Paracoccus marginatus (papaya mealybug)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Vectors
- Plant Trade
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Paracoccus marginatus Williams & Granara de Willink, 1992
Preferred Common Name
- papaya mealybug
International Common Names
- French: cochenille du papayer
- PACOMA (Paracoccus marginatus)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hemiptera
- Suborder: Sternorrhyncha
- Unknown: Coccoidea
- Family: Pseudococcidae
- Genus: Paracoccus
- Species: Paracoccus marginatus
Notes on Taxonomy and NomenclatureTop of page Paracoccus marginatus was described relatively recently (Williams and Granara de Willink, 1992) and the identity of the species is clear. There are no scientific synonyms.
DescriptionTop of page
Crawlers (0.3 mm long) are yellow. Immature and newly matured females have yellowish bodies dusted with mealy white wax that is often thinner between the segments, giving the body a slightly barred appearance. Short, waxy filaments develop around the margin in the adult female, each less than a quarter as long as the body. The adult female is 2.5-4 mm long, soft-bodied, elongate oval and slightly flattened; on maturation she begins to secrete sticky, elastic, white wax filaments from the edges of her abdomen to form a protective ovisac for her yellow eggs. The ovisac can be as much as twice as long as the body, or more (Miller et al., 2001). Sometimes the body colour of the mature female is not immediately apparent if she has become buried under white, waxy secretions. The male is a short-lived, small insect with long, segmented antennae; six legs each bearing a single claw; one pair of simple wings coated with white wax powder; a pair of long, white waxy filaments at the posterior of the abdomen; and no mouthparts.
A redescription of this species is provided by Miller and Miller (2002) including decriptions of the immature stages and adult male.
DistributionTop of page
P. marginatus probably originated in Mexico or a nearby country in Central America. It was not recorded from the Caribbean islands before 1994, but has been extending its range in the region ever since. It was first recorded from USA (Florida) in 1998 (Miller et al., 2001). Matile-Ferrero et al. (2001) mention that P. marginatus is present in St Martin, St Barthélémy and most of the islands in the Greater and Lesser Antilles; their record of the species from French Guyana is the first from northern South America.
P. marginatus has recently been reported spreading in Asia and West Africa. In addition to the countries listed, it has also been reported from Nigeria and Gabon, and has now been discovered in Tanzania (IITA, 2015).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Bangladesh||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Cambodia||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|China||Present||Wu FuZhong et al., 2014|
|-Yunnan||Present||Wu FuZhong et al., 2014|
|India||Present||Invasive||Muniappan, 2009a; Suresh et al., 2010; CABI/EPPO, 2012|
|-Karnataka||Present||Shekhar et al., 2011; CABI/EPPO, 2012|
|-Kerala||Present||Krishnakumar and Rajan, 2009; Lyla and Philip, 2010; CABI/EPPO, 2012|
|-Rajasthan||Present||Mani et al., 2012|
|-Tamil Nadu||Present||Invasive||Muniappan, 2009a; Ayyasamy and Regupathy, 2010; Suresh et al., 2010; CABI/EPPO, 2012|
|-West Bengal||Present||Lalitha et al., 2015|
|Indonesia||Restricted distribution||Muniappan, 2009a; IPPC, 2009; CABI/EPPO, 2012|
