Annu. Rev. Entomol. 2002. 47:701–31
Copyright c ? 2002 by Annual Reviews. All rights reserved
BIOLOGY AND MANAGEMENT OF ECONOMICALLY
IMPORTANT LEPIDOPTERAN CEREAL STEM
BORERS IN AFRICA
1ARC-Plant Protection Research Institute, Private Bag X134, Pretoria 0001,
South Africa; e-mail: firstname.lastname@example.org
2International Centre of Insect Physiology and Ecology, P.O. Box 30772,
Nairobi, Kenya; e-mail: email@example.com; firstname.lastname@example.org
3CABI Bioscience UK Centre, Entomology Department, Imperial College at
Silwood Park, Ascot, Berkshire SL5 7PY United Kingdom; e-mail: email@example.com
maize, diapause, competition, pheromones, cultural control, biological
grown in Africa for human consumption. Of the various insect pests attacking cereal
crops in Africa, lepidopteran stem borers are by far the most injurious. All 21 eco-
nomically important stem borers of cultivated grasses in Africa are indigenous except
has recently been found in sugarcane in Mozambique. C. partellus is competitively
displacing indigenous stem borers in East and southern Africa. A parasitoid, Cotesia
a 32–55% decrease in stem borer densities. This article is an attempt to summarize
the status of knowledge about economically important cereal stem borers in Africa
with emphasis on their distribution, pest status and yield losses, diapause, natural ene-
mies, cultural control, host plant resistance, and biological control. Special attention is
given to Busseola fusca and C. partellus, the most important pests of maize and grain
Cereals (maize, sorghum, millet, rice) are extremely important crops
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 702
STEM BORERS AND THEIR DISTRIBUTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . 702
DIAPAUSE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 704
DISPLACEMENT OF NATIVE STEM
BORERS BY C. PARTELLUS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 706
INDIGENOUS NATURAL ENEMIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707
Parasitoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707
Predators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707
Nematodes and Pathogens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707
KFIR ET AL.
PEST STATUS AND YIELD LOSSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 708
MANAGEMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 709
Utilization of Synthetic Sex Pheromones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 709
Cultural Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 710
Intercropping and Habitat Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 711
Management of Crop Residues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 712
Manipulation of Sowing Dates and Densities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 714
Fertilizer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715
Host Plant Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715
Breeding for Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 716
Biological Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 716
is also an important cash crop grown in some areas. The crops are grown primarily
far the most injurious (224). Because of their economic importance, stem borers
have been the subjects of many studies, which resulted in an enormous amount
of literature. This review summarizes the status of knowledge about cereal stem
borers in Africa.
STEM BORERS AND THEIR DISTRIBUTION
Maes (105) listed 21 economically important lepidopteran stem borers of culti-
21 species, 7 are primarily pests of rice, and one mainly attacks pearl millet in the
Sahelian region. Among the noctuids, Busseola fusca and six Sesamia spp. are
considered economically important. Two pyralids are serious pests: the rice borer,
Maliarpha separatella, and Eldana saccharina, a pest of sugarcane and maize.
The largest group (12 species) of injurious stem borers are crambids, with the ma-
jority (7 species) belonging to the genus Chilo Zincken. Within specific crops and
geographic regions, fewer species are considered to be important pests (Table 1).
In South Africa, B. fusca and Chilo partellus are the only important stem
borers of maize and sorghum (94), while in sugarcane in the same region, only
E. saccharina is considered to be a serious pest (47). In East Africa, C. partellus,
Chilo orichalcociliellus, E. saccharina, B. fusca, and Sesamia calamistis are men-
tioned as important and widely distributed stem borers of maize and sorghum
(168). Major stem borers of maize and sorghum in West Africa include B. fusca,
most important cereal crop, Coniesta ignefusalis is the dominant stem borer (73).
AFRICAN STEM BORERS
distributions, and major cultivated host plants
Important stemborer species in Africa and the Indian Ocean Islands, their
Family SpeciesDistributionHost plantsReference
eastern and southern
coastal eastern, Malawi,
West and Central
Indian Ocean Islands;
maize, sorghum, rice
Information on rice stem borers is primarily from West Africa and the Indian
Ocean Islands, as these are areas where rice is an important food crop. Chilo
zacconius, M. separatella, and S. calamistis are considered the most economically
important stem borers of rice in West Africa (13,37). M. separatella is the only
rice borer that has a widespread distribution in sub-Saharan Africa, and it also
occurs in the Comoro Islands and Madagascar (37). Other rice stem borers of
and Sesamia nonagriodes botanephaga (37,38). Additionally, Chilo aleniellus is
reported to be a rice stem borer in Ivory Coast (189,190).
