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Datasheet

Phenacoccus manihoti (cassava mealybug)

Summary

  • Last modified
  • 22 June 2017
  • Datasheet Type(s)
  • Pest
  • Invasive Species
  • Preferred Scientific Name
  • Phenacoccus manihoti
  • Preferred Common Name
  • cassava mealybug
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Cassava mealybug spread across the width of Africa in a period of 16 years. Its accidental introduction damaged a staple crop that is particularly important in times of drought, during a time of drought, leading to famine (...

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Pictures

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PictureTitleCaptionCopyright
Phenacoccus manihoti (cassava mealybug); adults on cassava.
TitleAdults
CaptionPhenacoccus manihoti (cassava mealybug); adults on cassava.
Copyright©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Phenacoccus manihoti (cassava mealybug); adults on cassava.
AdultsPhenacoccus manihoti (cassava mealybug); adults on cassava.©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Phenacoccus manihoti (cassava mealybug); adult females on cassava leaves.
TitleFemales
CaptionPhenacoccus manihoti (cassava mealybug); adult females on cassava leaves.
Copyright©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); adult females on cassava leaves.
FemalesPhenacoccus manihoti (cassava mealybug); adult females on cassava leaves.©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); eggs/woolly ovisacs.
TitleEggs
CaptionPhenacoccus manihoti (cassava mealybug); eggs/woolly ovisacs.
Copyright©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); eggs/woolly ovisacs.
EggsPhenacoccus manihoti (cassava mealybug); eggs/woolly ovisacs.©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); second, third and pre-ovipositing fourth instars feeding on cassava.
TitleSecond, third and fourth instars
CaptionPhenacoccus manihoti (cassava mealybug); second, third and pre-ovipositing fourth instars feeding on cassava.
Copyright©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Phenacoccus manihoti (cassava mealybug); second, third and pre-ovipositing fourth instars feeding on cassava.
Second, third and fourth instarsPhenacoccus manihoti (cassava mealybug); second, third and pre-ovipositing fourth instars feeding on cassava.©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Phenacoccus manihoti (cassava mealybug); cassava stem (planting material) distorted by mealybug infestation.
TitleDamage symptoms
CaptionPhenacoccus manihoti (cassava mealybug); cassava stem (planting material) distorted by mealybug infestation.
Copyright©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); cassava stem (planting material) distorted by mealybug infestation.
Damage symptomsPhenacoccus manihoti (cassava mealybug); cassava stem (planting material) distorted by mealybug infestation.©International Institute of Tropical Agriculture (IITA)
Epidinocarsis lopezi; adult, a natural enemy of the cassava mealybug.
TitleNatural enemy
CaptionEpidinocarsis lopezi; adult, a natural enemy of the cassava mealybug.
Copyright©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Epidinocarsis lopezi; adult, a natural enemy of the cassava mealybug.
Natural enemyEpidinocarsis lopezi; adult, a natural enemy of the cassava mealybug.©Georg Goergen/International Institute of Tropical Agriculture (IITA)/Insect Museum, Cotonou, Benin
Phenacoccus manihoti (cassava mealybug); 'Cassava trees' used at IITA to rear the parasitoid Epidinocarsis lopezi, the main natural enemy of P. manihoti.
Title'Cassava trees' used to rear parasitoids.
CaptionPhenacoccus manihoti (cassava mealybug); 'Cassava trees' used at IITA to rear the parasitoid Epidinocarsis lopezi, the main natural enemy of P. manihoti.
Copyright©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); 'Cassava trees' used at IITA to rear the parasitoid Epidinocarsis lopezi, the main natural enemy of P. manihoti.
'Cassava trees' used to rear parasitoids.Phenacoccus manihoti (cassava mealybug); 'Cassava trees' used at IITA to rear the parasitoid Epidinocarsis lopezi, the main natural enemy of P. manihoti.©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); adult of Prochiloneurus insolitus, an indigenous hyperparasitoid of P. manihoti.
TitleHyperparasitoid
CaptionPhenacoccus manihoti (cassava mealybug); adult of Prochiloneurus insolitus, an indigenous hyperparasitoid of P. manihoti.
Copyright©International Institute of Tropical Agriculture (IITA)
Phenacoccus manihoti (cassava mealybug); adult of Prochiloneurus insolitus, an indigenous hyperparasitoid of P. manihoti.
HyperparasitoidPhenacoccus manihoti (cassava mealybug); adult of Prochiloneurus insolitus, an indigenous hyperparasitoid of P. manihoti.©International Institute of Tropical Agriculture (IITA)

Identity

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Preferred Scientific Name

  • Phenacoccus manihoti Matile-Ferrero, 1977

Preferred Common Name

  • cassava mealybug

International Common Names

  • Spanish: chinche harinosa de la yuca
  • French: cochenille farineuse du manioc
  • Portuguese: cochonilha da mandioca

Local Common Names

  • Germany: Maniok-Schmierlaus

EPPO code

  • PHENMA (Phenacoccus manihoti)

Summary of Invasiveness

Top of page Cassava mealybug spread across the width of Africa in a period of 16 years. Its accidental introduction damaged a staple crop that is particularly important in times of drought, during a time of drought, leading to famine (Herren and Neuenschwander, 1991).