|-Java||Present||Muniappan, 2009a; CABI/EPPO, 2012|
|Israel||Present||Mendel et al., 2016|
|Malaysia||Present||Mastoi et al., 2011; CABI/EPPO, 2012|
|Oman||Present||unpublished data, R. Muniappan, Virginia Tech, USA, 2011|
|Philippines||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Sri Lanka||Present||Galanihe et al., 2010; CABI/EPPO, 2012|
|Taiwan||Present||Chen et al., 2011; CABI/EPPO, 2012|
|Thailand||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Benin||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Cameroon||Restricted distribution||Introduced||2015||IPPC, 2015||preliminary report|
|Gabon||Present||Introduced||2012||IPPC, 2012||present: under eradication|
|Ghana||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Kenya||Present||Macharia et al., 2017||via PestLens newsletter|
|Mauritius||Present||Germain et al., 2014|
|Réunion||Present||Germain et al., 2010; CABI/EPPO, 2012|
|Togo||Present||Muniappan et al., 2009b; CABI/EPPO, 2012|
|Mexico||Present||Native||Williams and Granara de Willink, 1992; Pollard, 1999; CABI/EPPO, 2012|
|USA||Restricted distribution||Pollard, 1999; CABI/EPPO, 2012|
|-Florida||Present||Pollard, 1999; CABI/EPPO, 2012|
|-Hawaii||Present||Muniappan, 2009a; CABI/EPPO, 2012|
Central America and Caribbean
|Antigua and Barbuda||Present||Pollard, 1999; CABI/EPPO, 2012|
|Belize||Present||Williams and Granara de Willink, 1992; Pollard, 1999; CABI/EPPO, 2012|
|British Virgin Islands||Present||Pollard, 1999; CABI/EPPO, 2012|
|Cayman Islands||Present||CABI/EPPO, 2012|
|Costa Rica||Present||Williams and Granara de Willink, 1992; CABI/EPPO, 2012|
|Dominican Republic||Present||Pollard, 1999; CABI/EPPO, 2012|
|French West Indies||Present||CABI/EPPO, 2012|
|Guadeloupe||Present||Matile-Ferrero and Etienne, 1998; CABI/EPPO, 2012|
|Guatemala||Present||Williams and Granara de Willink, 1992; CABI/EPPO, 2012|
|Haiti||Present||Pollard, 1999; CABI/EPPO, 2012|
|Netherlands Antilles||Present||Pollard, 1999; CABI/EPPO, 2012|
|Puerto Rico||Present||Pollard, 1999; CABI/EPPO, 2012|
|Saint Kitts and Nevis||Present||Pollard, 1999; CABI/EPPO, 2012|
|Saint Lucia||Present||Introduced||Invasive||Jn Pierre, 2008|
|Sint Maarten||Present||Pollard, 1999|
|United States Virgin Islands||Present||Pollard, 1999; CABI/EPPO, 2012|
|French Guiana||Present||Matile-Ferrero et al., 2001; CABI/EPPO, 2012|
|Guam||Present||Meyerdirk et al., 2004; CABI/EPPO, 2012|
|Northern Mariana Islands||Present||Muniappan, 2009a; CABI/EPPO, 2012|
|Palau||Present||Muniappan et al., 2006; CABI/EPPO, 2012|
Risk of IntroductionTop of page At present P. marginatus is confined to one small part of the globe, but if it spreads outside this area it could become an important tropical pest. It is a quarantine threat to farmers and horticulturalists in tropical countries around the world. P. marginatus is not included on any quarantine schedules because its increase in importance is so recent. This pest probably presents its greatest threat to countries within 30° of the equator.
HabitatTop of page P. marginatus has only been recorded feeding on above-ground parts of its hosts, particularly on leaves and fruit (Miller et al., 2001).
Hosts/Species AffectedTop of page The species was described from material collected on cassava and papaya, but it is now recognized to be highly polyphagous (Anon., 2000). It causes significant damage to cassava in Central America. Papaya, Hibiscus and Annona species are particularly badly affected (Matile-Ferrero et al., 2001). Papaya is a preferred host.