Two of the economically important cereal stem borers in Africa are introduced
species: C. partellus and C. sacchariphagus. C. partellus is an Asian species
(29) that invaded Africa sometime before 1930 when it was first recorded in
Malawi (186), but it was not reported again until some 20 years later in Tanzania
(53). The distribution of C. partellus now includes Ethiopia, Sudan, Somalia,
Kenya, Tanzania, Uganda, Mozambique, South Africa, Swaziland, Lesotho,
Zimbabwe, Zambia, Malawi, and Botswana (41,71,79,127). Using GIS tools,
Overholt et al. (145) predicted the eventual distribution of C. partellus in Africa
based on the climate at locations where it was known to occur and then extrap-
olating to other locations with similar climates. The prediction included several
countries in southwestern and West Africa where C. partellus is not yet known to
KFIR ET AL.
The other invasive species, C. sacchariphagus, is a serious pest of sugarcane
in the Indian Ocean Islands. C. sacchariphagus may have been accidentally in-
troduced into the islands either from Sri Lanka or Java with the introduction of
sugarcane around 1850 (222). There are recent confirmed reports of C. saccha-
riphagus attacking sugarcane in Mozambique (209).
others are found throughout sub-Saharan Africa. C. orichalcociliellus, a pest of
maize and sorghum, occurs in coastal East Africa, Malawi, and Madagascar at
altitudes below 600 m (51,108,127). C. aleniellus has been reported as a pest of
rice in West and Central Africa and more recently as an important pest of maize
in Ivory Coast (120). C. ignefusalis is the dominant stem borer of millet in the
Sahelian zone of West Africa (69).
grasses and sedges (23,45,61,78). In southern Africa, E. saccharina is a major
pest of sugarcane but rarely causes damage in maize (177). In contrast, E. sac-
charina is considered to be a pest of maize, sugarcane, and rice in West Africa
B. fusca and Sesamia spp. occur throughout subsaharan Africa (79). Of the
Sesamia spp., S. calamistis is the most widely distributed and economically im-
portant species, but several others, including S. cretica, which occurs in Somalia,
Sudan, and Ethiopia, and S. nonagrioides botanephaga, which is found in both
East and West Africa, are also important. As with E. saccharina, the pest status
of B. fusca varies by region. In East and southern Africa it is a pest at higher alti-
tudes (>600 m) (127,178), but in West Africa, B. fusca occurs from sea level to
>2000 m (187) but is primarily a pest in the dry savanna zone (68).
Many cereal stem borers have a resting period toward the end of the cropping
eral months in the dry season (130,161,163). However,S. calamistis was reported
not to enter diapause in Uganda (78) or in Nigeria (68). In West Africa, B. fusca
enters diapause during the dry season, and it takes up to six months to complete
development. With the initiation of the rains, the larvae pupate within the stems,
and 10–12 days later adult moths emerge (68). Similar observations were made
on C. ignefusalis, which has a facultative diapause in dry millet stems (226).
cold dry season (April–September), in the lower parts of the dry stalks, where they
are well protected from natural enemies and adverse climatic conditions (83–85).
B. fusca diapauses throughout its distribution in Africa, whereas C. partellus does
not diapause in the warm low-lying regions of the South African Provinces of
Kwazulu-Natal (203) and Mpumalanga (R. Kfir, unpublished data), Swaziland
AFRICAN STEM BORERS
(218), and southern Mozambique (27). In regions where there is an abundance of
host plants and the climate is warm, C. partellus normally develops continuously
all-year round. In other regions with long dry periods in winter or in summer, the
borer enters into a resting period. C. partellus was reported to diapause in the dry
season in India (187) and on several islands off the coast of Africa (39,51). How-
ever, populations without a resting period were reported from the Coast Province
lus found in the dry season on the Kenya Coast and at lower elevations in South
Africa. In the coastal area of Kenya, in periods between cropping seasons, some
stem borers diapause in maize stubble, whereas others remain actively feeding in
wild grasses, such as Napier grass, in the proximity of cultivated areas (124).
An increase in carbohydrates and decrease in protein and water of the food
plant are the main factors inducing diapause in B. fusca (193). Dry condition of
the host plant (127) and the general deterioration of the nutritive environment
(51,160) induced diapause in larvae of C. partellus even when climatic conditions
were favorable for development (161). Diapause could be artificially induced in
nondiapause larvae introduced into aged maize stems.
During diapause, larvae of B. fusca and C. partellus progressively decreased in
weight and had up to seven additional stationary molts. The longer time the larvae
remained in diapause, the lighter in weight the emerging adults were, with fewer
eggs and oocytes in their ovaries. After eight months in diapause, emerging adults
that emerged from nondiapausing larvae (86).
of time they were kept in the laboratory at 21◦C and 60% relative humidity. Lar-
vae of C. partellus collected in April–June emerged in November, whereas those
collected in July emerged in October and those collected in August emerged in
September. Regardless of the collection date, C. partellus started to emerge from
Kfir (85) concluded that B. fusca had an obligatory diapause and C. partellus a
facultative diapause. In the field, C. partellus starts to emerge from diapause in
the second part of August, and it continues until the first week of November, a
period of 12 weeks. In contrast, B. fusca pupates only during October–November
(83,85,95). This difference in the pattern of emergence from diapause by the two
lapping generations of C. partellus (85,205) occurring every year in South Africa.
