest or invasive)
LittoralCoastal areas Secondary/tolerated habitat Natural
LittoralCoastal dunes Secondary/tolerated habitat Harmful (pest or invasive)
LittoralCoastal dunes Secondary/tolerated habitat Natural
LittoralMangroves Secondary/tolerated habitat Harmful (pest or invasive)
LittoralMangroves Secondary/tolerated habitat Natural
FreshwaterIrrigation channels Secondary/tolerated habitat Harmful (pest or invasive)
FreshwaterIrrigation channels Secondary/tolerated habitat Natural

Hosts/Species Affected

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F. virgata is one of the most highly polyphagous mealybugs known, attacking plant species belonging to some 203 genera in 77 families (García et al., 2016). Many of the host species belong to the Fabaceae and Euphorbiaceae. Among the hosts of economic importance are avocado, banana, betel vine, black pepper, cassava, cashew, cauliflower, citrus, cocoa, coffee, cotton, custard apple, aunergine, grapevine, guava, jute, lantana, Leucaena, litchi, mango, oil palm, pigeon pea, pineapple, soyabean and tomato. Acalypha species are apparently a favoured host-plant in many places (Kaydan and Gullan, 2012).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Abelmoschus esculentus (okra)MalvaceaeMain
    Acalypha (Copperleaf)EuphorbiaceaeMain
      Albizia lebbeck (Indian siris)FabaceaeMain
        Anacardium occidentale (cashew nut)AnacardiaceaeMain
          Ananas comosus (pineapple)BromeliaceaeMain
            AnnonaAnnonaceaeMain
              Annona reticulata (bullock's heart)AnnonaceaeOther
                Annona squamosa (sugar apple)AnnonaceaeUnknown
                  Arachis hypogaea (groundnut)FabaceaeOther
                    Brassica oleracea (cabbages, cauliflowers)BrassicaceaeUnknown
                      Cajanus cajan (pigeon pea)FabaceaeMain
                        Carica papaya (pawpaw)CaricaceaeMain
                          CitrusRutaceaeMain
                            Coccoloba uvifera (sea grape)PolygonaceaeMain
                              Cocos nucifera (coconut)ArecaceaeMain
                                Codiaeum variegatum (garden croton)EuphorbiaceaeMain
                                  Coffea (coffee)RubiaceaeMain
                                    Colocasia esculenta (taro)AraceaeMain
                                      Corchorus (jutes)TiliaceaeMain
                                        Cucurbita maxima (giant pumpkin)CucurbitaceaeMain
                                          Cucurbita pepo (marrow)CucurbitaceaeMain
                                            Dioscorea esculenta (Asiatic yam)DioscoreaceaeUnknown
                                              DracaenaAgavaceaeMain
                                                Elaeis guineensis (African oil palm)ArecaceaeMain
                                                  FicusMoraceaeMain
                                                    Glycine max (soyabean)FabaceaeUnknown
                                                      Gossypium (cotton)MalvaceaeMain
                                                        Hevea brasiliensis (rubber)EuphorbiaceaeUnknown
                                                          Hibiscus (rosemallows)MalvaceaeOther
                                                            Ipomoea batatas (sweet potato)ConvolvulaceaeMain
                                                              Lantana camara (lantana)VerbenaceaeOther
                                                                Leucaena leucocephala (leucaena)FabaceaeMain
                                                                  Litchi chinensis (lichi)SapindaceaeMain
                                                                    Malpighia glabra (acerola)MalpighiaceaeOther
                                                                      Mangifera indica (mango)AnacardiaceaeMain
                                                                        Manihot esculenta (cassava)EuphorbiaceaeMain
                                                                          ManilkaraSapotaceaeMain
                                                                            Musa (banana)MusaceaeMain
                                                                              Nephelium lappaceum (rambutan)SapindaceaeOther
                                                                                Nicotiana tabacum (tobacco)SolanaceaeMain
                                                                                  Persea americana (avocado)LauraceaeOther
                                                                                    Phaseolus (beans)FabaceaeMain
                                                                                      Phoenix dactylifera (date-palm)ArecaceaeMain
                                                                                        Phyllanthus acidus (star gooseberry)EuphorbiaceaeOther
                                                                                          Piper betle (betel pepper)PiperaceaeMain
                                                                                            Piper nigrum (black pepper)PiperaceaeMain
                                                                                              Pongamia pinnata (Indian beech)FabaceaeOther
                                                                                                Psidium guajava (guava)MyrtaceaeMain
                                                                                                  Punica granatum (pomegranate)PunicaceaeMain
                                                                                                    Saccharum officinarum (sugarcane)PoaceaeOther
                                                                                                      Solanum lycopersicum (tomato)SolanaceaeMain
                                                                                                        Solanum melongena (aubergine)SolanaceaeMain
                                                                                                          Solanum nigrum (black nightshade)SolanaceaeMain
                                                                                                            Theobroma cacao (cocoa)MalvaceaeMain
                                                                                                              Vigna unguiculata (cowpea)FabaceaeMain
                                                                                                                Vitis vinifera (grapevine)VitaceaeMain
                                                                                                                  Zea mays (maize)PoaceaeOther
                                                                                                                    Zingiber officinale (ginger)ZingiberaceaeMain

