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Datasheet

Dysmicoccus neobrevipes
(grey pineapple mealybug)

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Datasheet

Dysmicoccus neobrevipes (grey pineapple mealybug)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Vector of Plant Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Dysmicoccus neobrevipes
  • Preferred Common Name
  • grey pineapple mealybug
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Dysmicoccus neobrevipes is a mealybug with a pantropical distribution. It is an economically important pest that can feed on and damage dozens of hosts, principally pineapple and the banana Musa × paradisia...

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Pictures

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PictureTitleCaptionCopyright
Heavy colonization of mealybugs can be seen on base of fruits, leaves and crowns.
TitleD. neobrevipes on pineapple
CaptionHeavy colonization of mealybugs can be seen on base of fruits, leaves and crowns.
CopyrightU.B. Gunasinghe
Heavy colonization of mealybugs can be seen on base of fruits, leaves and crowns.
D. neobrevipes on pineappleHeavy colonization of mealybugs can be seen on base of fruits, leaves and crowns.U.B. Gunasinghe

Identity

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Preferred Scientific Name

  • Dysmicoccus neobrevipes Beardsley

Preferred Common Name

  • grey pineapple mealybug

International Common Names

  • English: annona mealybug; gray pineapple mealybug

EPPO code

  • DYSMNE (Dysmicoccus neobrevipes)

Summary of Invasiveness

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Dysmicoccus neobrevipes is a mealybug with a pantropical distribution. It is an economically important pest that can feed on and damage dozens of hosts, principally pineapple and the banana Musa × paradisiaca. The main damage caused by D. neobrevipes is due to its role as a vector of mealybug wilt (Plant Health Australia, 2013). Qin et al. (2010) considered it a dangerous alien species with a high risk of invasion in China. Although D. neobrevipes can colonize without the help of associated caretaker ants, most commonly Pheidole and Solenopsis, the ants’ presence can help them to invade new areas by providing shelter and protecting them from natural enemies and adverse weather conditions.

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: Dysmicoccus
  •                                         Species: Dysmicoccus neobrevipes

Notes on Taxonomy and Nomenclature

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All members of what we now consider the pineapple mealybug complex were originally referred to as a single species called Pseudococcus brevipes (Cockerell) (Carter, 1933), which was later renamed Dysmicoccus brevipes. D. neobrevipes was found to have morphological differences from D. brevipes and was therefore described as a distinct species by Beardsley in 1959 from samples collected in Hawaii (Beardsley, 1992). However, this species had been recognised earlier by Ito (1938), who found two forms of pineapple mealybug in Hawaii, a pink form (D. brevipes) and a grey form (subsequently described as D. neobrevipes).

Description

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Live adult females are oval, grey, and coated with white mealy wax which forms small tufts (Beardsley 1959). They are 1.5 mm long and 1.0 mm wide. Authoritative identification requires slide-mounted adult females under a compound light microscope. See Beardsley (1959) for a detailed description of the D. neobrevipes.

Distribution

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D. neobrevipes is thought to be native to tropical America, but now has a pantropical distribution, with a small number of records from subtropical or Mediterranean localities (Kessing and Mau, 1992). It has been found in all zoogeographic regions on a wide diversity of hosts, but has a smaller geographical range than the related pink form, D. brevipes (Jahn et al., 2003). D. neobrevipes is known to have been introduced to China, Japan, Sri Lanka and Lithuania.

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentIntroducedHe et al., 2014
IndiaPresentGarcía et al., 2016
JapanPresentIntroducedTanaka and Uesato, 2012; García et al., 2016
-Ryukyu ArchipelagoPresentTanaka and Uesato, 2012
MalaysiaPresentGarcía et al., 2016
PakistanPresentGarcía et al., 2016
PhilippinesPresentAguilar et al., 2014; García et al., 2016
SingaporePresentGarcía et al., 2016
Sri LankaPresentIntroducedSirisena et al., 2013
ThailandWidespreadSarkar et al., 2014; García et al., 2016
VietnamPresentGarcía et al., 2016

Africa

UgandaPresentBua et al., 2013

North America

MexicoPresentBeardsley, 1965; García et al., 2016
USAPresentPresent based on regional distribution.
-FloridaPresentUnited States Department of Agriculture, 1979; García et al., 2016
-HawaiiPresentHu et al., 2009; García et al., 2016