Conditions of continuous moisture during the long rainy season in Kenya play
a significant role in the termination of diapause by B. fusca (135,137). Rain-
fall alone did not appear to be the main factor (192). Contact with water was a
more significant factor in diapause termination than water uptake (136). Applica-
tion of water or rainfall played an important role in promoting pupation during
postdiapause dormancy of B. fusca in Ethiopia (59) and Ivory Coast (119). De-
lay in wetting larvae after diapause and access to water early in diapause had a
KFIR ET AL.
deleterious effect on the larvae. The main factor enabling diapausing B. fusca
larvae to survive adverse conditions appears to be efficient water conservation
A combination of temperature and photoperiod played an important role in
termination of diapause in B. fusca in South Africa, and water was important
as a stimulus for morphogenesis following diapause (89). Long days accelerated
termination of diapause in C. partellus, but under 16-h daylight, termination of
diapause was faster than when under constant illumination. In contrast, tempera-
ture, relative humidity, and day length did not affect diapause of C. partellus and
C. orichalcociliellus in Kenya (161). C. partellus collected in South Africa at
25◦38?S, 27◦47?E (90) is more affected by day length than the Kenyan population
located near the equator. Kfir (89,90) suggested that the right combination of day
length and temperature could be used for breaking diapause to provide large num-
bers of insects for artificial infestations for plant-resistance trials. This requires
only simple facilities and can save the large expense of keeping a continuous
culture in the laboratory on artificial diet.
DISPLACEMENT OF NATIVE STEM
BORERS BY C. PARTELLUS
The invasive stem borer, C. partellus, has proved to be a highly competitive col-
onizer in many of the areas it has invaded in eastern and southern Africa, often
becoming the most injurious stem borer (93,163) and displacing native species
(92,141). In coastal Kenya, there is evidence that C. partellus has partially dis-
placed the indigenous stem borer, C. orichalcociliellus (130–132, 141). However,
C. orichalcociliellus continues to be found at a relatively high frequency (10–
30% of the borer complex), which suggests that the displacement of C. orichal-
cociliellus will not proceed to extirpation (228). Recent investigations have found
that C. orichalcociliellus completed development in two native grasses, in which
C. partellus could not develop (131), which may be one factor that allows their
ment of native stem borers in two other areas in Africa. In the Eastern Province
of Kenya, C. partellus was present in the early 1980s but was less abundant than
B. fusca (163). However, in the same area in the period 1996–1998, B. fusca was
rare and C. partellus was dominant (183). Similarly, in the eastern Highveld re-
gion of South Africa, C. partellus partially displaced B. fusca over a period of
seven years (92). The displacement was most evident in grain sorghum where the
proportion of C. partellus in the total stem borer population increased from about
3% in 1986 to 91% in 1992.
Several factors have been investigated that may be responsible for the competi-
a generation in less time than C. orichalcociliellus (101,131), which may result
AFRICAN STEM BORERS
in a higher population growth rate. Moreover, C. partellus terminates diapause
more rapidly than C. orichalcociliellus (130) or B. fusca (92), which may allow C.
partellus to colonize host plants before the two native species at the beginning of
growing seasons. Kfir (92) speculated that B. fusca avoids plants already infested
by C. partellus, using odors associated with host plant feeding. Ofomata (129)
showed that when equal numbers of C. partellus and C. orichalcociliellus infest
the same maize, sorghum, or wild-sorghum plant, more C. partellus successfully
more neonate C. partellus larvae dispersed from the plant where they hatched, and
they dispersed greater distances than C. orichalcociliellus, which may allow C.
partellus to colonize more plants than the native borer (129).
INDIGENOUS NATURAL ENEMIES
The indigenous parasitoids associated with African cereal stem borers have been
treated in Polaszek (150). All stem borer parasitoids belong to either the
Hymenoptera or Diptera. Their effectiveness in keeping stem borer populations
below acceptable thresholds has been doubted by several authors (93,142). How-
ever, no studies are yet available that examine the effect of the absence of par-
asitoids on stem borer populations. Recent investigations (R. Kfir, manuscript
submitted) into the effects of removal or partial removal of parasitoids from stem
There have been a number of studies of the effectiveness of indigenous predators
on several borer species in the past decade (30,31,93,103,162). Virtually all
studies have concluded that indigenous predators are not able to keep stem borer
populations below economic injury levels (31,162).
Nematodes and Pathogens
have been summarized recently (149). These authors also provided summaries for
the study and identification of these agents. In doing so, they also concluded that
Knowledge concerning viruses associated with African cereal stem borers has
been reviewed (128). Iridoviruses, granulosis viruses, cytoplasmic viruses, and
nuclear polyhedrosis viruses are known to attack and kill borer larvae in Africa,
but no detailed studies have been conducted.
KFIR ET AL.
PEST STATUS AND YIELD LOSSES
Feeding and stem tunneling by borer larvae on plants results in crop losses as a
consequence of destruction of the growing point, early leaf senescence, interfer-
ence with translocation of metabolites, and nutrients that result in malformation
of the grain, stem breakage, plant stunting, lodging, and direct damage to ears
(18,33,36,94). Infestations by stem borers increase the incidence and severity of
stalk rots (33).