Host Plants and Other Plants AffectedTop of page
|Acacia (wattles)||Fabaceae||Wild host|
|Acalypha (Copperleaf)||Euphorbiaceae||Wild host|
|Ananas comosus (pineapple)||Bromeliaceae||Other|
|Annona squamosa (sugar apple)||Annonaceae||Main|
|Cajanus cajan (pigeon pea)||Fabaceae||Other|
|Capsicum annuum (bell pepper)||Solanaceae||Other|
|Carica papaya (pawpaw)||Caricaceae||Main|
|Cestrum nocturnum (night jessamine)||Solanaceae||Wild host|
|Citrus sinensis (navel orange)||Rutaceae||Other|
|Dahlia pinnata (garden dahlia)||Asteraceae||Other|
|Eugenia uniflora (Surinam cherry)||Myrtaceae||Other|
|Gossypium hirsutum (Bourbon cotton)||Malvaceae||Other|
|Guazuma ulmifolia (bastard cedar)||Sterculiaceae||Wild host|
|Hibiscus rosa-sinensis (China-rose)||Malvaceae||Main|
|Hibiscus sabdariffa (Roselle)||Malvaceae||Main|
|Ipomoea (morning glory)||Convolvulaceae||Other|
|Jatropha curcas (jatropha)||Euphorbiaceae||Main|
|Lablab purpureus (hyacinth bean)||Fabaceae||Other|
|Malpighia glabra (acerola)||Malpighiaceae||Wild host|
|Malvaviscus arboreus (wax mallow)||Malvaceae||Wild host|
|Mangifera indica (mango)||Anacardiaceae||Other|
|Manihot esculenta (cassava)||Euphorbiaceae||Main|
|Mimosa pigra (catclaw mimosa)||Fabaceae||Wild host|
|Morus alba (mora)||Moraceae||Other|
|Pachystachys lutea (lollypops)||Acanthaceae||Wild host|
|Parthenium hysterophorus (parthenium weed)||Asteraceae||Wild host|
|Persea americana (avocado)||Lauraceae||Main|
|Psidium guajava (guava)||Myrtaceae||Other|
|Punica granatum (pomegranate)||Punicaceae||Other|
|Solanum lycopersicum (tomato)||Solanaceae||Other|
|Solanum melongena (aubergine)||Solanaceae||Main|
|Solanum nigrum (black nightshade)||Solanaceae||Other|
|Solanum torvum (turkey berry)||Solanaceae||Other|
|Theobroma cacao (cocoa)||Sterculiaceae||Other|
Growth StagesTop of page Flowering stage, Fruiting stage, Post-harvest, Vegetative growing stage
SymptomsTop of page Heavy infestations by P. marginatus cause deformation of new growth, leaf yellowing, leaf curl and early fall of fruit (Anon., 2000). Fruit may become completely covered by a layer of mealybugs and wax secretions (Miller et al., 2001). Papaya, Hibiscus and Annona species are particularly badly affected.
List of Symptoms/SignsTop of page
|Fruit / abnormal shape|
|Fruit / external feeding|
|Fruit / honeydew or sooty mould|
|Fruit / premature drop|
|Fruit / reduced size|
|Growing point / dieback|
|Growing point / distortion|
|Growing point / external feeding|
|Inflorescence / distortion (non-graminaceous plants)|
|Inflorescence / external feeding|
|Inflorescence / fall or shedding|
|Inflorescence / honeydew or sooty mould|
|Inflorescence / twisting and distortion|
|Leaves / abnormal forms|
|Leaves / external feeding|
|Leaves / honeydew or sooty mould|
|Leaves / yellowed or dead|
|Stems / dieback|
|Stems / external feeding|
|Stems / honeydew or sooty mould|
|Stems / stunting or rosetting|
|Stems / wilt|
|Whole plant / distortion; rosetting|
|Whole plant / early senescence|
|Whole plant / external feeding|
Biology and EcologyTop of page The biology of P. marginatus has not been studied in detail. The species is known to reproduce sexually, and each female lays a considerable number of eggs in a white, waxy ovisac. Like other mealybugs, female P. marginatus can be expected to have three larval stages before moulting to the larviform adult stage. The male is likely to have two immature larval stages that feed, followed by non-feeding pre-pupal and pupal stages before it moults to a short-lived, winged adult. In tropical conditions the generations are not synchronized and there are several each year - possibly as many as 15 generations in favourable conditions, like the similar Maconellicoccus hirsutus.
The dispersal stage is the first-instar crawler which can survive a day or so without feeding while it locates a suitable feeding site. The larval stages and adult female (but not the male prepupa or pupa) are capable of crawling, but seldom do so unless conditions become unfavourable.
P. marginatus feeds on phloem sap and excretes sugary honeydew; this coats plant surfaces around and below the colony, and sooty mould often develops on the deposits. Ants may be attracted to colonies in search of honeydew, and their presence can deter natural enemies from attacking the mealybugs.
The climatic preferences of P. marginatus have not been documented, but its occurrence in countries that mostly lie less than 30° from the equator suggests that it probably does not tolerate cold conditions. As for most scale insects, heavy rain is likely to cause increased mortality of P. marginatus, especially of the mobile first-instar crawlers.