                                                                                                                      Growth Stages

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                                                                                                                      Flowering stage, Fruiting stage, Post-harvest, Vegetative growing stage

                                                                                                                      Symptoms

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                                                                                                                      Infestations of F. virgata remain clustered around the terminal shoots, leaves and fruit, sucking plant sap which results in yellowing, withering and drying of plants and premature shedding of leaves and fruit. The mealybugs do not feed on phloem very often, so unlike many mealybug species they do not produce huge quantities of sugary honeydew. What honeydew is produced can foul foliage and fruit and serve as a medium for the growth of black sooty moulds. Sooty moulds and wax deposits can block light and air from the plant, sometimes reducing photosynthesis and hence plant vigour and crop yield.

                                                                                                                      List of Symptoms/Signs

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                                                                                                                      SignLife StagesType
                                                                                                                      Fruit / discoloration
                                                                                                                      Fruit / external feeding
                                                                                                                      Fruit / honeydew or sooty mould
                                                                                                                      Growing point / dead heart
                                                                                                                      Growing point / external feeding
                                                                                                                      Leaves / abnormal colours
                                                                                                                      Leaves / honeydew or sooty mould
                                                                                                                      Leaves / wilting
                                                                                                                      Stems / discoloration
                                                                                                                      Stems / external feeding
                                                                                                                      Stems / honeydew or sooty mould

                                                                                                                      Biology and Ecology

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                                                                                                                      There are several papers on the biology of F. virgata but these need to be verified due to the confusion with F. malvastra, particularly in India where both species occur. F. virgata is biparental and F. malvastra is parthenogenetic.

                                                                                                                      Reproductive Biology

                                                                                                                      F. virgata reproduces sexually, with each female mating only once.  At 30-35 ºC and 65% relative humidity (RH), a complete life cycle required about 6-7 weeks, and at 16.6 ºC, 13-14 weeks (Gerson, 2016). When reared at 16-29 ºC and 54-71% RH on sprouting potatoes, there were three (Ammar et al., 1979) to five overlapping generations per year (Nayar et al., 1976; Awadallah et al., 1979), with the population increasing with each successive generation in the field (Ammar et al., 1979). The same study found that the production of males was highest in crowded conditions. In the field in Egypt, the preferred oviposition sites were on the lower leaf surfaces and at the junction between stem and leaf petiole on Acalypha macrophylla (Awadallah et al., 1979). Eggs are laid in groups beneath the body on a pad of cottony wax filaments (Kaydan and Gullan, 2012) over a period of 20-29 days (Schmutterer, 1969). Each female is ovoviviparous and, in the laboratory, produces from 64 eggs (Awadallah et al., 1979) up to 737 eggs that hatch within 30 minutes of being laid (Ghose and Paul, 1972) or after 3-4 hours (Schmutterer, 1969). Female and male nymphs moulted 3 and 4 times, respectively, and the development period varied from 26-47 and 31-57 days, respectively (Ghose and Paul, 1972).