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

Top of page The cassava mealybug was first reported in 1973 from the Kinshasa (Congo Democratic Republic [Zaire]) and Brazzaville (Republic of Congo) areas of Africa (Hahn and Williams, 1973; Sylvestre, 1973). The hitherto unknown insect was subsequently described and named in 1977 as Phenacoccus manihoti (Matile-Ferrero, 1977), originating from the neotropics. In the neotropics, the insect was first discovered in Paraguay in 1981 by A.C. Bellotti of CIAT. P. manihoti had earlier been confused with another mealybug species found on cassava in Guyana and northern Brazil. Whilst exploration for the natural enemies of P. manihoti continued, this other mealybug was identified as P. herreni (Cox and Williams, 1981). The taxonomic clarification greatly enhanced successful introductions and establishment of specific natural enemies of P. manihoti in an Africa-wide classical biological control programme.

Description

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Eggs

Eggs are oblong, golden yellow and enclosed in woolly ovisacs located at the posterior end of the adult females. Length and breadth measurements are 0.30-0.75 mm and 0.15-0.30 mm, respectively (Matile-Ferrero, 1978; Nwanze, 1978).

Larva

Antennae are 6-segmented in first instars and 9-segmented in subsequent instars. Body length and breadth measurements are, respectively, 0.40-0.75 mm and 0.20-0.30 mm for first instars/crawlers; 1.00-1.10 mm and 0.50-0.65 mm for second instars; 1.10-1.50 mm and 0.50-0.60 mm for third instars; and 1.10-2.6 mm and 0.50-1.40 mm for fourth instars/newly emerged adults (Matile-Ferrero, 1978; Nwanze, 1978).

Adult

Adult females of the cassava mealybug are ovoid, rose-pink and dusted with white, powdery wax; the eyes are relatively prominent, legs are well developed and of equal size (Matile-Ferrero, 1978). The mealybug's body segmentation is apparent. Body segments bear very short lateral and caudal white wax filaments in the form of swellings that produce a toothed appearance to the body outline.

- antennae often 9-segmented, occasionally with 7 or 8 segments
- denticle usually present on claw - body generally with 18 pairs of cerarii
- quinquelocular pores usually present on venter - dorsal and cerarian setae lanceolate
- cerarii usually each with 2 lanceolate setae and no auxillary setae, except on anal lobes

All of the microscopic features listed above as typical for genus Phenacoccus are present in P. manihoti. Other important characters of P. manihoti are:

- underside of head with 32-68 quinquelocular pores immediately anterior to the clypeolabral shield
- circulus 'ox-yoke' shaped
- no translucent pores on hind tibiae

The above characters will facilitate recognition of many Phenacoccus species, especially the economically important ones. The lanceolate setae are especially distinctive for this genus. Regional keys to mealybug faunas, such as the one provided by Williams and Granara de Willink (1992), should, however, be used to support an identification of Phenacoccus, as some species have only a few of the morphological features which are typically found in this genus. A useful key to identify P. manihoti may be found in Williams and Granara de Willink (1992).

Distribution

Top of page P. manihoti is indigenous to South America, where it is found in Argentina, Bolivia, Brazil, Colombia, Guyana and Paraguay. It was accidentally introduced from South America to the Congo Republic in 1973 (Herren and Neuenschwander, 1991). It has spread in Africa to practically all countries where cassava is grown, in a broad belt from West through to East Africa and down to the eastern edge of South Africa. A distribution map is provided by CIE (1993).