In South Africa, estimated yield losses from B. fusca damage ranges between
10% and total loss (24,106,109,206). Yield loss in maize by B. fusca was signif-
icantly correlated with leaf damage, but a higher correlation was observed with
stem-boring damage (24). The estimated yield losses due to C. partellus in maize
and sorghum exceed 50% (153). A positive correlation between infestation level
populations using artificial infestation on sorghum indicated that C. partellus was
more injurious than B. fusca (200,201). More damage by C. partellus was ob-
served on long-season, grain-sorghum cultivars because of exposure over a longer
period in the susceptible preflowering stages (199).
In Lesotho, seasonal variation in yield loss due to B. fusca damage on maize
ranged between 0.4% and 36.6% (55). In Mozambique, larvae of third-generation
C. partellus, the most important stem borer, were reported to infest 87% of cobs of
late-planted maize and to severely damage 70% of grain (27). Infestations of up to
100% of plants, with considerable yield losses, have been recorded in the Maputo
and Gaza Provinces (125), the Limpopo Valley (27), and in southern Mozambique
In Zimbabwe, C. partellus caused yield loss of 50–60% in sorghum (178).
In maize, borer infestations range from 30% to 70% in fields of resource–poor
farmers but are less than 30% in commercial farms where insecticides are used
(177). In Tanzania (215) and Kenya (216), loss of about 12% maize grain for
every 10% plants infested by B. fusca were reported. In Kenya, 18% yield losses
were attributed to C. partellus and C. orichalcociliellus in maize (217) and 88%
in sorghum owing to C. partellus (165).
Maximum stalk damage in maize and up to 80% grain yield loss in sorghum
by C. partellus were observed in Kenya on 20-day-old crops, whereas similar
In Tanzania, Jepson (81) reported 40–100% sorghum plants infested by B. fusca,
whereas in Ethiopia movement of B. fusca larvae into the base of the sorghum
head resulted in undersized heads and grain loss of 15% (112).
AFRICAN STEM BORERS
of Sesamia sp. larvae and the extent of damage to maize stems, and a negative re-
lationship between damage to maize stems and maize yield were shown (64). The
calculated yield loss caused by Sesamia sp. to maize in the rain forest coastal, de-
rived, and Guinea ecological zones were 27%, 15%, 18%, and 14%, respectively
(64). In Burundi, insecticides and exclusion-cage trials indicated maize yield re-
ductions of 12–15% by E. saccharina and 30–50% by B. fusca (121). In Burkina
Faso and Niger, yield loss in sorghum by C. partellus and in millet by C. igne-
fusalis was estimated by using carbofuran to protect the crops and by infesting the
crops at different growth stages. The highest grain yield was obtained when the
crop was protected between 15 and 30 days after emergence; artificial infestation
at 15 days after emergence resulted in the greatest damage. The infestation in un-
protected sorghum plots was 60–62% (188). Stem borer control in sorghum in the
southern Guinea savanna of Nigeria, where S. calamistis predominates, improved
yields by 16–19% (1), whereas in the northern Guinea savanna, where B. fusca
and unsprayed sorghum in 22 farmers’ fields were studied in Nigeria, and a mean
yield loss of 21% was determined (11). Yield loss in sorghum depends on the time
of infestation (12). Sorghum infested by B. fusca before the booting stage suffered
greater yield losses (110). Proportion of internodes bored in the lower part of the
stalk had a more consistent negative correlation with grain yield than did propor-
tion of stem tunneled (110). A recent study in Cameroon found that stem borers,
and caused an 11% loss of plants owing to deadheart (43).
Utilization of Synthetic Sex Pheromones
Pheromone-baited traps are useful devices for monitoring moth population levels
of stem borers. Trap catches of male moths can provide useful information for
the timing of insecticide applications (154,208,210). Campion & Nesbitt (42)
reviewed the progress in the identification and the utilization of sex pheromones
for stem borer monitoring and concluded that mass trapping is unlikely to provide
satisfactory control but that mating disruption is more likely to be effective. Syn-
Chilo auricilius and C. zacconius have shown satisfactory attractiveness to male
moths in the field (25). Sex pheromones for B. fusca, C. partellus, S. calamistis,
(95,96) revealed that the first flight resulting from moths emerging from hiber-
nating borers peaked about the middle of November. The second flight, usually
larger, peaked in the second half of February; a third flight, which occurred in
KFIR ET AL.
Mpumalanga Province, peaked around the middle of April. No catches of moths
were recorded during winter (June–September). Larval peaks of B. fusca in the
field lagged about 4–6 weeks behind the corresponding moth flight peaks (95,96).
Omni-directional traps were superior to delta traps for quantitative and qualitative
estimation of B. fusca moth populations (95,96,207).
tions in the field are required before trapping C. partellus can be used to determine
economic threshold levels.
A slow-release pheromone formulation gave high levels of communication
disruption in B. fusca when applied at 40 g a.i. ha−1as 250 or 500 point sources
ha−1(49). This effect was observed to persist for at least 18 weeks and from
release rate studies was predicted to last for 6 months. In field trials in Kenya,
some reduction in damage levels was observed, indicating that mating disruption
had occurred (49).
