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Ann. Rev. Entomol. 1986. 31 :95-119

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INSECT PESTS OF COWPEAS

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L. E. N. lackai
by Universidade Estadual do Maranhao on 04/05/11. For personal use only.

International Institute of Tropical Agriculture, Oyo Road, PMB 5320, Ibadan, Nigeria

R. A. Daousr
Boyce Thompson Institute for Plant Research at Cornell University, Tower Road,
Ithaca, New York 14853

PERSPECTIVES AND OVERVIEW

Cowpeas (blackeye peas, or simply beans in many parts of Africa), Vigna
unguiculata, are widely grown in the tropics and subtropics for human as well
as for animal food. Nigeria, Brazil, and Niger are among the major producers
and account for over 70% ofthe world crop (137, 174). Nigeria alone produces
ca 900,000 tons annually(172). Grown mainlyas a secondary crop in associa­
tion with other staples such as maize, sorghum, millet, and cassava, cowpeas
constitute the cheapest source of dietary protein and energy for most poor
people in the tropical world (132). They are eaten as green seeds, green pods,
and dry grains, and tender leaves are used as a vegetable (101, 104, 132). In
addition, the haulm is fed to cattle in a number of countries (132).
Cowpeas originated in the savannah region of West and Central Africa (39)
and, like manyother crops with a long historyof cultivation, theyare subject to
heavylosses or entire crop failure (24, 85, 169, 189) as a result of severe insect
pest predation (e.g. 24, 25, 174, 175).
The pest spectrum is wide, and practically everypart of the cowpea plant has
an adopted pest species. While the pest status of the different insects may vary
from one country or region to another (169, 175), the losses reported suggest
that anyone major pest of cowpeas can cause substantial economic loss if left
uncontrolled (32, 169).

'Formerly stationed, by Boyce Thompson Institute, at the National Rice and Bean Research
Center (CNPAF), Goiiinia, Goilis, Brazil.

95
0066-4170/86/0 I 01-0095$02.00
 96 JACKAl & DAOUST

In traditional cropping systems, cowpea yields are consistently low, between

100 and 250 kg/ha. These low yields have been attributed for the most part to
insect pest depredation (24, 115, 168, 170, 175). This assertion is supported by
the dramatic increases in yield, sometimes up to tenfold, obtained with applica­
tion of insecticides (24, 104, 175, 186). Despite this yield potential, cowpea is
still considered a high-risk crop in many producing regions (168) and many
farmers are reluctant to invest in its production. Adoption of insecticide tech­
nology has been slow, mainly because of inconsistent supplies, soaring costs,
and the safety risks involved (96).
This situation, however, is rapidly changing because of the introduction of
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pest-resistant varieties and the greater availability of (often subsidized) in­

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secticides (95, 172). Cowpeas are increasingly grown as a monocrop in areas

where other crops fail as a result of drought.
Research on ways to improve cowpea production and utilization has received
considerable attention in recent years with the increased participation of in­
ternational agricultural research centers such as the International Institute of
Tropical Agriculture (UTA), national research programs [many of them sup­
ported by bilateral aid agreements such as the Bean/Cowpea Collaborative
Research Support Program (CRSP) sponsored by the USAID] , the International
Development Research Center (IDRC), the Canadian International Develop­
ment Center (CIDA), and many other international bodies.
This paper is an attempt to review the current status of research on cowpea
pests including the major pests, their biology and distribution, damage, control
methods, and the potential of integrated pest management (IPM) for the small
grower.

DISTRIBUTION, DAMAGE, AND PEST BIOLOGY

Lepidoptera
Lepidopterous species in several families cause serious damage to cowpeas.
Throughout tropical Africa, the legume pod borer, Maruca testulalis, is the
most important lepidopterous cowpea pest and causes severe damage (189). In
Kenya, for example, losses of up to 80% occur on indigenous cowpea varieties
as a result of pod borer attack (130); in Nigeria, losses have ranged from 30 to
86% even on high-yielding varieties (85, 87, 186).
The biology of M. testulalis has been extensively studied in several regions
of tropical Africa (8, 10, 89, 111, 130, 131, 174, 189). Adult moths usually
deposit their eggs on flower buds, although cowpea leaves, flowers, and
abscission scars also serve as oviposition sites (90, 98). Young larvae feed on
tender plant stems, terminal shoots, and peduncles during vegetative growth
and on flowers as plants mature (90,98, Il l , 173). Older pod borer larvae are
 COWPEA PESTS 97

highly mobile; feeding continuously on flowers and newly formed pods, they
cause severe damage throughout the reproductive cycle of the crop (10, 98,
130). In general, larval infestation has been found to be highest on flowers,
followed by flower buds, terminal shoots, and pods (89). M. testulalis larvae
emerge in the early evening from flowers, pods, stems, and the soil beneath
plants, and they wander on plant surfaces throughout the night (190). In the
early morning, larvae return to their shelters. Because of the strong photonega­
tive response of pod borer larvae, pods located within the leaf canopy and those
touching other plant parts are more heavily infested and are more severely
damaged (173, 190). When humidity is high, the infestation of pods by M.
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testulalis larvae may lead to an increased incidence of Choanephora pod rot