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

CambodiaPresentIntroduced2010 Invasive Parsa et al., 2012; EPPO, 2014
IndonesiaRestricted distributionEPPO, 2014
-JavaPresentEPPO, 2014
LaosPresentEPPO, 2014
MalaysiaPresentDewi Sartiami et al., 2015
ThailandPresentIntroduced2008 Invasive Parsa et al., 2012; EPPO, 2014
VietnamPresentIntroduced2012 Invasive Parsa et al., 2012; EPPO, 2014

Africa

AngolaPresentIntroduced1975 Invasive Zeddies et al., 2001; EPPO, 2014
BeninPresentIntroduced1979 Invasive Akinlosotu and Leuschner, 1981; Zeddies et al., 2001; EPPO, 2014
BurundiAbsent, unreliable recordIntroduced1987 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; EPPO, 2014
CameroonRestricted distributionIntroduced1985 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; Zeddies et al., 2001
Central African RepublicPresentIntroduced1984 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; Zeddies et al., 2001; EPPO, 2014
CongoPresentIntroduced1973 Invasive Zeddies et al., 2001; EPPO, 2014
Congo Democratic RepublicPresentIntroduced1973 Invasive , ; , ; Hahn and Williams, 1973; Matile-Ferrero, 1978; Nwanze et al., 1979; Hennessey and Muaka, 1987; EPPO, 2014
Côte d'IvoirePresentIntroduced1985 Invasive Neuenschwander and Herren, 1988; Zeddies et al., 2001; EPPO, 2014
Equatorial GuineaRestricted distributionIntroduced1989 Invasive Neuenschwander and Herren, 1988; Zeddies et al., 2001; EPPO, 2014
GabonPresentIntroduced1976 Invasive Boussienguet, 1986; Zeddies et al., 2001; EPPO, 2014
GambiaPresentIntroduced1976 Invasive Herren and Neuenschwander, 1991; Neuenschwander, 2003; EPPO, 2014
GhanaPresentIntroduced1982 Invasive Korang-Amoakoh et al., 1987; Zeddies et al., 2001; EPPO, 2014
GuineaPresentIntroduced1986 Invasive Neuenschwander and Herren, 1988; Zeddies et al., 2001; EPPO, 2014
Guinea-BissauPresentIntroduced1982 Invasive Zeddies et al., 2001; EPPO, 2014
KenyaPresentIntroduced1989 Invasive Herren and Neuenschwander, 1991; Neuenschwander, 2003; IPPC-Secretariat, 2005; EPPO, 2014
LiberiaPresentIntroduced1990 Invasive Neuenschwander and Herren, 1988; Zeddies et al., 2001; EPPO, 2014
MadagascarRestricted distributionIntroduced Invasive
MalawiPresentIntroduced1985 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; Neuenschwander, 2003; EPPO, 2014
MaliPresentEPPO, 2014
MozambiquePresentIntroduced1986 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; EPPO, 2014
NigerPresentIntroduced1986 Invasive Zeddies et al., 2001
NigeriaPresentIntroduced1979 Invasive , ; Zeddies et al., 2001; EPPO, 2014
RwandaPresentIntroduced1984 Invasive Birandano, 1986; Herren and Neuenschwander, 1991; EPPO, 2014
SenegalPresentIntroduced1976 Invasive Herren and Neuenschwander, 1991; Neuenschwander, 2003; EPPO, 2014
Sierra LeonePresentIntroduced1982 Invasive James, 1987; James and Fofanah, 1992; Zeddies et al., 2001; EPPO, 2014
South AfricaRestricted distributionIntroduced Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991
SudanPresentIntroduced Invasive Neuenschwander and Herren, 1988; Neuenschwander, 2003; EPPO, 2014
TanzaniaPresentIntroduced1987 Invasive Neuenschwander and Herren, 1988; Herren and Neuenschwander, 1991; EPPO, 2014
-ZanzibarRestricted distributionIntroduced Invasive Neuenschwander and Herren, 1988
TogoPresentIntroduced1980 Invasive Neuenschwander and Herren, 1988; Zeddies et al., 2001; EPPO, 2014
UgandaPresentIntroduced1992Zeddies et al., 2001; EPPO, 2014
ZambiaPresentIntroduced1984 Invasive Neuenschwander and Herren, 1988; Chakupurakal et al., 1994; Zeddies et al., 2001; EPPO, 2014
ZimbabwePresentIntroduced Invasive Neuenschwander and Herren, 1988; Giga, 1994; EPPO, 2014

South America

ArgentinaPresentNative Not invasive de Santis, 1963
BoliviaPresentNative Not invasive Löhr and Varela, 1987; IIE, 1993; EPPO, 2014
BrazilRestricted distributionEPPO, 2014
-AmazonasPresentNative Not invasive Foldi, 1988; IIE, 1993; EPPO, 2014
-Mato Grosso do SulPresentNative Not invasive IIE, 1993; EPPO, 2014
-ParaPresentNative Not invasive Matile-Ferrero, 1977; EPPO, 2014
-ParanaPresentNative Not invasive Löhr and Varela, 1987
-Rio Grande do SulPresentNative Not invasive Foldi, 1988
ColombiaPresentNative Not invasive ,
GuyanaPresentEPPO, 2014
ParaguayPresentNative Not invasive Löhr and Varela, 1987; Löhr et al., 1990; EPPO, 2014

Risk of Introduction

Top of page P. manihoti poses a threat to other cassava-growing regions of the world, such as Indonesia.