Central America and Caribbean

Antigua and BarbudaPresentGarcía et al., 2016
BahamasPresentGarcía et al., 2016
Costa RicaPresentGarcía et al., 2016
Dominican RepublicPresentGarcía et al., 2016
El SalvadorPresentGarcía et al., 2016
GuatemalaPresentGarcía et al., 2016
HaitiPresentGarcía et al., 2016
HondurasPresentGarcía et al., 2016
JamaicaPresentBeardsley, 1975; García et al., 2016
PanamaPresentGarcía et al., 2016
Puerto RicoPresentGarcía et al., 2016
Trinidad and TobagoPresentGarcía et al., 2016
United States Virgin IslandsPresentGarcía et al., 2016

South America

BrazilPresentGarcía et al., 2016
ColombiaPresentGarcía et al., 2016
EcuadorPresentGarcía et al., 2016
PeruPresentGarcía et al., 2016
SurinamePresentGarcía et al., 2016

Europe

ItalyPresentTranfaglia, 1983; García et al., 2016
-SicilyPresentGarcía et al., 2016
LithuaniaPresentIntroducedMalumphy et al., 2008

Oceania

American SamoaPresentGarcía et al., 2016
Cook IslandsPresentGarcía et al., 2016
FijiPresentBeardsley, 1965; García et al., 2016
GuamPresentBeardsley, 1965
KiribatiPresentBeardsley, 1965; García et al., 2016
Marshall IslandsPresentGarcía et al., 2016
Northern Mariana IslandsPresentGarcía et al., 2016
SamoaPresentGarcía et al., 2016

History of Introduction and Spread

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D. neobrevipes is thought to be native to tropical America and to have been introduced to Hawaii some time before the establishment of plant quarantine regulations (Rohrbach et al., 1988). It has presumably since been introduced to all its other recorded localities outside the Neotropics via trade in infested plant material.

Risk of Introduction

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D. neobrevipes has the potential to cause harm to its hosts in areas where it is introduced that lack natural enemies, or where it is protected from natural enemies by caretaker ants, most commonly species of Pheidole and Solenopsis (Jahn et al., 2003). D. neobrevipes is under quarantine restrictions in the USA, where it has been intercepted many times, particularly on samples from the Philippines (Ben-Dov, 2001). Qin et al. (2010) conducted an investigation into the risk of invasion by D. neobrevipes into China. Their results suggested that D. neobrevipes is a dangerous alien species with a high risk of invasion.

Habitat

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D. neobrevipes has a pantropical distribution, with a small number of records from subtropical or Mediterranean localities. It is mostly found wherever its main host, pineapple, is grown (Kessing and Mau, 1992).

Habitat List

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CategoryHabitatPresenceStatus
Terrestrial-managed
Cultivated / agricultural land Principal habitat Harmful (pest or invasive)
Cultivated / agricultural land Principal habitat Natural