                                                                                                                      Longevity

                                                                                                                      Longevity of the adult female was 36-53 days, and that of the male 1-3 days (Ghose and Paul, 1972).

                                                                                                                      Activity patterns:

                                                                                                                      As with all mealybugs, only the short-lived males can fly.

                                                                                                                      In Egypt in the field, F. virgata overwintered (probably as adult females) in cracks and junctions of trunks and larger branches and on fallen leaves. In the laboratory, females migrated to the soil in winter (Ammar et al., 1979).

                                                                                                                      Nutrition

                                                                                                                      F. virgata feeds on the sap of a wide variety of host plants.

                                                                                                                      Environmental requirements:

                                                                                                                      F. virgata is found predominantly in the tropics, but its range does extend well into some temperate regions; it is found as far north as New York state in the USA (Ben-Dov., 1994).

                                                                                                                      A significant positive correlation was found (in Egypt) between population density and daily maximum and minimum temperatures, but not between population density and relative humidity (Ammar et al., 1979).

                                                                                                                      Climate

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                                                                                                                      ClimateStatusDescriptionRemark
                                                                                                                      Af - Tropical rainforest climate Preferred > 60mm precipitation per month
                                                                                                                      Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
                                                                                                                      As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (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])
                                                                                                                      BW - Desert climate Tolerated < 430mm 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 Tolerated 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

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                                                                                                                      Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
                                                                                                                      43 27

                                                                                                                      Natural enemies

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                                                                                                                      Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                                                                                                      Acerophagus texanus Parasite California most flowering plants
                                                                                                                      Aenasius advena Parasite Adults/Nymphs California; Hawaii most flowering plants
                                                                                                                      Alloagrapta javana Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Alloagrapta obliqua Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Anagyrus brevicornis Parasite Adults/Nymphs
                                                                                                                      Anagyrus qadrii Parasite Adults/Nymphs
                                                                                                                      Anaysis alcocki Parasite Adults/Nymphs
                                                                                                                      Anusioptera aureocincta Parasite Adults/Nymphs
                                                                                                                      Azya luteipes Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Blepyrus insularis Parasite Nymphs
                                                                                                                      Brumoides suturalis Predator Adults/Nymphs
                                                                                                                      Cephaleta brunniventris Parasite
                                                                                                                      Cheilomenes sexmaculata Predator Adults/Nymphs
                                                                                                                      Chrysopa flaveola Predator Adults/Nymphs
                                                                                                                      Chrysopa orestes Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Chrysoperla carnea Predator Adults/Nymphs
                                                                                                                      Coccinella septempunctata Predator Adults/Nymphs
                                                                                                                      Coelinius Parasite Larvae
                                                                                                                      Cryptolaemus montrouzieri Predator Adults/Eggs/Larvae/Nymphs/Pupae Indonesia, India coffee, guava
                                                                                                                      Eublemma costimacula Predator Adults/Nymphs
                                                                                                                      Exochomus flaviventris Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Geocoris tricolor Predator Adults/Nymphs
                                                                                                                      Gyranusoidea citrina Parasite Nymphs
                                                                                                                      Hyperaspis notata Predator Adults/Nymphs
                                                                                                                      Hyperaspis senegalensis hottentotta Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Mallada boninensis Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Myiopharus doryphorae Parasite Adults/Nymphs
                                                                                                                      Neozygites fresenii Pathogen Adults/Nymphs
                                                                                                                      Nephus regularis Predator Adults/Nymphs
                                                                                                                      Ocyptamus argentinus Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Ocyptamus mentor Predator Adults/Nymphs
                                                                                                                      Odontochrysa lacciperda Predator Adults/Nymphs
                                                                                                                      Olla v-nigrum Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Patiyana coccorum Parasite Adults/Nymphs
                                                                                                                      Pseudaphycus debachi Parasite Adults/Nymphs
                                                                                                                      Scymnus apiciflavus Predator Adults/Nymphs
                                                                                                                      Scymnus castaneus Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Scymnus coccivora Predator Adults/Eggs/Larvae/Nymphs/Pupae
                                                                                                                      Scymnus roepkei Predator Adults/Nymphs
                                                                                                                      Spalgis epeus Predator Adults/Nymphs
                                                                                                                      Triommata coccidivora Predator Adults/Nymphs