Accidental introduction to new territories is possible through the movement of infested living cassava material for propogative purposes through shipping or air transport/mail.

It may be advisable for plant quarantine services to regulate trade in fresh planting material of cassava from Africa and tropical South American countries to other tropical countries. Trade may still be possible through a system of inspection of source areas and pre-export certification of shipments being free of infection. Planting material should be inspected in the growing season previous to shipment and be found free of infestation. A phytosanitary certificate should guarantee absence of the pest from consignments of planting material.

Habitat

Top of page In Africa, P. manihoti survives and occurs on cassava in all agroecosystems where it has spread.

Hosts/Species Affected

Top of page The cassava mealybug strongly prefers cassava and other Manihot species; the other host crops and wild hosts are only marginally infested. Talinum triangulare, Croton and Poinsettia species are particularly suitable for laboratory rearing and experiments.

Records from other plants are apparently accidental (Herren and Neuenschwander, 1991). Although it has been collected on plants in various families, such as citrus and tomato, there is no evidence that it can survive for more than one generation on plants other than Manihot and perhaps certain other Euphorbiaceae (Williams and Granara de Willink, 1992).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Boerhavia diffusa (red spiderling)NyctaginaceaeWild host
Capsicum spp.SolanaceaeOther
CitrusRutaceaeOther
Cyperus (flatsedge)CyperaceaeWild host
Euphorbia pulcherrima (poinsettia)EuphorbiaceaeWild host
Glycine max (soyabean)FabaceaeOther
Ipomoea batatas (sweet potato)ConvolvulaceaeOther
Manihot esculenta (cassava)EuphorbiaceaeMain
Manihot glaziovii (ceara rubber)EuphorbiaceaeWild host
Sida acuta (sida)MalvaceaeWild host
Solanum (nightshade)SolanaceaeOther
Solanum lycopersicum (tomato)SolanaceaeOther

Growth Stages

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

Symptoms

Top of page In cassava and on Manihot glaziovii, the pest causes stunting, leaf distortion and loss, dieback and weakening of stems used for crop propagation. The insect does not cause any significant damage to its other known host crops/plants which may only serve as a temporary support for 'drifting' populations of the insect that fall on them.

List of Symptoms/Signs

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Growing point

  • dieback
  • distortion

Leaves

  • abnormal colours
  • abnormal forms
  • abnormal leaf fall
  • honeydew or sooty mould
  • leaves rolled or folded
  • wilting
  • yellowed or dead

Roots

  • reduced root system

Stems

  • dieback
  • distortion
  • stunting or rosetting
  • witches broom

Whole plant

  • dwarfing
  • plant dead; dieback
  • seedling blight

Biology and Ecology

Top of page Herren and Neuenschwander (1991) reviewed the biology of cassava mealybug. The life cycle has been studied in the Congo by Fabres (1980) and by Fabres and Boussiengue (1981). Mealybug populations begin to build up in February, and there are nine generations. The largest generation is that during the dry season. Population numbers drop at the onset of the rainy season, when many mealybugs are washed off the plant. Within cassava fields, this mealybug occurs in a markedly aggregated distribution pattern, which differs between seasons.

P. manihoti reproduces by parthenogenetic oviparity. The life cycle consists of an egg and four instar stages with the fourth being the adult mealybug. Various laboratory experimental results (Nwanze et al., 1979; Iheagwam, 1981; Lema and Herren, 1985; Le Rü and Fabres, 1987; Schulthess et al., 1987) summarize that the insect has a lower thermal threshold of 14.7°C, an optimal temperature of about 28°C, no development above 35°C and a net reproductive rate of about 500 eggs in an average life span of 20 days. Egg incubation lasts approximately 8 days and the insect usually dies 1-3 days after it ceases egg production (Nwanze, 1978). The average development period of egg to adult lasts for about 33 days. The most favoured sites for oviposition are terminal shoot tips, lower leaf surfaces and leaf petioles. The eggs hatch into crawlers (= first instars) and the insect moults thrice in its development to a fourth instar (= adult mealybug). Except for crawlers, all instars prefer the lower surfaces of fully expanded leaves (Nwanze, 1978) from where they move sluggishly to the stems and shoot tips. At low population densities, therefore, the insect is most abundant in the shoot tips (Schulthess et al., 1987; Neuenschwander and Hammond, 1988).