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disease (10). Upon reaching maturity, larvae drop from flowers or pods onto the
soil and pupate beneath the plant under leaf debris (98). The pupal period lasts
from 6 to 10 days (129).
The cowpea seed moth, Cydia ptychora, is another pod borer that attacks
cowpeas in parts of tropical Africa and India, but it is more restricted in its
distribution than M. testulalis (142). In southern Nigeria, Taylor (185) es­
timated that larvae damage between 20 and 30% of the pods during infestations.
The moth can complete its life cycle on a wide range of grain legumes, but
cowpea is its principal host plant (11). C. ptychora females have been shown to
oviposit on senescent sepals and, to a lesser extent, on the distal ends of
peduncles, flower petals, and pods (11), although some researchers have found
that the young or yellowing pods are the preferred oviposition sites (13 4, 185).
Upon hatching, larvae enter the pods and feed upon the seeds (185, 186), but
they can also attack foliar buds and young flowers (10, 134). Larval develop­
ment within the pod lasts between 11 and 14 days under field conditions, with
up to 10 larvae developing in each pod (185). Mature larvae leave pods to
pupate in the soil. After about 6 days, adults emerge, mate, and deposit eggs on
new plants in the early evening hours to complete the cycle (134).
Among stem borers, the lesser cornstalk borer, Elasmopalpus lignosellus, is
widespread in the Americas and attacks at least 62 species of plants in the
southern United States alone, including many legumes (179). In Brazil, E.
lignosellus is the most important lepidopterous species on cowpeas (47).'
Larvae bore into the cortical region of the plant at ground level or slightly
below; they tunnel up the stem of seedlings and cause wilting, yellowing of
foliage, and plant death (32, 35, 47). Generally, infestations do not exceed 5 to
10%, and 80% of the damage occurs during the first few weeks after plant
emergence (161). However, in dryer areas with sandy soils, yield losses caused
by a reduction in plant stand can be severe (32, 47). The biology of E.
lignosellus has been studied on many crops and, on cowpeas, the life cycle and
parasite complex have been investigated (107).
Since most lepidopterous species are polyphagous, numerous sporadic, but
 98 JACKAl & DAOUST

devastating, attacks by caterpillars occur on cowpeas in various locations. For

example, the limabean pod borer, Etiella zinckenella, attacks cowpeas in Asia,
Egypt, and the southern United States and occasionally causes severe damage
(80, 174). The velvetbean caterpillar, Anticarsia gemmatalis, can constitute up
to 88.5% of the lepidopterous defoliators on cowpeas in the southern United
States (123). Anticarsia irrorata and Plusia signata attack cowpeas in Ghana,
but neither species is a major pest (4). In contras.t, Amsacta moorei, the hairy
caterpillar, causes extensive damage to cowpea seedlings and is considered the
most important cowpea pest in Senegal (1 22) Heliothis and Spodoptera species
.

are sporadic cowpea pests throughout the Americas and tropical Africa (4, 35,
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47, 103a, 173). In Nigeria, larvae of Heliothis armigera bore into cowpea pods
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but remain outside the pod while eating whole seeds (10). In the United States
and several Latin American countries, H. zea adults deposit eggs on cowpea
foliage during the flowering stage, and young larvae, upon hatching, feed for a
short time on leaves and then bore into pods (32,47). Spodoptera littoralis is a
serious cowpea pest in Egypt and in parts of East Africa (103a, 174). In Brazil,
S. frugiperda, S. latifascia, Agrotis ipsilon, Urbanus proteus, and Hedylepta
indicata occur as sporadic cowpea pes ts (47).

Coleoptera
A diverse complex of coleopterous pests attack cowpeas in many regions of the
world. Tn the southern United States, Brazil, and several Caribbean islands, the
cowpea curculio, Chalcodermus aeneus, is the most important pest species (31,
32,34,42,61, 161). The pest occurs on cowpeas in the southern Atlantic and
Gulf States from Maryland to Texas. In Georgia alone, it has caused economic
losses in excess of one million dollars annually (32). In northeast Brazil where
most of the cowpeas are grown (900,000 hectares annually), C. aeneus is
widespread and destructive (161).
Adult curculios rest at the base of plants during the day and at night emerge to
feed on the developing pods in the "greensnap" stage (31, 33). They then
puncture pods through the hull with their rostrums (15, 32) and continue to
excavate the developing pea onto which eggs are deposited. As grubs develop,
they consume one or more seeds before completing development (32). Pre­
pupae then drop from pods onto the soil beneath plants, penetrate the soil to a
depth of between 1 and 6 em, and pupate (21). The entire cycle from egg
deposition to adult emergence lasts an average of 30 days, depending upon
temperature and location (15). When green pods are not available in the field,
adult curculios feed on cowpea foliage and stems (31). Females are generally
more active than males and move from pod to :-Jvd in response to available
oviposition sites (33). Feeding scars on pods, in addition to causing direct
damage, provide entry points for the pod rot fungus, Choanephora sp. (42, 43).
Economic losses resulting from curculio damage have not been assessed in
Brazil, but it is known that, in addition to direct losses in seed weight, the seed
 COWPEA PESTS 99

quality and germination capacity is significantly reduced (196). In the United

States, one to two stings and one grub in a can of processed peas result in
condemnation of the entire batch (32, 61).
Other Curculionidae attacking cowpeas in Brazil include the defoliator and
pod borer Pantomorus glaucus (29, 161), Aracanthus sp., and Promecops sp.
(30, 53), but these species normally are minor pests.
The only true weevil causing significant damage to cowpeas in Africa is the
apionid Piezotrachelus varium (122, 186). In northern Nigeria, the pest is
common and has been known to damage up to 92% of the seeds in some
locations at certain times; generally, however, only 13 to 26% of the pods are
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damaged and 3 to 4 seeds per pod are lost in heavy infestations (24, 1 86). The
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weevil is particularly severe in Senegal where populations last throughout the