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Acacia (wattles)FabaceaeMain
Acacia farnesiana (huisache)FabaceaeMain
Acacia koa (koa)FabaceaeMain
AgaveAgavaceaeMain
Agave sisalana (sisal hemp)AgavaceaeMain
AglaonemaAraceaeMain
AlpiniaZingiberaceaeMain
Alpinia purpurata (red ginger)ZingiberaceaeMain
Anagallis arvensis (scarlet pimpernel)PrimulaceaeMain
Ananas comosus (pineapple)BromeliaceaeMain
AnnonaAnnonaceaeMain
Annona reticulata (bullock's heart)AnnonaceaeMain
Annona squamosa (sugar apple)AnnonaceaeMain
ArachisFabaceaeMain
Artocarpus (breadfruit trees)MoraceaeMain
Artocarpus altilis (breadfruit)MoraceaeMain
Artocarpus heterophyllus (jackfruit)MoraceaeMain
BarringtoniaLecythidaceaeMain
Barringtonia asiatica (sea poison tree)LecythidaceaeUnknown
CajanusFabaceaeMain
CitrusRutaceaeMain
Citrus aurantiifolia (lime)RutaceaeMain
Citrus reticulata (mandarin)RutaceaeMain
Citrus sinensis (navel orange)RutaceaeMain
Clerodendrum (Fragrant clerodendron)LamiaceaeMain
Coccoloba (sea grape)PolygonaceaeMain
Cocos nucifera (coconut)ArecaceaeMain
Codiaeum (ornamental croton)EuphorbiaceaeMain
Coffea (coffee)RubiaceaeMain
Crescentia alataBignoniaceaeOther
Crescentia alataBignoniaceaeUnknown
Cucurbita maxima (giant pumpkin)CucurbitaceaeMain
Garcinia mangostana (mangosteen)ClusiaceaeMain
Gossypium (cotton)MalvaceaeMain
GuettardaRubiaceaeUnknown
HeliconiaHeliconiaceaeMain
Mangifera indica (mango)AnacardiaceaeMain
Manilkara zapota (sapodilla)SapotaceaeMain
Musa (banana)MusaceaeMain
Musa acuminata (wild banana)MusaceaeMain
Musa x paradisiaca (plantain)MusaceaeMain
Nephelium lappaceum (rambutan)SapindaceaeMain
Opuntia (Pricklypear)CactaceaeMain
Opuntia megacanthaCactaceaeUnknown
Pandanus (screw-pine)PandanaceaeMain
Phaseolus (beans)FabaceaeMain
PhilodendronAraceaeMain
Pipturus argenteusUrticaceaeUnknown
Piscidia piscipulaFabaceaeUnknown
PolianthesAgavaceaeMain
Polianthes tuberosa (tuberose)AgavaceaeMain
Psidium guajava (guava)MyrtaceaeMain
PunicaPunicaceaeMain
Punica granatum (pomegranate)PunicaceaeMain
SamaneaFabaceaeMain
Samanea saman (rain tree)FabaceaeMain
Solanum lycopersicum (tomato)SolanaceaeMain
Solanum melongena (aubergine)SolanaceaeMain
TectonaLamiaceaeMain
TheobromaSterculiaceaeMain
Theobroma cacao (cocoa)SterculiaceaeMain
ThespesiaMalvaceaeMain
Thespesia populnea (portia tree)MalvaceaeMain
Tournefortia argenteaBoraginaceaeUnknown
Vigna unguiculata (cowpea)FabaceaeMain
Wrightia arborea (lanete)ApocynaceaeMain
YuccaAgavaceaeMain

Growth Stages

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

Symptoms

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D. neobrevipes is usually found near the top of the host plant and feeds by sucking phloem sap from the plant tissue. This may cause local lesions to form at the site of feeding on some hosts. These lesions are bizonate, with a dark green centre surrounded by a lighter green area (Dasgupta, 1988). D. neobrevipes also affects the plant’s photosyntheitic ability by excreting sugary honeydew that fouls plant surfaces, forming a medium for the growth of sooty mould, which blocks sunlight and air from reaching the leaves, impairing photosynthesis (Tabata and Ichiki, 2015).

The main damage that pineapple mealybugs such as D. neobrevipes cause is as a result of their role as a vector of pineapple wilt. This devastating disease is caused by Pineapple mealybug wilt associated virus-2 (PMWaV-2), a mealybug-transmitted ampelovirus (Subere et al., 2011). There are two types of wilt, quick wilt and slow wilt. Quick wilt, also known as mealybug wilt, develops around 2 months after a short attack by a large colony of mealybugs, whereas slow wilt is caused by many mealybugs feeding on the plant tissue over many months (Jahn et al., 2003). Slow wilt causes the inner leaves to turn dry and brown, and outer leaves to lose their turgidity and droop (Jahn et al., 2003). Unlike slow wilt, quick wilt causes leaves to turn a light-green to yellow-pink colour in plants younger than 6 months. In older plants, quick wilt causes leaves to droop, turn pink and dry out (Carter, 1932).

Both types of wilt cause leaves to droop and dry out. They also both affect the fruit yield of the plant, especially if symptoms are seen early in the season. Affected plants either produce smaller fruit or produce no fruit at all. Pineapple wilt may also result in the invasion of saprophytic organisms, which leads to collapse of the roots (Kessing and Mau, 1992). Ultimately, plants may die as a result of infection by pineapple wilt transmitted by D. neobrevipes.

D. neobrevipes also causes green spot disease of pineapple, which is characterized by galls on leaves caused by a reaction between the plant and a secretion from the mealybugs.