The stunted, distorted cassava tips which result from feeding offer a protective environment for the mealybugs which feed on the sap (Calatayud and Le Rü, 1997). Crawlers move actively within the plant, usually to upper leaf surfaces from where they easily get blown away by wind. Movement in air currents and transportation of infested stem planting material by man are the main methods by which the insect is dispersed over long distances.

Ants attending mealybugs for their honeydew are known to defend the pests from natural enemies that would otherwise attack them. They have been observed interfering with biological control of cassava mealybug in Ghana (Cudjoe et al., 1993).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Allobaccha eclara Predator Adults/Nymphs
Anagyrus nyombae Parasite
Anagyrus pseudococci Parasite Gambia; Nigeria; Senegal cassava
Anagyrus pullus Parasite
Blaptostethus pallescens Predator Adults/Nymphs Rwanda cassava
Brumoides foudrasii Predator Adults/Nymphs Gambia; Nigeria cassava
Cacoxenus oligodous Predator
Cardiastethus exiguus Predator Adults/Nymphs Nigeria; Rwanda; Zaire cassava
Ceratochrysa antica Predator Adults/Nymphs Nigeria cassava
Cheilomenes lunata Predator Adults/Nymphs Zaire cassava
Cheilomenes propinqua Predator Adults/Nymphs Nigeria; Zaire cassava
Cheilomenes propinqua vicina Predator Adults/Nymphs Guinea Bissau cassava
Cheilomenes sulphurea Predator Adults/Nymphs Nigeria; Rwanda; Zaire cassava
Chilocorus angolensis Predator Adults/Nymphs Zaire cassava
Choristoneura freemani Predator Adults/Nymphs Nigeria; Zaire cassava
Clitostethus neuenschwanderi Predator Adults/Nymphs Congo; Zaire cassava
Coccodiplosis citri Predator Adults/Nymphs
Cryptoblabes gnidiella Predator Adults/Nymphs Nigeria cassava
Declivitata uncifera Predator Adults/Nymphs Zaire cassava
Dicrodiplosis manihoti Predator Adults/Nymphs Gambia; Ghana; Guinea Bissau; Nigeria; Zaire cassava
Diomus hennesseyi Predator Adults/Nymphs
Diomus neuenschwanderi Predator Adults/Nymphs Burundi; Central African Republic; Cote d'Ivoire; Gambia; Ghana; Guinea; Guinea-Bissau; Malawi; Mozambique; Nigeria; Rwanda; Senegal; Sierra Leone; Togo; Zaire; Zambia cassava
Domomyza perspicax Predator
Epidinocarsis diversicornis Parasite Africa cassava
Epidinocarsis lopezi Parasite Adults/Nymphs
Exochomus flavipes Predator Adults/Nymphs Gambia; Ghana; Guinea Bissau; Nigeria; Senegal; Togo; Zaire cassava
Exochomus flaviventris Predator Adults/Nymphs Congo
Exochomus promtus Predator Adults/Nymphs Ghana; Nigeria; Rwanda; Zaire cassava
Exochomus troberti Predator Adults/Nymphs
Exochomus ventralis Predator Adults/Nymphs Zaire cassava
Geocoris amabilis Predator Adults/Nymphs Nigeria; Senegal cassava
Hyperaspis aestimabilis Predator Adults/Nymphs Zaire; Zambia cassava
Hyperaspis delicatula Predator Adults/Nymphs
Hyperaspis jucunda Predator Adults/Nymphs Malawi
Hyperaspis notata Predator Adults/Nymphs Africa; Burundi; Central African Republic; Ghana; Guinea; Malawi; Mozambique; Sierra Leone; Zaire; Zambia cassava
Hyperaspis pumila Predator Adults/Nymphs Gambia; Guinea Bissau; Nigeria; Senegal; Togo cassava
Hyperaspis raynevali Predator Adults/Nymphs Congo
Hyperaspis raynevali Predator Adults/Nymphs Congo cassava
Hyperaspis senegalensis Predator Adults/Nymphs Congo; Gambia; Nigeria; Zaire cassava
Hyperaspis senegalensis hottentotta Predator Adults/Nymphs
Hyperaspis vinciquerrae Predator Adults/Nymphs
Isora circularis Zaire cassava
Mallada boninensis Predator Adults/Nymphs
Melanostoma annulipes Predator Adults/Nymphs Zaire cassava
Micraspis striata Predator Adults/Nymphs Zaire cassava
Neozygites fumosa Pathogen Adults/Nymphs Congo cassava
Nephus flavomaculatus Predator Adults/Nymphs Guinea Bissau; Nigeria; Zaire cassava
Nephus phenacoccophagus Predator Adults/Nymphs Gabon; Nigeria; Rwanda; Senegal; Zaire cassava
Nephus reunioni Predator Adults/Nymphs Zaire cassava
Nephus vetustus Predator Adults/Nymphs
Orius albidipennis Predator Adults/Nymphs Nigeria cassava
Parapyrus manihoti Parasite
Platynaspis capicola Predator Adults/Nymphs Zaire cassava
Platynaspis vittigera Predator Adults/Nymphs Zaire cassava
Ptyonocera atrifusella Nigeria cassava
Pyroderces hemizopha Predator Nigeria; Senegal; Togo; Zaire cassava
Rodolia occidentalis Predator Adults/Nymphs Ghana; Nigeria cassava
Scymnus kibonotensis Predator Adults/Nymphs Gambia; Nigeria; Rwanda; Zaire cassava
Scymnus levaillanti Predator Adults/Nymphs Nigeria; Zaire cassava
Scymnus quadrivittatus Predator Adults/Nymphs Nigeria cassava
Scymnus scapuliferus Predator Adults/Nymphs Nigeria cassava
Scymnus viduus Predator Adults/Nymphs Zaire cassava
Spalgis lemolea Predator Adults/Nymphs Ghana; Guinea Bissau; Nigeria; Rwanda; Zaire cassava
Sympherobius maculipennis Predator Adults/Nymphs
Xylocoris afer Predator Adults/Nymphs Nigeria cassava