year (122).
The pest status and biology of P. varium h ave been reviewed by Singh & van
Emden (174). Shiny black adult females bore holes in fresh green cowpea pods
using their long slender rostrums (173). Eggs are deposited in small packets into
the holes; larvae feeding on the seeds consume several seeds before completing
their development (122). Pupation occurs within pods, and adults emerge
through holes that they cut in the pod wall. The entire life cycle is generally
completed in 3 to 4 weeks (24, 122).
The cowpea weevil, or the cowpea seed beetle as it is more accurately
referred to, Callosobruchus maculatus, is the principal post-harvest pest of
cowpeas. This bruchid species occurs wherever the crop is grown and frequent­
ly infests up to 100% of the stored seeds within 3 to 5 months under ordinary
storage conditions (25, 28, 152, 166, 168, 178). In Egypt, losses in seed weight
commonly reach 50% after only 3 months of storage (84). In Nigeria alone,
over $30 million per annum is lost as a result of cowpea weevil damage (172).
The damage pattern caused by C. maculatus to stored cowpeas was studied in
northern Nigeria in an open market survey over an 8-yr period (27). This study
showed that each larva consumed an average of 10% of one seed and that an
estimated 30,000 tons was lost to bruchids annually (27). Since the reduction in
seed weight due to bruchids is directly proportional to the number of holes or
"windows" produced in seeds, seed losses can be easily predicted for different
levels of attack by C. maculatus (55, 158, 159, 172). However, economic
losses that are due to a direct reduction in seed quality and lowered seed
germination may be even more important. This was shown in an economic
evaluation made in Ceara, Brazil, in which cowpea seeds with 5% bruchid
damage were devalued by 53% in an open market (19). Losses in seed germina­
tion due to bruchid attack may reach 100% for grains with four holes per seed
(158).
The biology and life cycle of the cowpea seed beetle has been extensively
studied in Latin America (158), Asia (148, 164, 177, 199), and other regions
(25, 76, 178, 198). Seed beetles attack cowpeas both in the field (25, 178) and
 100 JACKAl & DAOUST

in storage by means of a crowding-induced adult polymorphism (117, 192).

The "active" or flying morph is specialized for dispersing from crowded
populations in storage and ovipositing on pods and seeds in the field. Newly
harvested seeds are usually infested at a low level (25), but the high fecundity
and short generation time of the sedentary "normal" morph promotes rapid
population buildup during storage. This is particularly true if access to seeds has
been increased by threshing (25, 135) or if pods shatter under dry conditions
(27). In storage, C. maculatus females attach eggs individually to the exterior
surface of seeds, but they do not oviposit on broken or damaged seeds or flour
derived from cowpeas or other legumes (18). Hatching larvae penetrate grains
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and complete development within the seeds (135, 152, 178). Both temperature
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and relative humidity greatly affect bruchid development (76).

Callosobruchus rhodesianus and Bruchidius atrolineatus also infest cow­
peas under field conditions. Generally they develop poorly on stored grains (25,
83), although the latter species has been reported as an important storage pest of
cowpeas in northern Nigeria (145). The bean weevil, �canthoscelides obtectus,
is a secondary pest of cowpeas in Mexico (97).
Chrysomelidae are by far the most important cowpea defoliators. The most
damaging species in Africa, Ootheca mutabilis, causes extensive defoliation
throughout Nigeria, Ghana (4, 128, 173, 186), and East Africa (103a). Adult
beetles attack seedlings and consume the palisade layer of the leaf between the
veins (143, 173). High beetle populations can cause complete defoliation, but
older plants escape serious damage (145). In addition to being an important
defoliator, O. mutabilis is an efficient vector of the southern bean mosaic and
cowpea mottle viruses of cowpeas (12). Medythia quaterna also transmits the
cowpea mottle virus to cowpeas, but with much less efficiency than O. mutabil­
is (12). In India, the chrysomelid Madurasia obscurella has been reported as a
major cowpea pest (79).
In Central America, Diabrotica balteata, D. adelpha, and Cerotoma ruficor­
nis are the predominant chrysomelid species on grain legumes, while D.
speciosa, Cerotoma sp., and Andrector arcuatus are the most important leaf
beetle species attacking cowpeas and beans in Brazil (47). In drybeans in Costa
Rica, yield losses resulting from chrysomelids reached 60% during the first 15
days of plant growth (77). In northern Brazil, cowpea losses probably reach
100% at certain times (47). As adults, these species cause severe defoliation to
plants, and as larvae, they consume stems, roots, and nodules (40, 47). In
addition to the direct damage to plants, chrysomelid beetles are also highly
efficient vectors of the cowpea mosaic virus throughout Latin America (40,
193, 194). The biology and habits of diabroticite beetles on cowpeas in Latin
America are not fully known.
Among Coleoptera in other families, meloid beetles, Mylabris sp. and
Coryna hermanniae, have been reported as serious flower pests and occasional
 COWPEA PESTS 101

pod pests in Nigeria and elsewhere in Africa, but attack is generally sporadic
(l0, 103a, 175). A lagriid beetle, Lagria villosa, causes defoliation and
sporadic damage to flower buds, flowers, and green pods in Ghana (4, 173).
The Mexican bean beetle, Epilachna varivestis, feeds primarily on cowpea
foliage during the larval and adult stages and is a vector of viral diseases in the
southern United States (32).