Various methods of cultural control of stem borers in Africa have been reviewed
(102,113,164,166,196). It is the most relevant and economic method of stem
borer control available for resource-poor farmers in Africa. Other control options
cultivars are not widely available, and biological control of stem borers is only
partially successful. Cultural control is among the oldest traditional practices and
normally cannot be used as a tactical means of control (52). Cultural control is
considered the first line of defense against pests and includes techniques such as
dates, and tillage methods (196). The latter three agronomic practices can directly
affect crop yield (52). Many cultural control practices are labor intensive, but they
have little adverse effects on the environment and are readily available without
extra investment in equipment.
However, an understanding of stem borers’ behavior and the relationships with
their respective crops are important for the development of efficient management
strategies. The differences in the behavior of E. saccharina in South and eastern
Africa provide an example of the importance of understanding pest behavior in
making control decisions. In South Africa, larvae of E. saccharina mainly infest
lower parts of sugarcane stalks. It is recommended that farmers cut stalks low
and leave the tops, which are then cut off and left in the field. In East Africa, on
the other hand, recommendations are to not leave tops of plants in fields because
E. saccharina larvae largely occur in upper-plant parts, and these residues would
house the carryover population (196).
most African farmers have not adopted them (126). Cultural control is severely
constrained by the lack of management capabilities of farmers, especially in areas
where farming communities lack the support of an adequate extension service
AFRICAN STEM BORERS
Intercropping and Habitat Management
Intercropping or mixed cropping has been widely practiced for centuries by small-
scale farmers in Africa to reduce risk of crop failure, attain higher yields, and
improve soil fertility (155,196). Although some of these practices also lead to
suppression of cereal stem borer populations, no studies have shown that farmers
grow specific intercrops to exploit this effect. Many field studies have been con-
ducted in Africa during the past two decades in an effort to identify the best crop
combinations for reducing stem borer populations on cereal crops.
Studies in Kenya have concentrated both on the practice of intercropping cow-
be adopted by small-scale farmers in eastern Africa (16,138,148,162). Most con-
cluded that intercropping reduced the incidence of stem borers. In West Africa,
maize, millet, and sorghum intercropping were studied (3,57,82). Maize/cassava-
intercropping systems in Nigeria reduced by half larval numbers of E. saccharina,
B. fusca, and S. calamistis populations (82). Many of these intercropping studies
did not seek to determine the underlying mechanisms behind the effect of inter-
cropping on stem borer population levels. Intercropping maize with cowpea was
an effective way of reducing damage caused by C. partellus larvae migrating from
neighboring plants (146,148). This finding was confirmed by the reports that 30%
of C. partellus oviposition in maize/sorghum/cowpea-intercropping systems was
on cowpea, and the number of larvae reaching host plants from cowpea decreased
with distance (15,16).
A recent study from Kenya has reported the effectiveness of intercropping
maize with the nonhost molasses grass, Melinis minutiflora (98,99). In field tri-
als, molasses grass showed no colonization by stem borers, and when used as an
intercrop with maize, it significantly reduced stem borer infestation of the main
crop. A significant increase in parasitism of stem borers by the larval parasitoid
Cotesia sesamiae was also observed. Volatile agents produced by the molasses
grass repelled stem borers but attracted foraging C. sesamiae. Female C. sesamiae
were attracted to (E)-4, 8-dimethyl-1,3,7-nonatriene, one of the volatile compo-
nents released by intact molasses grass. Nonatriene has been implicated as an
SOS signal for recruiting predators and parasitoids and is also produced by stem
borer–damaged maize plants. Intact plants with an inherent ability to release such
as an effective cover crop, the molasses grass provides good fodder for livestock.
The grass is now being tested in on-farm trials in Kenya to control stem borers on
Another useful diversionary tactic for stem borer control is planting an outer
encircling row of some highly preferred host to act as a trap plant. Napier grass,
Pennisetum purpureum, and Sudan grass, Sorghum vulgare sudanense, common
fodder plants in Africa, were reported from Kenya to provide natural control to
stem borers by acting as trap plants (99,100). Although the stem borers oviposit
heavily on the attractive Napier grass, only few larvae complete their life cycles
(76). Napier grass has its own defense mechanism against crop borers. When the
KFIR ET AL.
larvae enter the stem, the plant produces a gummy substance that causes the death
of the pest (76). In on-farm trials in Kenya, planting Napier grass around maize
fields significantly increases crop yields by reducing the stem borer population in
maize. Sudan grass, also a fodder grass, provided natural control of stem borers
by acting as a trap plant and as a reservoir for its natural enemies (99). In field
trials conducted in Kenya, planting Sudan grass around maize fields decreased
grass around maize field also increased efficiency of natural enemies.