Notes on Natural Enemies

Top of page In Africa, P. manihoti is attacked by the usual guild of polyphagous or oligophagous predators and parasitoids of mealybugs in Africa (Bartlett, 1978; Moore, 1988; Ben-Dov and German, 2003), which switched over to it as a new food source, and by indigenous entomopathogenic fungi, for example, Neozygites fumosa, which causes epizootics in the pest populations under warm and very humid conditions (Le Rü et al., 1985; Le Rü, 1986).

Prior to the introduction of Apoanagyrus lopezi (Hymenoptera: Encyrtidae) in Africa, lists of natural enemies of the pest were compiled for Congo (Matile-Ferrero, 1977; Fabres and Matile-Ferrero, 1980), Congo Democratic Republic (Nsiama Shè et al., 1984), Nigeria (Akinlosotu and Leuschner, 1981; Iheagwam., 1981) and Gabon (Boussienguet, 1986). Following the establishment of A. lopezi, the cassava mealybug food web was investigated over the whole continent and found to comprise about 130 species (Neuenschwander et al., 1987; Nsiama Shè, 1987; Biassangama et al., 1989). Many of the associated fauna are opportunistic and do not reproduce on the cassava mealybug; some are attracted more to the bunchy tops with the rich organic material from living and dead cassava mealybugs. Only about 20 species are common and seem to have some impact (Neuenschwander et al., 1987).

Wherever A. lopezi was established, it became the most important parasitoid and abundant natural enemy (Neuenschwander and Hammond, 1988; Hammond and Neuenschwander, 1990). Indigenous polyphagous hyperparasitoids adopt A. lopezi as an alternate host. The common hyperparasitoids in West Africa are Prochiloneurus insolitus and Chartocerus hyalipennis (Neuenschwander et al., 1987; Goergen and Neuenschwander, 1990; 1992; 1994). In Central Africa the hyperparasitoid P. aegyptiacus is usually most common (Fabres and Matile-Ferrero, 1980; Boussienguet, 1986; Neuenschwander et al., 1987; Biassangama et al., 1989). The most important predators are coccinellids, e.g., Hyperaspis spp., Exochomus sp., and Diomus sp., which usually occur at high densities of cassava mealybug (Fabres and Kiyindou, 1985; Boussienguet, 1986; Neuenschwander et al., 1987; Neuenschwander and Hammond, 1988; Stäubli Dreyer et al., 1997a,b). Results from olfactometer studies indicate that exotic and indigenous species of Diomus react more strongly to cassava mealybug, honeydew and exuviae than do indigenous Exochomus spp. (van den Meiracker et al., 1988).

Ants attending mealybugs for their honeydew are known to defend the pests from natural enemies that would otherwise attack them. They have been observed interfering with biological control of cassava mealybug in Ghana (Cudjoe et al., 1993).

Means of Movement and Dispersal

Top of page Natural dispersal

The dispersal stage of mealybugs is the first-instar crawler stage; these are often dispersed passively in the wind.

Vector transmission

Crawlers may also be carried passively by passing animals and people that brush past the host plant.

Agricultural practices

Harvesting infested plant material aids dispersal by scattering the crawlers into the air, where the wind may carry them away. Prunings of infested plants, and the clothing, tools and vehicles of agricultural workers can become contaminated with the crawlers and so aid in their dispersal.