                                                                                                                      Notes on Natural Enemies

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                                                                                                                      Bartlett (1978) gave an account of the introduced parasitoids and predators used to control F. virgata. Noyes (2016) provides a list of known hymenopteran parasitoids of F. virgata.

                                                                                                                      Means of Movement and Dispersal

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                                                                                                                      Natural Dispersal

                                                                                                                      As for most mealybugs, local dispersal of F. virgata is mainly by the crawling of the first instars. However, all the female developmental stages can walk, at least until eggs are laid (Kaydan and Gullan, 2012), and they will move to avoid unfavourable conditions. Additionally, wind may pick up and carry crawlers longer distances (Kaydan and Gullan, 2012).

                                                                                                                      Vector Transmission

                                                                                                                      In unfavourable conditions, attendant ants sometimes pick up mealybugs and carry them to new feeding sites (Missouri Botanical Garden, 2016). Sometimes mealybug crawlers walk onto the feet of birds perching on infested trees, and get carried to new plants.  

                                                                                                                      Accidental Introduction

                                                                                                                      All life stages of F. virgata may be carried in shipments of fresh plant material and fruit (Kaydan and Gullan, 2012); as for other mealybug species, human transport of infested plants is a common way in which F. virgata gets transported over long distances, in-country or internationally, and presumably for short distances as well.  This includes trade in nursery stock, sharing or trade of infested material by plant fanciers, import of unusual plants to botanical gardens and food plant material to zoos, and the 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.

                                                                                                                      Humans and farm machinery working in infested fields can accidentally carry crawlers to other sites.

                                                                                                                      Pathway Causes

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                                                                                                                      CauseNotesLong DistanceLocalReferences
                                                                                                                      Botanical gardens and zoosAccidental introduction on plants Yes
                                                                                                                      Crop productionAccidental introduction on plants; transport on used farm machinery Yes Yes
                                                                                                                      Cut flower tradeAccidental introduction on plants Yes Yes
                                                                                                                      HitchhikerAccidental introduction on plants, and also on machinery, tools and workers Yes Yes
                                                                                                                      HorticultureAccidental introduction on plants, and also on machinery, tools and workers Yes Yes
                                                                                                                      Landscape improvementAccidental introduction on plants Yes Yes
                                                                                                                      Nursery tradeAccidental introduction on plants, and also on machinery, tools and workers Yes Yes
                                                                                                                      People sharing resourcesPlant fanciers sharing plant material Yes Yes
                                                                                                                      Self-propelled Yes

                                                                                                                      Pathway Vectors

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                                                                                                                      VectorNotesLong DistanceLocalReferences
                                                                                                                      ConsumablesAccidental introduction on plant material Yes
                                                                                                                      Host and vector organismsAccidental introduction on plant material Yes Yes
                                                                                                                      Plants or parts of plantsAccidental introduction on plant material Yes Yes
                                                                                                                      Wind Yes

                                                                                                                      Plant Trade

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                                                                                                                      Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
                                                                                                                      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
                                                                                                                      Leaves 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

                                                                                                                      Impact Summary

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                                                                                                                      CategoryImpact
                                                                                                                      Economic/livelihood Negative