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
Roots / external feeding
Stems / discoloration of bark
Stems / external feeding
Stems / honeydew or sooty mould
Whole plant / discoloration
Whole plant / external feeding

Biology and Ecology

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Reproductive Biology

D. neobrevipes males and females reproduces sexually. Unmated females will not produce young (Beardsley, 1960). D. neobrevipes is ovoviviparous, with each female giving birth to approximately 350 live young in a lifetime, although it can be as many as 1000 (Kessing and Mau, 1992). Females go through 3 larval instars which last for 11 to 23 days, 6 to 20 days and 7 to 28 days, respectively. The total larval period is 35 days on average but can range from 26 to 52 days.

Males go through 4 larval instars before becoming winged adults. These instars last for 11 to 19 days, 7 to 19 days, 2 to 7 days and 2 to 8 days, respectively. The total larval period ranges from 22 to 53 days (Kessing and Mau, 1992).

Longevity

Adult females can live for 48 to 72 days, whereas the winged males live for 2 to 7 days (Kessing and Mau, 1992). Qin et al. (2011) found that the longevity of D. neobrevipes can vary depending on the mealybug’s host; the longevity of D. neobrevipes on most hosts tested was 51.0 days, but they recorded a longevity of 62.5 days for female mealybugs on Ananas comosus Baili.

Associations

D. neobrevipes has a mutualistic symbiotic relationship with ants. The genera of ants most commonly associated with D. neobrevipes are Pheidole and Solenopsis (Jahn et al., 2003).

Ants benefit from the mutualism by feeding on the sugary honeydew produced by the mealybugs and also by feeding on some of the mealybugs’ natural enemies. The removal of honeydew reduces the likelihood of fungal attack on the mealybugs, and the reduction of natural enemies protects the mealybugs from predation and parasitism. Jahn et al. (2003) reviewed a number of studies on Pheidole megacephala, the big-headed ant, and found that this species, which is dominant in pineapple crops in Hawaii, deterred natural enemies of D. neobrevipes. In addition to these benefits, the ants also protect mealybugs from inclement weather by providing earthen shelters, or by transporting them to safer areas. Jahn and Beardsley (1996) reported that in the absence of both natural enemies and adverse weather, P.megacephala provided no benefit to the mealybug population.

Climate

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ClimateStatusDescriptionRemark
Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aenasius brasiliensis Parasite
Anagyrus ananatis Parasite
Argiope appensa Predator
Blepyrus propinquus Parasite Hawaii apples
Chelisoches morio Predator
Conocephalus saltator Predator
Cryptolaemus montrouzieri Predator
Curinus caeruleus Predator
Diadiplosis pseudococci Predator
Dicrodiplosis guatemalensis Predator
Euborellia annulipes Predator
Exochomus concavus Predator
Hambeltonia pseudococcinna Parasite
Hyperaspis c-nigrum Predator
Hyperaspis silvestri Predator
Labidura riparia Predator
Nephus bipunctatus Predator
Pseudaphycus dysmicocci Parasite
Pseudiastata nebulosa Predator
Rhyzobius forestieri Predator
Rhyzobius ventralis Predator
Scymnus apiciflavus Predator
Scymnus margipellens Predator
Scymnus uncinatus Predator
Tropidophryne melvillei Parasite
Zaplatycerus fullawayi Parasite

Means of Movement and Dispersal

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Natural Dispersal

First-instar nymphs of D. neobrevipes crawl upwards and can be dispersed by the wind (Jahn and Beardsley, 2000). Ants have been observed aiding the mealybugs’ colonization of new plants by carrying mealybugs in their mandibles, helping them to disperse within and between fields (Phillips, 1934). However, in laboratory experiments by Jahn and Beardsley (1996), the ant Pheidole megacephala did not transport mealybugs in any significant numbers.

First instar D. neobrevipes, also known as crawlers, can be dispersed by the wind (Jahn and Beardsley, 2000).

Impact Summary

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CategoryImpact
Environment (generally) Negative

Economic Impact

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D. neobrevipes is a pest of many economically important crops, particularly pineapple and banana. In addition to the direct damage it causes (see Symptoms), it also causes green spot disease and transmits pineapple wilt which is caused by Pineapple mealybug wilt associated virus-2 (PMWaV-2). Sether and Hu (2002) showed that pineapple wilt can cause yield loss of 35% in pineapple, representing significant losses to producers.