Homoptera and Hemiptera

Throughout equatorial Africa, pod sucking bugs (PSBs) cause serious damage
to cowpeas (4, 10, 24, 85, 112, 186). In Kenya and Nigeria, the coreids
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Clavigralla tomentosicollis and Riptortus dentipes are the most important

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species ( l 0, 94, 103a, 104). Nymphs and adults attack young, tender pods and
cause them to become shrivelled and the seeds to become malformed (6, 10,
94). Singh & van Emden (174) have reviewed the distribution and life cycles of
some coreids occurring on tropical legumes, and others (6, 7, 112, 128) have
studied their biology extensively. Studies at lITA in Nigeria showed that two or
more pairs of C. tomentosicollis adults per 10 plants caused economic yield
reductions (86).
The pod sucking Pentatomidae are of greater economic importance in the
United States (123) than in other locations. In South Carolina, an average of
three adult southern green stink bugs (SGSB), Nezara viridula, per plant caused
yield losses of up to 100% (162). Similar results were obtained in Georgia
(125). It has been shown that one SGSB adult is capable of damaging up to 59
cowpea seeds (124). Even at low population levels, the stink bug Euschistus sp.
and the leaf-footed bug Leptoglossus phyllopus are also capable of causing
distorted pods, blemished peas, pod abscission, and seed abortion (35, 162).
In Latin America, Piezodorus guildinii and Acrosternum sp. cause severe
damage to cowpeas in the highland regions of Peru and in the humid tropical
regions of north Brazil (47). The coreid Crinocerus sanctus is the most impor­
tant pod bug in Brazil. This species inflicts severe damage to cowpeas by
sucking the sap from stems, foliage, and pods (47).
In the family Miridae, Lygus hesperus is generally considered the most
important cowpea and bean pest in California (26, 120). These bugs feed on
meristematic tissues, flower buds, and developing seeds of grain legumes
(120). Crop losses result from the shedding of buds and flowers, the pitting of
seeds, and a reduction in the number of seeds per pod (35, 120).
Leafhoppers in the genus Empoasca are among the most important piercing
and sucking insects in the drier parts of tropical and subtropical regions where
pulses are produced. Nymphs and adults damage cowpeas by sucking juices
from foliage. The severely attacked crop shows a marked curling or rolling of
foliage and a rough appearance on the yellowing leaf tissue (150). Loss of vigor
leads to stunted growth, a lowered number of pods per plant, and, ultimately,
 102 JACKAl & DAOUST

plant death (4). Leafhoppers have not been implicated as vectors in the
transmission of any serious cowpea disease (138, 161).
In Latin America, the green leafhopper, Empoasca kraemeri. is among the
principal pest species on both dry beans (149) and cowpeas (161). The pest
occurs throughout the semi-arid "Sertao" of northeastern Brazil and causes
production losses in cowpeas of 60% or more (161). The seedling stage is the
most susceptible period of plant growth (52). If the crop is protected during this
period, dramatic increases in productivity can be obtained (from 498 kg/ha to
over 1200 kg/ha) (52). Yield losses due to E. kraemeri are highly correlated to
pest density (54).
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In Ghana, Empoasca dolichi is the dominant leafhopper species present on

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cowpeas, and this species constitutes 99% of the leafhopper population in

southwestern Nigeria (4, 137, 139). E. dolichi was formerly considered to be a
minor pest in Nigeria, but more recent observations showed that during the drier
months this species is capable of causing serious damage (173). Yield losses of
up to 40% have been reported in susceptible cultivars during the preflowering
period as a result of this pest (150). Older plants attacked by high leafhopper
populations (60 to 90 adults/plant) did not suffer losses in seed yield (138).
Other leafhopper species in Nigeria include Empoasca christiani, E. knudseni.
and E. pikna (173); in India, E. kerri is more common (79).
The black cowpea aphid, Aphis craccivora. is also a widespread pest on
cowpeas and causes significant damage in India, the Philippines, Thailand, the
southern United States, and throughout much of tropical Africa and Latin
America (32, 56, 79, 102, 160, 166). In Nigeria, aphids are important on
cowpeas grown in the dry season (the principal planting season) under irriga­
tion, and they have also been reported in outbreaks resulting from heavy
pesticide application (59, 114, 145). In several tropical regions, aphids are
more important as agents in the transmission of viral diseases of cowpeas than
as direct plant feeders (32, 145, 166), even though a heavy attack on young
seedlings can cause death of the plant (35, 160). Aphids feed on stems, terminal
shoots, and petioles of seedlings and, as plants mature, they move to pods and
flowers. Heavy feeding causes the stunting of plants and delay in the initiation
of flowering.

Thysanoptera
Thrips are among the most widespread and important pests of cowpeas in
tropical Africa. The cowpea flower thrips, Megalurothrips sjostedti, occurs
throughout tropical Africa and causes yield losses of up to 100% in Tanzania,
Ghana, Cameroon, and Nigeria (4, 68, 143, 173, 184). During the preflower­
ing period, M. sjostedti nymphs and adults may damage the terminal leaf buds
and bracts/stipules, causing the latter to become deformed with a brownish­
yellow mottled appearance (68). However, the principal point of plant attack is
 COWPEA PESTS 103

on the flower buds and, later, on the flowers themselves (173). Attacked flower
buds become brown and eventually abort (166), leaving behind dark red scars
(10). Flower damage is characterized by a distortion, malformation, and
discoloration of floral parts (173). Flower thrips populations are higher during
the dry season, which favors rapid multiplication of thrips (4, 68).
Other thrips species also occur on cowpeas in Africa, including the legume
foliage thrips, Sericothrips occipitalis (68, 173). This species normally feeds
and develops on leaves and foliage buds. In India, Megalurothrips distalis has
been reported (79), while Thrips palami is known to occur in the Philippines
(109). Frankliniella tritici is the most abundant species of foliage thrips in the
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southern USA (32, 123).