Natural enemiesTop of page
Notes on Natural EnemiesTop of page P. marginatus has only recently been categorized as a pest, so little information is available on its natural enemies. Two hyperparasite species were collected from heavy infestations of P. marginatus on papaya from US Virgin Islands (St Thomas) in 1998 (ME Schauff, US Department of Agriculture, Museum of Natural History, Washington, D.C., unpublished data, 1998). Although no primary parasites were found, it is likely that they were present and probably belonged to the superfamily Chalcidoidea.
Four hymenopteran parasitoids of P. marginatus have been found in Mexico, and are being investigated for use in biological control (Anon., 2000).
The predatory ladybeetle used to control pest mealybugs, Cryptolaemus montrouzieri, was observed to be reluctant to feed on P. marginatus in the British Virgin Islands (Wayne deChi, Agricultural Health Specialist, Interamerican Institute for Cooperation on Agriculture (IICA), personal communication). A predatory dipteran larva (Diadiplosis sp.) has been observed eating the eggs of P. marginatus in Antigua (Wayne deChi, personal communication), and Matile-Ferrero et al. (2001) recorded two unidentified predators from French Guyana belonging to the Diptera: Cecidomyiidae, and the Coleoptera: Coccinellidae.
Means of Movement and DispersalTop of page
Natural dispersal (non-biotic)
As for most mealybugs, the tiny first-instar crawlers of P. marginatus disperse short distances by walking; those on exposed parts of the plant may be carried to other hosts over somewhat greater distances by gusts of wind.
Passing animals, including man, may accidentally pick up crawlers of P. marginatus as they brush past infested plants, and transfer them to new host plants by the same means. Birds such as crows and mynah, and bats play an important role in disseminating P. marginatus from infested plants with ripe fruits to healthy plants (Sarma, 2013).
Vehicles moving through a crop, or pruning and harvesting activities, can help carry P. marginatus crawlers from one plant to another.
Movement in trade
As for most mealybugs, transport of infested fresh plant material can transmit P. marginatus over greater distances, and potentially from one country to another.
Pathway VectorsTop of page
|Clothing, footwear and possessions||Air and surface transport of ornamental and crop plant material.||Yes|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Flowers/Inflorescences/Cones/Calyx||adults; eggs; larvae; nymphs; pupae||Yes||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Fruits (inc. pods)||adults; eggs; larvae; nymphs; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Leaves||adults; eggs; larvae; nymphs; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Stems (above ground)/Shoots/Trunks/Branches||adults; eggs; larvae; nymphs; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Plant parts not known to carry the pest in trade/transport|
|True seeds (inc. grain)|
ImpactTop of page
Colonies of P. marginatus excrete honeydew, which coats nearby plant surfaces. Fouling of plant leaves by honeydew and sooty mould blocks out air and light, impairing photosynthesis and reducing plant productivity. Fouled fruit, for example, papaya, may be of reduced value or unmarketable. Heavy infestations of thin-skinned fruit crops such as papaya can make them unmarketable. P. marginatus causes significant damage to cassava in Central America, and has the capacity to cause serious damage to papaya, other tropical fruit and ornamentals such as Annona and Hibiscus spp. (Anon., 2000; Miller et al., 2001). The species is a quarantine threat to farmers and horticulturalists in tropical countries around the world. Since its accidental introduction to Cuba in 1999, P. marginatus has been under close surveillance; so far no economic damage has been observed on major crops there (Anon., 2000).
The accidental introduction of P. marginatus to Indonesia and India presents a serious economic threat to the agricultural industry in these countries (Muniappan, 2009).
DiagnosisTop of page Authoritative identification requires preparation of good quality slide mounts of adult females, and identification using the key provided by Williams and Granara de Willink (1992). This publication also provides advice on how to prepare slide mounts of mealybugs, as do Watson and Chandler (2000).