For the control of stem borers in resource-poor, maize farming systems in
eastern Africa, novel habitat-management strategies have been developed using
“push-pull” or stimulo-deterrent diversionary tactics (100). These strategies in-
a push-pull strategy, are Napier grass, Sudan grass, molasses grass, and silverleaf
desmodium, Desmodium uncinatum. Napier grass and Sudan grass are used as
trap plants, whereas molasses grass and silverleaf desmodium repel ovipositing
stem borers. Molasses grass, when intercropped with maize, not only reduced in-
festation of the maize by stem borers but also increased stem borer parasitism by
a natural enemy, C. sesamiae (98). All four plants are of economic importance to
farmers in eastern Africa as livestock fodder.
The economic gain from the use of intercrops depends on the balance between
a lowered cost of control of stem borers and the increased cost of maintaining an
intercropped field, along with any decrease in yield of the main crop from greater
balance between income and costs. Economic data assessing the financial returns
as well as the biological effects are therefore most useful in making decisions on
the use of intercrops and trap plants for stem borer control.
Management of Crop Residues
Crop residues are important for carrying over stem borer larval populations from
one growing season to the next. In Nigeria, larvae of B. fusca, E. saccharina,
and S. calamistis were found in crop residues below the soil surface, and higher
incidences of these borers were observed in no-tillage plots (82). In Kenya,
C. orichalcociliellus, C. partellus, E. saccharina, and S. calamistis were observed
to mulch the next crop (115). Under these conditions, borers from the old stalks
constantly infested the newly planted crops.
An effective control option would be to reduce the first generation of adult
population by destroying the larvae in old stalks (78,84,97,192). Ploughing in
order to bury maize stubble was an effective control measure used early in the
twentieth century in South Africa to control B. fusca (106). In Zimbabwe, it was
AFRICAN STEM BORERS
observed that B. fusca moths emerging through 5 cm of soil were crippled and
that deeper burial of the stalks under 10–15 cm of soil ensured that no adult moths
226,000 larvae overwinter per hectare (84). Slashing maize and sorghum stubble
destroyed 70% of C. partellus and B. fusca populations, and additional plough-
ing and disking destroyed a further 24% of the pest population in sorghum and
19% in maize (84,97). Tillage may reduce borer populations through mechanical
damage either by burying them deeply into the soil or by breaking the stems and
system is not widely practiced in South Africa because of the advent of minimum
tillage and the importance of winter grazing of maize to beef farmers (88,95).
In rural Africa, farmers often use dry stems of maize, sorghum, and millet for
building houses and fences, as fuel, bedding for livestock, boundaries of terraces
on slopes, and as stakes (68,156,161). Farmers normally stack dry stalks in the
field where they are kept until commencement of rains before being taken to vil-
lages, thus creating a reservoir for infestation in the following season (196). To
solve this problem, early cutting of stalks and horizontal placement on the soil
surface has been recommended. This was observed to cause mortality of 97% of
stem borers in maize and 100% in sorghum in Ethiopia (58) and to reduce the
residual population of borers in millet from 16% in uncut stems to 3% (227). The
high level of mortalities of C. partellus, C. orichalcociliellus, and S. calamistis in
horizontally placed stalks was ascribed to the effects of sun and heat, more specif-
ically, the reaching of the thermal threshold for survival (147). On the other hand,
in Nigeria, the control of S. calamistis, B. fusca, C. ignefusalis, and E. saccha-
rina on maize by removing stalks and stubble after harvest did not reduce stem
borer populations significantly apparently because of migration of adults into the
immediately after harvest has been recommended (8,68,80,192). In Tanzania,
almost complete eradication of C. partellus on maize and sorghum was achieved
by burning (53). About 95% of farmers in Nigeria kept sorghum stalks and did
not follow the recommended practice of burning them after harvest (4). As a
compromise, partial burning was recommended while the leaves are dry but the
stalks are not (4,8). Heat generated from the burning leaves kills up to 95% of
larvae and at the same time cures the stalks, making them stronger as building
materials and more resistant to termite attack (4,12). Crop residues are the only
organic matter added into soils in many small-scale African farms. Burning crop
residues can create problems in farms where the organic content of soils is low
and soil erosion from wind and rain is severe (196).
KFIR ET AL.
Manipulation of Sowing Dates and Densities
Growing crops when the pest is least abundant ensures that the most susceptible
the February–May period, when infestation is normally low. There is no available
information on the effectiveness of these measures, and the last implementation
of this legislation was in 1937–1938 (72).
Several workers have reported more damage by stem borers to late-planted maize
(62,64), Nigeria (5,44,68), and Zaire (77). Thus, in some areas of West Africa
This affects the borer populations in the rain forest zone where alternative wild
host plants in the dry season are scarce (34). This strategy is also desirable in the
semi-arid tropics, where rainfall is variable and unpredictable; late sowing is not
desirable because the yield of late sown crops is low, even if free from stem borers
In the Highveld region of South Africa, the second-generation population B.
fusca is larger and can cause more damage than the first generation (211). The
best strategy to reduce damage would be to plant early in the growing season.
Similar observations were made in Lesotho (55), Zimbabwe (179), and Ethiopia
where second-generation larvae caused crop losses of 22.5–100% compared with
0–22.6% by first generation (60).