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Diptera
The bean fly, Ophiomyia phaseoli (syn. Melanagromyza vignicollis), is the
most important member of the Diptera that attack cowpeas (80, 175, 182). In
Southeast Asia this species is capable of causing significant yield reductions
(175). Plants are attacked within a week to 10 days following germination (3).
Eggs are deposited on the lower leaf surface. These hatch after 3-4 days, and
larvae bore into the slender stem tissues and form a tunnel in the pith or cortical
tissues (3). This behavior disrupts the normal transport of water and nutrients
and results in seedling mortality.
The leafminer Liriomyza trifolii has been reported to cause almost near
collapse of cowpea crops in Tanzania (144, 163). This same species is also
known to attack cowpeas in the United States and in Central and South
America, but economic losses are seldom reported (32, 47).

PEST CONTROL
The degree of success achieved in cowpea production in most of the producing
regions is a function of the level of pest control. Among the options available
for cowpea insect pest control are insecticides (in many cases to the exclusion of
other methods), the use of cultural control methods, and host plant resistance
(HPR). Attempts at biological control have also been reported, but to a more
limited extent than the other methods, and little research has been conducted to
assess the real potential of this control strategy.

Chemical Control
Most cowpea growers in the tropics are small holders who generally do not use
insecticides on their crop (95). However, as farm sizes increase and as farmers
are educated regarding the benefits of insecticide usage, chemical control
strategies are being increasingly used (62, 96, 109, 146) against seedling,
flower, and pod pests, as well as against storage beetles.
 104 JACKAl & DAOUST

SEEDLING PESTS This group includes the bean fly, aphids, leafhoppers,
foliage beetles, and many others. Good control is easily obtained with either
systemic or contact insecticides such as endosulfan, dimethoate, monocro­
tophos, thiometon, phorate, and carbofuran (10, 169). Control of pests during
this period is essential because high pest populations can terminate plant growth
if unchecked.
The cowpea aphid is most easily controlled with pirimicarb (a highly selec­
tive aphicide), dimethoate, and thiometon (65, 87, 96). Other products may
also be used to control aphids, but many, including DDT and Permethrin, have
undesirable side effects (59, 145). Bean flies, leafhoppers, and foliage beetles
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can be adequately controlled with a wide range of products including monocro­

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tophos, carbofuran, dimethoate, most synthetic pyrethroids, endosulfan, car­

baryl, and methomyl (17, 57, 67, 109, 195). These products are generally
applied as foliar sprays, although granular formulations, e.g., carbofuran, are
also available.

FLOWER AND POD PESTS Pests in this group constitute the most important
insect species attacking cowpea and other legumes worldwide (174, 175, 189).
Control of these pests, among which Maruca testulalis, Megalurothrips sjos­
tedti, pod sucking bugs (PSBs), and the cowpea curculio are most important,
can be effectively obtained with many synthetic pyrethroids, dimethoate,
endosulfan, chlorpyrifos, or formulations that combine pyrethroids with other
products (4, 31, 32, 58, 87, 96, 115). While these insecticides have a broad
spectrum of activity, not all products in this group are effective against every
pest. For example, monocrotophos gives excellent control of thrips and PSBs,
but it has been found to be ineffective against M. testulalis at a dosage of 500
gal/ha (92, 170). Deltamethrin has also been used effectively to control PSBs,
but only when the application interval did not exceed ,8 days (74). However, ·
Clavigralla tomentosicollis was not effectively controlled by this synthetic
pyrethroid (176). In laboratory and greenhouse tests (L. E. N. lackai and T. M.
Ndlovu, unpublished), deltamethrin was shown to have poor ovicidal activity
against C. tomentosicollis eggs, but it gave good control of adult PSBs for up to
4 days after application. During extremely high PSB populations, other more
effective chemicals, including Cymbush Super® and Sherpa Plus®, should be
substituted for deltamethrin applications (87, 96).
Application of insecticides on cowpeas has been traditionally carried out
with conventional high volume, hydraulic, knapsack sprayers. However, other
application techniques, including the spinning disc and ultra-low-volume and
Electrodyn® sprayers (62, 67, 96, 147), are gradually replacing the knapsack
sprayer in arid zones, e.g. the African sahel, where water is scarce. While these
techniques have many advantages, such as absence of mixing and ease of
handling (62, 96, 147), their use, compared to use of the knapsack sprayer, has
 COWPEA PESTS 105

not resulted in increases in cowpea yields (87; M. A. Gowman et aI, un­

published).
In cases where chemical control has been used by cowpea producers, (and it
should be emphasized that there is a trend of increasing use among farmers), the
problem of pesticide residues on cowpea leaves, pods, and grains has become
an important consideration. A number of studies, conducted mainly in India,
have examined insecticide residues of numerous products; the results have
generally shown that by 7-10 days after insecticide application residues have
been below tolerable limits (16, 63, 151). Washing and cooking can further
reduce pesticide residues. The highest residue levels were recovered from
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leaves and stems of plants, rather than from the fruits (176). Since numerous
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factors can alter pesticide persistence profiles in plants or in soil, chemicals

with low mammalian toxicity are generally advocated for cowpea pest control
(62, 96).