                                                                                                                      Economic Impact

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                                                                                                                      Pelley (1968) gave a general discussion of the pest status of F. virgata on coffee. Keuchenius (1915) and Ultee (1924) discussed it forming damaging mixed populations with Planococcus citri (Risso) on coffee in Java, Indonesia. Schmutterer (1969) stated it was a major pest of irrigated guava trees in the drier areas of the Sudan where it is common on many other crop, shade, ornamental and wild plants. In Tanzania it is a pest of cashew and in some parts of the world it is a pest of cotton (Williams, 1996). In India it has been widely reported as a pest of ornamental plants (Ghose and Paul, 1972) and a range of crops including coffee (Chacko and Bhat, 1976), custard apple (Mani and Krishnamoorthy, 1989), betel vine (Patil et al., 1987), black pepper (Sarma et al., 1987), pigeon pea (Gautam and Saxena, 1986) and milk tree (Manilkara hexandra), a rootstock for sapodilla (Manilkara achras) (Jhala et al., 1988). It is also recorded as a pest of kenaf (Hibiscus cannabinus) and mesta (H. sabdariffa) in Bangladesh (Jalil and Kabir, 1971), of Leucaena leucocephala in Taiwan (Chang and Sun, 1985), and of glasshouse ornamental plants in Egypt (Nada, 1986).

                                                                                                                      As well as direct damage caused by feeding, F. virgata is a known vector of Cacao swollen shoot virus (CSSV) in West Africa, cocoa Trinidad virus (CTV, Diego Martin valley isolate) in Trinidad (Thorold, 1975), and a badnavirus disease of black pepper in India (Bhat et al., 2003). Sooty mould fouling of ornamental plants and produce reduces their market value or can make them unmarketable.

                                                                                                                      Threatened Species

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                                                                                                                      Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
                                                                                                                      Serianthes nelsoniiCR (IUCN red list: Critically endangered); USA ESA listing as endangered speciesGuam; Northern Mariana IslandsPest and disease transmissionUS Fish and Wildlife Service (1994)

                                                                                                                      Risk and Impact Factors

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                                                                                                                      Invasiveness
                                                                                                                      • Invasive in its native range
                                                                                                                      • Proved invasive outside its native range
                                                                                                                      • Has a broad native range
                                                                                                                      • Highly adaptable to different environments
                                                                                                                      • Tolerant of shade
                                                                                                                      • Capable of securing and ingesting a wide range of food
                                                                                                                      • Benefits from human association (i.e. it is a human commensal)
                                                                                                                      • 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
                                                                                                                      Impact mechanisms
                                                                                                                      • Pest and disease transmission
                                                                                                                      • Fouling
                                                                                                                      • Herbivory/grazing/browsing
                                                                                                                      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

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                                                                                                                      Heavy infestations are conspicuous because of the white waxy secretions, white masses of male tests (waxy filamentous cocoons) and sooty moulds growing on the excreted honeydew. Colonies often occur at the growing points, around the stem nodes, on the undersides of leaves and on the fruit.

                                                                                                                      Similarities to Other Species/Conditions

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                                                                                                                      F. virgata should be distinguished from F. malvastra, recorded from many parts of the world, and F. dasylirii in the neotropics (García et al., 2016), which may be more widespread than is presently known. These species are morphologically very similar and cannot be distinguished in the field by simple superficial features. Slide-mounted preparations are needed for examination. Other species closely related to F. virgata have been described in recent years from South America and should be separated using the key given by Kaydan and Gullan (2012). Kaydan and Gullan (2012) also provide detailed morphological descriptions, illustrations and discussion of F. virgata, F. dasylirii and F. malvastra; information from their description of F. virgata is provided in the 'Description' section of this datasheet.

                                                                                                                      Prevention and Control

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

                                                                                                                      Cultural Control and Sanitary Methods

                                                                                                                      At the beginning of a local outbreak, severely infested branches should be cut and burnt immediately (Schmutterer, 1969). Mealybugs can walk from one canopy to another when they are touching, so pruning can prevent spread of an infestation.