In the lower elevations of South Africa, it is recommended that sorghum
be planted after mid-October to avoid infestation from the first moth peak of
season was more infested by B. fusca than maize sown later. In Malawi, planting
date influenced infestation levels of B. fusca and C. partellus on sorghum; how-
ever, the choice of optimum sowing date depends on the sorghum cultivar (111).
In contrast, in the Sahelian region, manipulating the planting dates of millet is not
a good method to reduce infestation by C. ignefusalis (227).
Sowing density may affect the crop and its pest population, as well as the
behavior of the pest in searching for food or an oviposition site (102). Young
C. partellus migrate from their hatching site to the funnel leaves or to other plants
within the vicinity. During this migration, up to 100% mortality may occur (108).
The lowest incidence of deadheart caused by B. fusca occurred at the lowest
plant density in sorghum in South Africa (198) and maize in Nigeria (134). A
reduction in row width increased the number of larvae able to reach adjacent plant
rows through migration, and this in turn resulted in more damaged plants (211).
B. fusca larvae migrate up to 2.4 m (68) and at a 0.9 m interrow spacing, which is
common in commercial agriculture; they migrated over 4 rows from the hatching
AFRICAN STEM BORERS
E. saccharina damage (118). Studies on C. partellus in maize planted at different
densities (56) and on C. ignefusalis in millet (57) showed no significant difference
in oviposition or borer incidence.
In subsistence farming systems in Africa where farmers normally intercrop
cereals with other crops and lack of water is a major constraint, manipulation of
sowing dates and management of plant densities is not always practical as farmers
often plant after first rains (196). Farmers’ choice of sowing dates is affected by
many constraints, and it is unlikely that a change in sowing date alone will result
in higher sustainable yields (126).
increased with the application of fertilizer (191), and in South Africa sorghum
plants without fertilizers were less preferred for oviposition by C. partellus (197),
in the survival of S. calamistis larvae and acceleration in larval development with
in the number of generations per year (171).
Nitrogen fertilization enhances borer development as well as the plant’s toler-
ance to borer attack. Yield losses decreased linearly from 20% with no fertilizer
to 11% with 120 kg of nitrogen per hectare (172). Ajayi (10) observed that time of
that the manipulation of time of nitrogen application may achieve a compromise
between using low levels of nitrogen for low stem borer infestation and using high
levels for better yields.
Host Plant Resistance
Host plant resistance as an approach to pest management in gramineous crops
confers many advantages (34). Resistant crop varieties provide an inherent con-
trol that involves no environmental problems, and they are generally compatible
with other insect-control methods. The cultivation of resistant crop plants is not
subject to the vagaries of weather as are chemical-control measures, and in certain
circumstances it is the only effective means of control. Resistant varieties control
even a low pest density, whereas insecticide use is justifiable only when the den-
sity reaches the economic injury level. Efforts are underway in Africa to identify
sources of stem borer resistance in cereal crops, but high levels of resistance have
not been found.
KFIR ET AL.
Breeding for Resistance
Efforts have been made to incorporate resistance of maize and sorghum to
C. partellus into a good agronomic background during the past two decades
(6,7,14), and many genotypes are already in national yield trials. Resistant lines/
hybrids with good general combining ability have been identified (167). Several
hybrid sorghums bred in South Africa exhibited greatest tolerance to stem borer
damage and therefore suffered low yield losses (202).
Over the past 60 years, there have been numerous attempts to introduce exotic
parasitoids into Africa and the Indian Ocean Islands for biological control of
exotic and native stem borers, but only a few species have established (Table 2).
On the mainland, only C. flavipes has established.
In 1993, the gregarious larval endoparasitoid, C. flavipes, was introduced
into Kenya from Pakistan for biological control of C. partellus. Releases were
made in the southern coastal area of Kenya (144), and the parasitoid was re-
covered during the season of release from C. partellus and two native stem bor-
ers, C. orichalcociliellus and S. calamistis (143). In 1994 only one stem borer
parasitized by C. flavipes was found despite intensive sampling. In 1995 and
1996 a few recoveries were made but parasitism remained low (143). In 1997
the number of recoveries increased dramatically and parasitism at 30 sites av-
eraged about 6%. Parasitism continued to increase during the next two years
with average parasitism of about 13% in 1999 (228). Surveys in other maize-
(183) and in the area bordering Lake Victoria in western Kenya (139). In the
Eastern Province, C. flavipes was found in low densities in 1996 and then
released at three sites in 1997. Parasitism during the season following
the releases was about 14% (183). C. flavipes was never intentionally released
in western Kenya, but Omwega et al. (139) speculated that the establish-
ment was the result of insects that escaped from a local laboratory colony in
1992. However, parasitism in western Kenya has not increased to the levels
observed in coastal Kenya or the Eastern Province (133). In western Kenya, four
stem borers are common in maize, C. partellus, S. calamistis, B. fusca, and E.
saccharina (163), all of which are attractive and acceptable hosts for C. flavipes.