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bark larvae No Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Flowers/Inflorescences/Cones/Calyx adults; eggs; larvae; nymphs Yes Yes Pest or symptoms usually visible to the naked eye
Fruits (inc. pods) adults; eggs; larvae; nymphs No Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants larvae No Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Leaves adults; eggs; larvae; nymphs No Yes Pest or symptoms usually visible to the naked eye
Roots larvae No Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Seedlings/Micropropagated plants adults; eggs; larvae; nymphs No 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 No Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bulbs/Tubers/Corms/Rhizomes
True seeds (inc. grain)
Wood

Wood Packaging

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Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood
Solid wood packing material with bark
Solid wood packing material without bark

Impact Summary

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CategoryImpact
Animal/plant collections None
Animal/plant products Negative
Biodiversity (generally) None
Crop production Negative
Environment (generally) None
Fisheries / aquaculture None
Forestry production None
Human health Negative
Livestock production None
Native fauna None
Native flora None
Rare/protected species None
Tourism None
Trade/international relations Negative
Transport/travel None

Impact

Top of page In 1973, P. manihoti was reported as an introduced arthropod species on cassava in Congo (Sylvestre, 1973; Matile-Ferrero, 1978) and Congo Democratic Republic (Hahn and Williams, 1973). Within a few years after these first reports, the insect became the major cassava pest and spread rapidly through most of the African cassava belt. By the end of 1986, for example, it had reached about 25 countries and covered 70% of the African cassava belt (Neuenschwander and Herren, 1988). In most countries the mealybug caused severe damage by stunting the growth points of cassava plants, sometimes totally defoliating the plants. Storage root yield losses of 84% have been reported (Nwanze, 1982). The pest-induced defoliation reduces availability of healthy leaves which are consumed as leafy vegetables in most of West and Central Africa. After the pest cripples plant growth, weed and erosion problems sometimes lead to total destruction of the crops. Additionally, pest-infested plants produce poor quality stem cuttings for use as planting material. The insect is more abundant and its damage severity is greater in the dry than in the wet season.

Examples of monetary values of damage are given by Norgaard (1988) and Neuenschwander (1990).

Zeddies et al. (2001) analysed the cost benefits of the biological control programme against cassava mealybug in Africa over a period of 40 years. Losses of cassava yield in the year of introduction were estimated at 80%; within 5 years, more tolerant varieties of cassava were cultivated and indigenous predators adapted to a new diet, so reducing annual losses to 20% (in rain forest) to 40% (in highlands and savanna).

Social Impact

Top of page The accidental introduction of P. manihoti to Africa damaged a staple crop that is particularly important in times of drought, during a time of drought, leading to famine (Herren and Neuenschwander, 1991).

Detection and Inspection

Top of page Colonies of this mealybug occur on the undersides of cassava leaves and on the shoot tips, and these will readily be seen during inspection. Minute crawlers, which may be present on plants before colonies are established, will only be detected by careful examination with the aid of a strong light and magnification. The plant tips are favoured feeding sites.

Similarities to Other Species/Conditions

Top of page P. manihoti is similar to P. madeirensis which also occurs on cassava. In P. madeirensis the body colour is greenish white and the ovisacs are much denser than those of P. manihoti. The presence of males in P. madeirensis is another distinguishing feature.

A very similar species, Phenacoccus herreni, is also common on cassava, but is yellow, produces males as well as females, and only occurs in South and Central America (Cox and Williams, 1981). Authoritative identification is difficult and should be done by an expert, using slide-mounted specimens. P. herreni may be identified using the key provided by Cox and Williams (1981) and Williams and Granara de Willink (1992).