                                                                                                                      Biological Control

                                                                                                                      Noyes (2016) provides a list of known hymenopteran parasitoids of F. virgata. In Egypt, three species of Coccinellidae, two species of Neuroptera and three parasitoid wasp species (Leptomastix sp. and two species of Tetrastichus) were reported attacking F. virgata (Awadallah et al., 1979). In India,  a coccinellid predator (Scymnus sp.) has been reported attacking F. virgata, controlling a population kept in culture (Ghose and Paul, 1972); also, the predatory caterpillar of Spalgis epius has been recorded feeding on the mealybugs (Chako and Bhat, 1976).

                                                                                                                      According to Zimmerman (1948), F. virgata was first recorded in the Hawaiian Islands in 1898 but was a widespread and common pest in the islands long before this. It is no longer common there as it has been controlled by the coccinellids Cryptolaemus montrouzieri, Olla v-nigrum and Azya luteipes, together with the syrphid Alloagrapta obliqua.

                                                                                                                      Pesticide-resistant populations of F. virgata were effectively controlled in guava orchards in India by the release of C. montrouzieri. The local natural enemies Aenasius advena (an encyrtid parasitoid) and Scymnus coccivora (a cocinellid predator) also helped control the mealybugs (Mani et al., 1990).

                                                                                                                      Bartlett (1978) gave an account of the introduced parasitoids and predators used to control F. virgata.

                                                                                                                      Chemical Control

                                                                                                                      Diazinon, malathion and dimethoate are effective in controlling F. virgata; however, these chemicals have to be sprayed repeatedly to achieve satisfactory control (Schmutterer, 1969). Prothiofos, either alone or with mineral oil, gave better control of F. virgata on guava in South Africa than did malathion, and resulted in negligible residues (Villiers and Stander, 1978).

                                                                                                                      In Florida, F. virgata on Caladium foliage was controlled by four sprays of permethrin at weekly intervals. Diflubenzuron reduced populations by about half. Permethrin provided the most effective control 21 days after dip treatments of tubers, and bendiocarb dip treatment was also very effective (Price, 1979).

                                                                                                                      Gaps in Knowledge/Research Needs

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                                                                                                                      There is a lack of documentation of the environmental requirements/limitations for F. virgata.  It would be helpful to have a CLIMEX analysis done to assess how much further it is likely to spread, and which areas are at risk of invasion.

                                                                                                                      References

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                                                                                                                      Price JF, 1979. Control of mealybugs on caladiums. Proceedings of the Florida State Horticultural Society, 92:358-360

                                                                                                                      Rosen D, 1981. A new species of Pseudaphycus (Hym.: Encyrtidae), with notes on the angelicus group. Entomophaga, 26(3):251-263

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                                                                                                                      Trjapitzin SV, Trjapitzin VA, 1999. Parasitoids of the mealybugs on cultivated grapes in Argentina, with description of a new species of the genus Aenasius Walker (Hymenoptera, Encyrtidae). Entomologicheskoe Obozrenye, 76:174-179

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

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                                                                                                                      Villatoro-Moreno, H., Cisneros, J., Gómez, J., Infante, F., Castillo, A., 2016. Mealybugs (Hemiptera: Pseudococcidae) associated with rambutan (Nephelium lappaceum L.) in Chiapas, Mexico. Journal of the Kansas Entomological Society, 89(4), 289-296. http://www.bioone.org/loi/kent doi: 10.2317/0022-8567-89.4.289

                                                                                                                      Villiers EA de, Stander GN, 1978. Control of the striped mealybug Ferrisia virgata on guavas. Citrus and Subtropical Fruit Journal, 541:16-17

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

                                                                                                                      Watson GW, Ooi PAC, Girling DJ, 1995. Insects on plants in the Maldives and their management. Ascot, UK: International Institute of Biological Control (IIBC), 124 pp