Similarities to Other Species/ConditionsTop of page P. marginatus may show very similar symptoms to pink hibiscus mealybug (Maconellicoccus hirsutus) on Hibiscus, Annona species and other ornamental plants. M. hirsutus is the more serious pest of the two (Pollard, 1999). Although superficially similar in the field, they can be distinguished by the colour of the body contents when crushed on white paper: P. marginatus is yellow; M. hirsutus is pink. When preserved in 80% alcohol, P. marginatus turn black (Miller et al., 2001) within 24-48 h, whereas M. hirsutus specimens turn darker brown but do not go black. When adult females are mounted on microscope slides, the species can be easily distinguished: P. marginatus has eight-segmented antennae and dorsal oral rim ducts located only in marginal areas; M. hirsutus has nine-segmented antennae and rows of dorsal oral rim ducts across all the body segments.
There are numerous other species of Paracoccus native to the Neotropical region, and they can be difficult to separate. Authoritative identification requires preparation of good quality slide mounts of adult females, and identification using the key provided by Williams and Granara de Willink (1992).
Prevention and ControlTop of page
As an exotic introduction to the Caribbean islands, there are good prospects for classical biological control of P. marginatus by hymenopteran parasitoids originating from its area of origin in Central America (Pollard, 1999). In situations where the mealybugs are heavily attended by ants, for their honeydew, control of ants by the use of sticky bands on the trunk or branches may help enhance the effectiveness of natural enemies. Spacing or pruning of trees, to ensure their canopies do not touch, will reduce the spread of mealybugs between trees.
A biological control programme developed by the USDA/APHIS has been successfully implemented against P. marginatus in Florida, Caribbean Islands, countries in South America, Guam and Palau and is currently being implemented in the Hawaiian Islands and Tinian in the Mariana Islands (Muniappan, 2009). Three parasitoids have been identified (Anagyrus loecki, Pseudleptomastix mexicana and Acerophagus papayae) and are being reared in Puerto Rico in order to supply countries in need.
ReferencesTop of page
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Muniappan R, Shepard BM, Watson GW, Carner GR, Rauf A, Sartiami D, Hidayat P, Afun JVK, Goergen G, Ziaur Rahman AKM, 2009. New records of invasive insects (hemiptera: Sternorrhyncha) in Southeast Asia and West Africa. Journal of Agricultural and Urban Entomology, 26(4):167-174.
Muniappan R, Shepard BM, Watson GW, Carner GR, Sartiami D, Rauf A, Hammig MD, 2008, publ. 2009. First report of the papaya mealybug, Paracoccus marginatus (Hemiptera: Pseudococcidae), in Indonesia and India. Journal of Agricultural and Urban Entomology, 25(1):37-40. http://entweb.clemson.edu/scesweb
Noyes JS, Schauff ME, 2003. New Encyrtidae (Hymenoptera) from papaya mealybug (Paracoccus marginatus Williams and Granara De Willink) (Hemiptera: Sternorrhyncha: Pseudococcidae). Proceedings of the Entomological Society of Washington, 105(1):180-185.
Pantoja A, Abreu E, Peña J, Robles W, 2007. Paracoccus marginatus Williams and Granara de Willink (Homoptera: Pseudococcidae) affecting papaya in Puerto Rico. Journal of Agriculture of the University of Puerto Rico, 91(3/4):223-225.
Regupathy A, Ayyasamy R, 2009. Need for generating baseline data for monitoring insecticide resistance in new invasive mealy bug Paracoccus marginatus Williams and Granara de Willink (Insecta: Hemiptera: Pseudococcidae), the key pest of papaya and biofuel crop, Jatropha curcas. Resistant Pest Management Newsletter, 19(1):37-40. http://whalonlab.msu.edu/rpmnews/general/rpm_archive.htm
Suresh S, Jothimani R, Sivasubrmanian P, Karuppuchamy P, Samiyappan R, Jonathan EI, 2010. Invasive mealybugs of Tamil Nadu and their management. Karnataka Journal of Agricultural Sciences, 23(1):6-9. http://220.127.116.11/ojs/index.php/kjas/article/viewFile/1266/1210
Surís M, Lafergue S, 2005. Influence of the host in the biological development of Paracoccus marginatus (Williams and Granara de Willink). (Influencia del hospedante en el desarrollo biológico de Paracoccus marginatus (Williams y Granara de Willink).) Revista de Protección Vegetal, 20(3):169-172.
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