Prevention and Control

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Biological Control

On the basis of the exotic origin and rapid spread of the cassava mealybug in Africa, classical biological control has been the main and most appropriate approach to the pest problem. Among several natural enemies introduced to combat the pest (Herren and Lema, 1982; Lema and Herren, 1985; Herren et al., 1987a; Neuenschwander and Zweigert, 1994), the solitary endophagous parasitoid Apoanagyrus lopezi, specific to P. manihoti, has been the most successful. Herren and Neuenschwander (1991) reviewed the biological control campaign against cassava mealybug in Africa. A. lopezi, collected from South America (Löhr and Varela, 1987; Löhr et al., 1988; Löhr et al., 1989; Löhr et al., 1990), has been the main natural enemy reared (Haug et al., 1987; Haug and Mégevand, 1989; Neuenschwander et al., 1989a, 1989b) and released across the cassava belt in Africa (Herren and Lema, 1982; Lema and Herren, 1985; Bird, 1987; Herren et al., 1987a,b). It was introduced to Nigeria in 1981 and is now established in at least 26 African countries (Ganga, 1984; Herren et al., 1987b; Korang-Amoakoh et al., 1987; Biassangama et al., 1988; Neuenschwander and Herren, 1988; Neuenschwander et al., 1989a, 1989b; Boussienguet et al., 1991; Hennessey et al., 1990; Herren and Neuenschwander, 1991; Neuenschwander and Zweigert, 1994). The biological and ecological impact of A. lopezi has been assessed in several laboratory and field experiments. In some studies, the results indicate a successful role of A. lopezi (Neuenschwander et al., 1986; Neuenschwander and Sullivan, 1987; Sullivan and Neuenschwander, 1988; Goergen and Neuenschwander, 1990; 1992; 1994; Cudjoe et al., 1992; 1993), whereas others are critical of reported success by A. lopezi (Fabres, 1981; Odebiyi and Bokonon-Ganta, 1986; Fabres et al., 1989; Iziquel and Le Rü, 1989; 1992; Le Rü et al., 1990; Souissi and Le Rü, 1997; 1998). Large-scale and sustained field studies have, however, recorded excellent biological control of the pest by A. lopezi (Neuenschwander and Madojemu, 1986; Hammond et al., 1987; Gutierrez et al., 1988a,b; Neuenschwander and Hammond, 1988; Neuenschwander and Gutierrez, 1989; Neuenschwander et al., 1989a, 1989b; van Alphen et al., 1989; Hammond and Neuenschwander, 1990; Neuenschwander et al., 1990; Gutierrez et al., 1993; Chakupurakal et al., 1994; Neuenschwander and Ajuonu, 1995; Neuenschwander, 1996). Ants attending mealybugs for their honeydew are known to defend the pests from natural enemies that would otherwise attack them. They have been observed interfering with biological control of cassava mealybug in Ghana (Cudjoe et al., 1993). It may be advisable to discourage ants in cassava fields if this becomes a problem. The economic impact of biological control of the cassava mealybug, mainly by A. lopezi, has been judged to be excellent (Norgaard, 1988a, b; Zeddies et al., 2001). Nominal costs of the biological control programme 1979-2013 were estimated at US$ 34.2 million, with the peak annual cost of the programme coming to US$ 5.2 million in 1985. The benefit to cost ratio of biological control by Apoanagyrus (Epidinocarsis) lopezi was calculated as at least 199:1. Where the soil is very infertile, however, biological control has been shown to be unsatisfactory, unless it can be complemented by cultural practices such as soil improvement (Neuenschwander et al., 1990; Le Rü et al., 1991; Schulthess et al., 1997) and host-plant resistance (Le Rü and Tertuliano, 1993; Tertuliano et al., 1993; Souissi and Le Rü, 1998). Biological control (particularly using the parasitoid Apoanagyrus lopezi) and the use of resistant varieties to control the pest are briefly described by Calatayud and Le Rü (1997). The coccinellid Hyperaspis notata is associated with the mealybugs P. manihoti and P. herreni on cassava in southern Brazil and the highlands of Colombia. It was brought to Africa to help control the accidentally introduced P. manihoti (Staubli-Dreyer et al., 1997). The parasitoids A. diversicornis, Allotropa sp., and the neuropteran predator Sympherobius maculipennis apparently failed to establish following their releases in Africa (Neuenschwander and Zweigert, 1994).

Organic Chemical Control

Immersion of cassava cuttings in manipueira (a liquid extract from cassava roots) for 60 minutes was found to significantly reduce infestation (Razafindrakoto et al., 1999). Mourier (1997) found that cassava leaves treated with a 1% neem kernel water extract (NKWE) were less attractive to first-instar cassava mealybug than untreated leaves, and those that started feeding died in the second instar. Three NKWE treatments at weekly intervals protected cassava against established early instar nymphs; however, some phytotoxicity was observed.

Host-Plant Resistence

Cassava contains two significant compounds whose levels increase in response to mealybug infestation. Cyanide content acts as a phagostimulant for the mealybug, whereas rutin has an antibiotic effect on the pest. It was found that the use of mulch and manure increased cassava resistance against mealybug infestation (Tertuliano et al., 1999). The use of resistant varieties to control the pest are briefly described by Calatayud and Le Rü (1997).

Cultural Control

Use of manure or other fertilizers can result in a reduction in the mealybug population because improved nutrition results in the production of larger parasitoid wasps with higher fertility levels (Schulthess et al., 1997). Mulch and fertilizer use also enhances the antibiotic properties of cassava against mealybug infestation (Tertuliano et al., 1999).

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