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

                                                                                                                      Williams DJ, 1996. A synoptic account of the mealybug genus Ferrisia Fullaway (Hem., Pseudococcidae). Entomologist's Monthly Magazine, 132(3):1-10; many ref

                                                                                                                      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, 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

                                                                                                                      Wong CY, Chen SP, Chou LY, 1999. Guidebook to Scale Insects of Taiwan. Wufeng, Taichung, Taiwan: Taiwan Agricultural Research Institute, 98 pp

                                                                                                                      Zimmerman EC, 1948. Insects of Hawaii. A Manual of the Insects of the Hawaiian Islands, Including an Enumeration of the Species and Notes on their Origin, Distribution, Hosts, Parasites, etc. Volume 5. Homoptera: Sternorhyncha. Honolulu, Univ. Hawaii Press, vii, [1+] 464 pp. https://scholarspace.manoa.hawaii.edu/handle/10125/2643

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                                                                                                                      Kondo T, 2008. The scale insects of soursop: Annona muricata L. (Las escamas de la guanabana: Annona muricata L). In: Novedades Tecnicas, Revista Regional, Corpoica, Centro de Investigacion, 9 (10) Palmira, Colombia: 25-29.

                                                                                                                      Kosztarab M, 1996. Scale insects of northeastern North America: identification, biology, and distribution. Martinsville, USA: Virginia Museum of Natural History. vii + 650 pp.

                                                                                                                      Lad S K, Patil P D, Godase S K, 2013. Record of mealy bugs infesting fruit crops in Konkan Region of Maharashtra. Journal of Applied Zoological Researches. 24 (2), 141-145. http://www.azra-india.com/journal.html

                                                                                                                      Lincango P, Hodgson CJ, Causton C, Miller DR, 2010. An updated checklist of scale insects (Hemiptera: Coccoidea) of the Galapagos Islands, Ecuador. In: Galapagos Research, 67 3-7.

                                                                                                                      Lit IL, Calilung VJ, 1994. An annotated list of mealybugs (Pseudococcidae, Coccoidea, Hemiptera) from Mount Makiling and vicinity, Laguna, Philippines. In: Philippine Entomologist, 9 385-398.

                                                                                                                      Mangala N, Sundararaj R, Nagaveni H C, 2012. Scales and mealybugs (Coccoidea: Hemiptera) infesting Pongamia pinnata (L.) Pierre and their population dynamics in Karnataka, India. Annals of Forestry. 20 (1), 110-116.

                                                                                                                      Matile-Ferrero D, 1978. Coccoidea Homoptera of the Comoro archipelago. (Homoptères Coccoidea de l'Archipel des Comores. Faune Entomologique de l'Archipel des Comores). In: Mémoires du Museum National d'Histoire Naturelle (N.S.) Serie A, Zoologie, 109 39-70.

                                                                                                                      Moghadam M, 2004. New record on three mealy bugs (three species and two genera) (Hem.; Cocciodea: Pseudococcidae) in Iran. Applied Entomology and Phytopathology. 72 (1), Pe142, en23.

                                                                                                                      Muniappan R, Watson G W, Vaughan L, Gilbertson R, Noussourou M, 2012. New records of mealybugs, scale insects, and whiteflies (Hemiptera: Sternorrhyncha) from Mali and Senegal. Journal of Agricultural and Urban Entomology. 28 (1), 1-7. http://scentsoc.org/Volumes/JAUE/28/28001.pdf DOI:10.3954/1523-5475-28.1.1

                                                                                                                      Mustafee T P, 1970. Occurrence of Ferrisiana virgata (Cockerell) (Pseudococcidae: Homoptera) on ramie (Boehmeria nivea Gaud.) in Assam. Indian Journal of Entomology. 32 (4), 389-390.

                                                                                                                      Perez-Gelabert D E, 2008. Arthropods of Hispaniola (Dominican Republic and Haiti): a checklist and bibliography. Zootaxa. 1-530.