Threatened Species

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

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Tolerant of shade
  • Capable of securing and ingesting a wide range of food
  • Gregarious
Impact outcomes
  • Host damage
  • Negatively impacts agriculture
  • Negatively impacts livelihoods
  • Damages animal/plant products
Impact mechanisms
  • Pest and disease transmission
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally

Detection and Inspection

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D. neobrevipes crawlers (first-instar nymphs) can be detected in the field using blue sticky traps. Jahn and Beardsley (2000) found blue sticky traps better for trapping D. neobrevipes than yellow sticky traps, which attract high numbers of other insects, such as flies.

D. neobrevipes is only found on the aerial parts of the plant (Beardsley 1960) and is usually seen on the surface, but can feed deep in the leaf axils or within the blossom cups. Therefore, a plant may need to be dissected in order to find all of the mealybugs on it (Jahn et al., 2003).

Similarities to Other Species/Conditions

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There are over 140 species in the genus Dysmicoccus but D. neobrevipes is most similar to D. brevipes, which is known as the pink form of the pineapple mealybug. Beardsley (1959) described the morphological differences between these two species and therefore was able to define them as two separate species. In D. neobrevipes the ventral sclerotization of the anal lobes is elongate, whereas in D. brevipes these areas are quadrate. Beardsley (1959) also noted that D. brevipes has several long setae on either side of the mid-dorsal axis of the 9th abdominal segment (now called 8th abdominal segment) which ranged from 45-80 µm long, whereas in D. neobrevipes the longest setae in this region are about 15 µm long, no longer than the other dorsal setae.

Prevention and Control

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Prevention

Jahn (1990) showed that controlling mutualistic ants will prevent the build-up of mealybug populations, by reducing the protection the mealybugs benefit from, thereby allowing natural enemies to prey on any mealybugs present.

Physical barriers such as ant fences have shown partial effectiveness (Kessing and Mau, 1992), but the most effective ant control involves the use of insecticidal baits. A commonly used chemical for these baits is hydramethylnon (Jahn et al., 2003).

Control

Although it is thought that the control of ants is an effective way to prevent the build-up of D. neobrevipes, the mealybugs themselves may also need controlling.

Even if mealybugs have already established and symptoms of pineapple wilt have been observed, control should still be implemented as recovery can be rapid if the appropriate control measures are put in to place (Carter, 1967).

Movement Control

Controlling ants will prevent them from transporting mealybugs further in an area. The use of wind barriers around the edge of the field can reduce the number of mealybug crawlers (first-instar nymphs) that are spread by the wind to uninfested areas (Jahn and Beardsley, 2000).

Biological Control

D. neobrevipes has a range of natural enemies that, in the absence of caretaker ants, can effectively control populations of the mealybug. New predators can be introduced to an area in order to control the mealybugs, but without first controlling ant populations, these introductions will not be effective (Rohrbach et al. 1988).

Chemical Control

Chemicals can be effective at controlling D. neobrevipes, but if the mealybugs are feeding deep in the plant, they will be shielded from the application of insecticides. The waxy coating on the mealybugs may prevent penetration of chemical sprays, reducing their effectiveness. Insecticides that have been used in the control of mealybugs include the organophosphates malathion and diazinon (Jahn and Beardsley, 2003). The most effective ant control involves the use of insecticidal baits (Jahn et al. 2003).

References

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Aguilar CH, Lasalita-Zapico F, Namocatcat J, Fortich A, Bojadores RM, 2014. Farmers' perceptions about banana insect pests and integrated pest management (IPM) systems in SocSarGen, Mindanao, Philippines. International Proceedings of Chemical, Biological and Environmental Engineering (IPCBEE) [2014 International Conference on Intelligent Agriculture, Hong Kong, China, 13-14 February 2014.], 63:22-27. http://www.ipcbee.com/vol63/005-ICOIA2014-IA0015.pdf

Beardsley JW, 1959. On the taxonomy of Pineapple mealybugs in Hawaii, with a description of a previously unnamed species (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, 17:29-37.

Beardsley JW, 1960. A preliminary study of the males of some Hawaiian mealybugs (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, 17:199-243.

Beardsley JW, 1965. Notes on the Pineapple Mealybug Complex, with descriptions of two new species (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, 19:55-68.