STORAGE PESTS A wide range of products is available for the control of

storage beetles in the genus Callosobruchus. The most effective insecticides
include pirimiphos-methyl (Actellic®), cypermethrin (SH-1479®), carbon di­
sulfide, chlorpyrifos, and phosphine (1, 2, 28, 84, 174). Other chemicals are
also widely used, including methyl bromide, DDT, BHe, and malathion (e.g.
2, 20, 22). In many developing countries, these products are often sold in the
open market with little or no precautionary measures being taken.
In addition to chemical insecticides, a number of plant products (e.g. oils,
powders, ashes, and others) are commonly used, particularly at the village
level, to protect cowpeas from damage in storage (8, 140, 171, 174). Also,
black pepper, lemon oil, palm oil, soybean oil, citrus peels, and activated
kaolin and neem extracts have been shown to be effective on storage beetles
(41, 88, 140, 180, 181, 183, 187). Many of these plant derivatives appear to
have ovicidal and/or larvicidal properties and a number of them completely
inhibit oviposition on certain grains (88, 116, 171). These compounds are
generally much safer than chemical insecticides and their use should be further
exploited. However, research is needed on better methods of production and
storage of these natural protectants.

Host Plant Resistance (HPR)

Increased attention has been focused on HPR of cowpeas in recent years, and
this subject is well covered in a number of recent reviews (167, 170, 174, 176).
Effective levels of resistance have been identified for a number of foliage pests.
These include leafhoppers (Empoasca dolichi) for which the cultivars TVu 59,
123, 662, and VITA-3 have been found resistant. This work was first reported
from Nigeria (150, 166), but the same varieties have more recently been found
to be resistant to other Empoasca species in Brazil (51) and East Africa (99). In
 106 JACKAl & DAOUST

India, cowpea varieties developed at lITA have been reported to be resistant to

the local leatbopper Amrasca biguttula (156). It appears that resistance in many
cases has been mediated by antibiosis and/or tolerance (150, 168).
Several varieties have been identified with high levels of resistance to Aphis
craccivora in Nigeria, East Africa, Botswana, India, and other countries (37,
99, 166; A. K. Ansari, unpublished). Despite the availability of resistance
sources, aphids are still largely controlled by chemicals because of their role as
vectors of the cowpea aphid-borne mosaic virus. However, a number of
high-yielding varieties with aphid and virus resistance have now been de­
veloped at lITA and should be available to farmers in the near future (170,
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172).
by Universidade Estadual do Maranhao on 04/05/11. For personal use only.

Cowpea resistance to aphids is conferred by extremely high levels of anti­

biosis in some cultivars, e.g. TVu 310, 810 (110). The exact plant component
responsible for the observed antibiotic effect has not been identified, but
preliminary indications suggest that phenols and/or flavonoids may be involved
(110).
Cowpea resistance has also been reported for a number of other seedling
pests, e.g. leafminers (Liriomyza spp. ) (50) and the bean fly in Asia (87).
Resistance to flower and pod feeders has been more difficult to obtain, but
low to moderate levels of resistance have been found for the flower thrips M.
sjostedti, and these have been incorporated into high-yielding varieties (e.g.
TVx 3236), which are now widely grown in several African countries (85, 167,
168, 172). The basis for this resistance is not fully known, but antibiosis
(physical and chemical) and antixenosis may be involved (85, 87, 154; A. B.
Salifu, unpublished).
Among the lepidopterous pests, low levels of resistance have been reported
for M. testulalis (91, 95, 167, 168) and Cydia ptychora (69, 71, 142). The
mechanisms of resistance involved have also been shown to be a combination of
antibiosis and antixenosis (69, 141, 168). Other investigations are presently
being conducted at the International Center of Insect Physiology and Ecology
(ICIPE) in Kenya to determine the components responsible for the observed
levels of antibiosis (K. N. Saxena, personal communication).
The only coleopterous pod pest for which resistance has been identified is the
cowpea curculio, Chalcodermus aeneus (42, 44, 66, 123). This resistance
appears to be related to pod wall strength, pod-seed ratio (which is negatively
correlated with pod wall resistance) (44, 73), an antibiotic factor in the hull
(36), and a differential concentration of certain primary metabolites (34). The
presence of a high number of tannin sacs and greater secondary wall formation
have also been associated with pod resistance to this pest (66).
Little information exists about cowpea resistance to the PSB complex, owing
to our inadequate knowledge of the feeding behavior and damage partitioning
among pest species in this complex and owing to the absence of a reliable
screening methodology. However, recent work by several authors (6, 7, 64, 94)
 COWPEA PESTS 107

should facilitate the development of an adequate screening methodology.

Khaemba & Kamala (103) reported that the cultivar Ife Brown (rVu 3629)
suffered a lowered level of damage by Riptortus dentipes, but this was more
related to plant age than resistance per se. More recent findings showed that a
number of cowpea lines suffer reduced damage from C. tomentosicollis under
populations of more than 20 bugs per 2m-row length (85, 87). However,
resistance against PSBs in general needs further investigation.
Even though several bruchid species attack cowpeas in storage, Calloso­
bruchus maculatus has been the focus of the most intensive studies concerning
cowpea resistance (75, 82, 126, 127, 167, 168). A source of resistance, TVu
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2027 [identified at UTA (167, 168»), served as a base for several studies (85,
by Universidade Estadual do Maranhao on 04/05/11. For personal use only.