                                                                                                                      Rosen D, 1981. A new species of Pseudaphycus (Hym.: Encyrtidae), with notes on the angelicus group. Entomophaga. 26 (3), 251-263. DOI:10.1007/BF02371875

                                                                                                                      Shakti Khajuria, Rai A K, Kanak Lata, 2013. Occurrence and distribution of insect pests attacking solanaceous vegetables in semi-arid region of central Gujarat. Insect Environment. 19 (4), 248-249. http://www.currentbiotica.com/Insect/Volume19-4/IE-V19(4)-11.pdf

                                                                                                                      Sharma D R, 2011. New pest problems on fruit crops in Punjab. Journal of Insect Science (Ludhiana). 24 (3), 300-304.

                                                                                                                      Silva V C P da, Kaydan M B, Silva-Torres C S A da, Torres J B, 2019. Mealybug species (Hemiptera: Coccomorpha: Pseudococcidae) on soursop and sugar apple (Annonaceae) in North-East Brazil, with description of a new species of Pseudococcus Westwood. Zootaxa. 4604 (3), 525-538. DOI:10.11646/zootaxa.4604.3.8

                                                                                                                      Sirisena U G A I, Watson G W, Hemachandra K S, Wijayagunasekara H N P, 2013. Mealybugs (Hemiptera: Pseudococcidae) species on economically important fruit crops in Sri Lanka. Tropical Agricultural Research. 25 (1), 69-82. http://www.pgia.ac.lk/sites/default/files/congress/journel/v25/Journal-No%201/Papers/7%20U.G.A.I.%20Sirisena_final_final_edited.pdf

                                                                                                                      Suh SooJung, Bombay K, 2015. Scale insects (Hemiptera: Coccoidea) found on dracaena and ficus plants (Asparagales: Asparagaceae, Rosales: Moraceae) from southeastern Asia. Insecta Mundi. 1-10. http://centerforsystematicentomology.org/default.asp?action=insectamundi&id=insecta_new&year=2015

                                                                                                                      Trjapitzin SV, Trjapitzin VA, 1999. Parasitoids of the mealybugs on cultivated grapes in Argentina, with description of a new species of the genus Aenasius Walker (Hymenoptera, Encyrtidae). In: Entomologicheskoe Obozrenye, 76 174-179.

                                                                                                                      UK, CAB International, 1966. Ferrisia virgata. [Distribution map]. In: Distribution Maps of Plant Pests, Wallingford, UK: CAB International. Map 219. DOI:10.1079/DMPP/20056600219

                                                                                                                      Vidya M, Rajanna K M, Reddy M N N, Babu V, 2015. Insect pest complex of cashew and influence of biotic and abiotic factors on the incidence of cashew pests in maidan parts of Karnataka. Acta Horticulturae. 409-414. http://www.actahort.org/books/1080/1080_54.htm

                                                                                                                      Villatoro-Moreno H, Cisneros J, Gómez J, Infante F, Castillo A, 2016. Mealybugs (Hemiptera: Pseudococcidae) associated with rambutan (Nephelium lappaceum L.) in Chiapas, Mexico. Journal of the Kansas Entomological Society. 89 (4), 289-296. http://www.bioone.org/loi/kent DOI:10.2317/0022-8567-89.4.289

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

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

                                                                                                                      Williams D J, 1985. Australian mealybugs. London, United Kingdom: British Museum (Natural History). viii + 431 pp.

                                                                                                                      Williams D J, 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 D J, Watson G W, 1988. Scale insects of the tropical South Pacific region. Part 2. Mealybugs (Pseudococcidae). Wallingford, Oxon, UK: CAB International. 260 pp.

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

                                                                                                                      Wong CY, Chen SP, Chou LY, 1999. Guidebook to Scale Insects of Taiwan., Wufeng, Taichung, Taiwan: Taiwan Agricultural Research Institute. 98 pp.

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                                                                                                                      10/06/2016 Datasheet Updated by:

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

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