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

Beardsley JW, 1975. Homoptera: Coccoidea. Supplement. In: Insects of Micronesia 6. 657-662.

Beardsley JW, 1992. The pineapple mealybug complex; taxonomy, distribution and host relationships. International Pineapple Symposium, 334:383-386.

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.

Ben-Dov Y, 2001. Dysmicoccus neobrevipes. ScaleNet. http://www.sel.barc.usda.gov/scalekeys/mealybugs/key/mealybugs/media/html/Species/Dysmicoccus_neobrevipes/Dysmicoccus_neobrevipes.html

Bua B, Karungi J, Kawube G, 2013. Occurrence and effects of pineapple mealybug wilt disease in central Uganda. Journal of Agricultural Science and Technology A, 3(5):410-416. http://www.davidpublishing.com/show.html?12655

Carter W, 1932, December. The Pineapple Mealybug (Pseudococcus brevipes (CM.)) and Wilt of Pineapples. (Abstract.). Phytopathology, 20(12):996-997 pp.

Dasgupta MK, 1988. Principles of Plant Pathology. New Delhi, India: Allied Publishers Limited, 1043 pp.

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

He YanBiao, Liu YingHong, Zhan RuLin, Xu ZaiFu, Sun GuangMing, Zhao YanLong, Li GuoPing, Chang JinMei, 2014. The occurrence of two species of pineapple mealybugs (Dysmicoccus spp.) (Hemiptera: Pseudococcidae) in China and their genetic relationship based on rDNA ITS sequences. Caryologia, 67(1):36-44. http://www.tandfonline.com/doi/full/10.1080/00087114.2014.891698

Hu JS, Sether DM, Melzer MJ, Subere CV, Cheah K, Chen Y, Li Q, Borth W, Wang IC, Nagai C, Wang ML, 2009. Characterization and management of pineapple mealybug wilt associated viruses. Acta Horticulturae [Proceedings of the Sixth International Pineapple Symposium, Joao Pessoa, Brazil, 18-23 November 2007.], No.822:185-189. http://www.actahort.org

Ito K, 1938. Studies on the life history of the pineapple mealybug, Dysmicoccus brevipes (Ckll.). Journal of Economic Entomology, 31:291-298.

Jahn GC, Beardsley JW, 2000. Interactions of ants (Hymenoptera: Formicidae) and mealybugs (Homoptera: Pseudococcidae) on pineapple. Proceedings of the Hawaiian Entomological Society, 34:181-185.

Jahn GC, Beardsley JW, González-Hernández H, 2003. A review of the association of ants with mealybug wilt disease of pineapple. Proceedings of the Hawaiian Entomological Society, 36:9-28.

Kessing JLM, Mau RFL, 1992. Dysmicoccus neobrevipes (Beardsley). Crop Knowledge Master. http://www.extento.hawaii.edu/kbase/crop/type/d_neobre.htm

Malumphy C, Ostrauskas H, Pye D, 2008. A provisional catalogue of scale insects (Hemiptera, Coccoidea) of Lithuania. Acta Zoologica Lituanica, 18(2):108-121. http://versita.metapress.com/content/u651315215867370/fulltext.pdf

Phillips JS, 1934, August. Bull. Exp. Stn. Pineapple Prod. coop. Ass. Hawaii, 15. Honolulu, iii + 57 pp.

Plant Health Australia, 2013. Exotic threat: grey pineapple mealybug. Australia Plant Health Factsheet. http://www.planthealthaustralia.com.au/wp-content/uploads/2013/03/Grey-pineapple-mealy-bug-FS.pdf

Rohrbach KG, Beardsley JW, German TL, Reimer NJ, Sanford WG, 1988. Mealybug wilt, mealybugs, and ants of pineapple. Plant Disease, 72(7):558-565

Sarkar MA, Wiwat Suasa-ard, Sopon Uraichuen, 2014. Suitability of different mealybug species (Hemiptera: Pseudococcidae) as hosts for the newly identified parasitoid Allotropa suasaardi Sarkar & Polaszek (Hymenoptera: Platygasteridae). Kasetsart Journal, Natural Science, 48(1):17-27. http://kasetsartjournal.ku.ac.th/kuj_files/2014/A1404181034288437.pdf

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16/03/15 Original text by:

Claire Curry, CABI, UK

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