86), and it was found that this variety contained an elevated level of trypsin
inhibitor that caused antibiosis to the larvae (75). Other factors such as seed
texture, high protein content, and a pod wall factor have also been linked to
resistance of other cowpea varieties to C. maculatus (28, 126, 127, 197). New
high-yielding varieties have been developed incorporating the high levels of
resistance from TVu 2027 (87, 118, 172), but it is not known whether trypsin
inhibitor is the only factor involved in resistance in these lines. In addition, even
though cowpeas are not generally stored in pods, pod wall resistance should be
further investigated because it will reduce C. maculatus infestation in the field
(9, 28).
Screening techniques for the identification of resistance that can be in­
corporated into high-yielding varieties have been developed and extensively
used, both under field and laboratory or screenhouse conditions (33, 37, 45, 91,
170). Screening methods against leafhoppers, aphids, and the storage weevil
were among the first to be developed (l26, 127, 150, 166, 168, 170); those for
flower thrips and the legume pod borer have been described in more recent
publications (45, 91, 95, 168, 170). A field screening method has also been
described for PSBs (95, 170). In addition, a number of different approaches
have been used to screen cowpea for resistance to the cowpea curculio both in
the field and laboratory (33, 36, 42, 66, 73). These methods involve both
damage measurements and the use of chemical and physical plant properties.

Cultural Control
The advantages of cultural control have been stressed by several workers (5, 14,
72, 96, 188). This is probably the oldest control practice among cowpea
growers in Africa and elsewhere (132). Cowpeas are generally grown as a
companion crop with maize, cassava, sorghum, millet, and other crops. As a
result, studies on cultural control have tended to concentrate on mixed cropping
(96, 119). However, the literature is replete with contradictions with respect to
the response of a given pest to the same cropping system. For example, in some
of the most intensive studies, Amoako-Atta et al (14) reported reduced damage
by M. testulalis when cowpea was intercropped with maize or sorghum in
 108 JACKAl & DAOUST

Kenya. Similar findings were reported from Nigeria (72) and Brazil (5).
However, Matteson (114) found no effect of cropping system (cowpea +
sorghum and cowpea + maize) on a given species of insect in Nigeria. In
contrast, Ezueh & Taylor (72) indicated that simultaneous planting of cowpea
and maize increased M. testulalis infestation, and they suggested planting
cowpea 12 weeks after maize. In part, this divergence results from the use in
some instances of larval infestation in flowers as a basis for comparison (85,
114), whereas in other instances damage to pods is adopted as the criterion (14,
72). A similar disparity in results has occurred for PSBs. Matteson (114) and
Kayumbo et al (100, 102) reported an increase in the PSB populations in cereal
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+ cowpea intercrop in both northern Nigeria and Tanzania, but this effect was
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not observed in southern Nigeria (85). In this case, however, differences in

sampling techniques may have been responsible (100).
Under cassava + cowpea intercrop, lackai (93) observed no difference in M.
testulalis larval infestation in cowpea flowers but reported a reduction in
numbers of PSBs and flower thrips. All other studies on foliage pests (es­
pecially beetles) and on flower thrips have indicated a drastic reduction in
numbers of these pests in mixed cropping situations (14, 85, 96, 114, 153,
165). Increases in pest popUlation resulting from a cropping system have been
attributed in part to the provision of a more attractive oviposition substrate, as in
the case of Anoplocnemis curvipes (128, 168), or to the provision of an
alternative food source, as is the case for the meloid beetles (114, 168).
Companion cropping may also delay pest incidence on cowpea and thereby
reduce crop damage in precocious early maturing varieties, without reducing
overall pest populations over time (165). Natural enemy activity is also thought
to be higher under mixed crops, although not predictably higher (174).
The time of planting can adversely or positively affect the functioning of a
given cropping system (72, 121; L. E. N. lackai and W. N. O. Hammond,
unpublished). Ezueh (70) reported that planting date manipulation did not
reduce damage from M. testulalis and C. ptychora. Proper spacing and crop
orientation can further improve the efficiency of the intercrop (119).
Limited information exists on use of trap cropping and tillage systems to
control cowpea pests, but there are indications that some pest species can be
reduced with trap crops (93, 95) and with a no-till agricultural system in
combination with rice stubble (155). These areas are of potential practical
value, but they have not as yet been adequately explored.

Biological Control
Despite the vast amount of research that has been conducted on cowpea pests,
little attention has been given to control by natural enemies and microbial
agents. In addition, the information that does exist has not been successf�lly
integrated into pest control strategies.
 COWPEA PESTS 109

A number of parasites, predators, and microbial agents of potential im­

portance in cowpea pest suppression have been reported by various workers.
On hemipterous pod pests, high levels of parasitization have been reported in
Nigeria and Tanzania (113, 114), and aphid predation by coccinellid beetles
(e.g. Menochilus sexmaculatus and Coccinella reponda) has been observed in
other locations (47, 59, 157). Parasitization of lepidopterous and coleopterous
pests (4, 23, 47, 57, 60, 66, 81, 105, lO7, 122, 133, 182, 191), in addition to
that of the bean fly (4, 78, 182) and flower thrips (114), has also been reported.
Microbial agents may be potentially useful for cowpea pest suppression, (23,
46-49, 108, 136), but this area is still largely unexplored and needs increased
Annu. Rev. Entomol. 1986.31:95-119. Downloaded from www.annualreviews.org

attention in the future (47). A number of studies have also shown that biological
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control can be augmented by the presence of certain weed species (13, 168).
Most studies on biological control of cowpea pests have failed to quantify the
real impact of these agents, with a few exceptions (23, 46, 78, 106, 108, 113,
114, 133). Although descriptive reporting needs to be encouraged, there is also
need to emphasize that the mere presence of a natural enemy in a cowpea system
does not guarantee its potential value as a control agent. Nevertheless, in a
system in which insecticides are used to obtain high yields, care must be taken
to conserve the natural enemy complex that is now available. It is generally
accepted that insecticides are harmful to natural enemies of insect pests, and
specific studies along these lines have shown that synthetic pyrethroids and
DDT are harmful to a number of parasites and predators (38, 59). But in
addition to these studies, considerable circumstantial evidence exists regarding
the negative effects of chemicals on natural enemies; this subject needs further
investigation and validation (76, 93, 145). In this regard, entomopathogens
have an even greater potential, since many are less disrupted by insecticides.
The study of these agents constitutes an entirely new area of biological control
of cowpea pests for which only limited information is available. However,
foundation studies in Brazil (47-49) and Kenya (136) have shown a wide range
of potentially useful microbial agents that could be exploited for further integra­
tion into cowpea pest management programs.

INTEGRATED PEST MANAGEMENT (IPM)

Integrated Pest Management implies the effective use of as many control
measures as are compatible, in order to suppress pest populations below
damaging levels and optimize yields with minimum disruption of, or damage
to, the environment. Much discourse exists on the status of IPM in Africa and
elsewhere in the tropics (95, 114, 176), but little work has been done to
implement the ideas that are often expressed. The reason usually advanced for
this is that not enough information is available on the various components
necessary for the formulation of IPM packages. However, from the information
 110 JACKAl & DAOUST

introduced in this review it is clear that considerable knowledge already exists

concerning cowpea pest management systems, enough in fact to begin to
formulate sound IPM strategies. Modifications to IPM approaches can be made
later as our knowledge of the agroecosystem increases. It should be noted that
the IPM concept was originally initiated to counteract the disastrous effects of
excessive pesticide usage, and, therefore, any delays in developing sound IPM
strategies will result in an increased proliferation in insecticide usage.
Some initial efforts have been made to formulate IPM packages (e. g. 96), but
considerably more work is still needed. An illustration of an attempt to develop
an IPM strategy is given in Figure 1 in which a cowpea variety, TVx 3236
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(developed at lITA), was resistant to flower thrips but was susceptible to other
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important pests. This resulted in the development of an integrated approach

toward cowpea pest control. This system used a cowpea/cassava intercrop that
reduced the population of flower thrips and PSBs by as much as 50% (93). As a
result of varietal resistance to flower thrips and a decrease in overall pest
popUlations, a reduced number of chemical insecticide sprays (only three
sprays) was needed. The testing of a cowpea variety with resistance to Maruca
borers (the major problem in the cassava + cowpea intercrop) should be even
more suitable for this system. Other such possibilities could be included in
experimental IPM packages for on-farm evaluation.
IPM strategies should also be developed for use in monocropped cowpeas.
Host plant resistance, insecticides, biological control agents, and host evasion
techniques, e.g. planting date manipulation, also need to be consolidated into
practical IPM packages for this cropping system.
Several research priorities exist for continued progress toward regulating

A B c o

Variety
�
Cultural Control
WNN Host Plant Resistance
� Chemica I Control DAP-l('

}
Strategy

e.g. e.g.
TVx3236 Cowpea ICassava A,h;d,
LH, Foliage No Spray#1 - 10 -20
Beetles

Thrips - Yes No spray ---

(+B Effect ) (+8 Effect)

Maruca - No Spray#2 - 40-45

Pod Bugs_ No Spray#3 - 50-60

(+ B Effect) (+ 8 Effect) I

* Days After Planting

Figure 1 A schematic presentation of an integrated pest management model for growing cowpeas
in the southwestern part of Nigeria.
 COWPEA PESTS 111

cowpea pest populations. These include studies on economic damage

thresholds to facilitate decision making regarding control strategies, further
investigations of the role of natural enemies, especially microbial agents, and
studies on population dynamics and individual species behavior. Increased
knowledge of these components will undoubtedly contribute enormously to the
formulation of lasting solutions to cowpea insect pest problems.

ACKNOWLEDGMENTS
We gratefully acknowledge the International Institute of Tropical Agriculture
(IITA) in Nigeria, the Bean/Cowpea Collaborative Research Support Program
Annu. Rev. Entomol. 1986.31:95-119. Downloaded from www.annualreviews.org

(CRSP), and the National Rice and Bean Research Center (Centro Nacional de
by Universidade Estadual do Maranhao on 04/05/11. For personal use only.

Pesquisa de Arroz e Feijao-CNPAF) in Brazil for their commitment to and

support for the research being conducted on cowpeas by both authors. We
especially thank Drs. S. R. Singh of IITA and F. J. Messina of the Boyce
Thompson Institute for their critical review of the manuscript. Finally, the
assistance by our secretarial staffs is most gratefully acknowledged.

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