Two of which, B. fusca and E. saccharina, are not suitable for its development
hosts in an area would create a sink for C. flavipes eggs and depress population
The impact of C. flavipes on stem borer populations in coastal Kenya was
recently investigated (228). A ratio-dependent, host-parasitoid model was used to
1.6 stem borers/plant, equivalent to a 32–55% decrease in the stem borer density,
AFRICAN STEM BORERS
Exotic stem borer parasitoids established in mainland Africa and Indian Ocean Islands
aCP, Chilo partellus; CS, Chilo sacchariphagus; SC, Sesamia calamistis; SB, Sciomesa biluma.
bOld/new is used when two stem borers were targets. The host listed first in the “target host” column is the old association host.
cThere is confusion regarding the origin of the C. flavipes populations in Mauritius and Reunion. Appert (19) indicated that C. flavipes was introduced into Mauritius in 1917 and then later
in Reunion, but provides no details or reference. Greathead (65) indicated that there was no record of the introduction, but he assumed that it was exotic. One possibility is that it arrived on the islands along with its host, C. sacchariphagus, around 1850.
dThe determination of old vs. new associations with C. sacchariphagus is difficult because the origin and taxonomic status of the population of C. sacchariphagus in Mauritius is unclear.
Williams (222) speculated that the borer either arrived in Mauritius from Sri Lanka in 1848, or from Java in 1850.
KFIR ET AL.
borers in the future.
In addition to the work conducted in Kenya, a survey in 1995 in northern and
bordering southwestern Kenya (140). Based on surveys conducted prior to 1994,
and on electrophoretic evidence, it was concluded that the most likely explanation
was that C. flavipes moved into Tanzania from Kenya (140).
Releases of C. flavipes were made in Mozambique in 1996 (50) and in Uganda
and Somalia in 1997 (141). Recoveries in Mozambique in 1999 indicated that the
parasitoid had established but parasitism was low. In Uganda, one year after the
release of C. flavipes, it had become the most common parasitoid of a complex
of four stem borers, and parasitism averaged about 20% (107). No postrelease
surveys have been conducted in Somalia, but recoveries in neighboring Ethiopia,
Releases in Zimbabwe, Zambia, Zanzibar, and Malawi were made in 1998–1999,
but to date, establishment has not been confirmed (W.A. Overholt, unpublished
Examination of the failures and successes in establishing exotic parasitoids
suggests several factors that may be related to establishment. The success rate on
the Indian Ocean Islands has clearly been much higher than on mainland Africa,
where the only confirmed establishment has been C. flavipes. The higher rate of
establishment on islands is true not only for stem borer parasitoids but for natural
enemies introduced against other pests as well. Greathead (65) reviewed classical
biological control in Africa and suggested that the greater success on the Indian
Ocean Islands could be explained by the theory of island biogeography (104).
Species are packed more tightly and are more specialized on continents than on
islands, and thus, specialist parasitoids encounter less resistance (competition)
when colonizing islands.
The geographic factor of islands versus the mainland is confounded with a sec-
control activities on the Indian Ocean Islands was against C. sacchariphagus in
a sugarcane ecosystem, whereas in mainland Africa the work has been directed
mostly in maize and sugarcane, with lesser effort in rice. The only successes
in maize have been the introductions of Pediobius furvus and C. sesamiae from
mainland Africa to Indian Ocean Islands for control of S. calamistis and the recent
establishment of C. flavipes in Kenya.
establishment. Sugarcane is grown throughout the year and thus provides a fairly
stable habitat. In contrast, the maize ecosystem is short-lived, providing a suitable
habitat for stem borers and their natural enemies for only two or three months.
(Comores) was the reason for several failures of the establish parasitoids against
AFRICAN STEM BORERS
C. partellus in maize. A review of success rates in biological control programs
in different categories of habitat stability supports the hypothesis that the chance
of establishment increases with an increase in habitat stability (66). However,
the lack of success in South Africa against E. saccharina in sugarcane, despite
15 years of intense effort (47), suggests that a semiperennial sugarcane habitat
alone is not sufficient for the establishment of parasitoids. Conlong (47) points out
that climate compatibility is also a major factor that influences establishment and
suggests that the temperate climate of South Africa has not been conducive for
the establishment of parasitoids from tropical and subtropical regions. Similar
explanations have been given for the lack of success in biological control of
C. partellus in South African maize (91,181).
Old host/parasitoid associations more likely result in establishment than do
new associations. All establishments on the Indian Ocean Islands and the only
confirmed establishment on mainland Africa included at least one old-association
relationship (see footnote c to Table 2). The behavioral and physiological com-
patibility of old-association parasitoids and their hosts are implicit, whereas in
new associations compatibility cannot be assumed (220). The greater success of
old associations supports the contention of several authors (66,67,214). How-
ever, as stem borers typically occur in complexes, at times including both exotic
and native pests, introduced parasitoids will often encounter both old- and new-
association hosts when colonizing a new area. The probability of establishment
and the level of suppression of the stem borer complex may depend not only on
the old host/parasitoid relationship(s) but also on the compatibility of the new
Finally, a notable aspect of the introduction of stem borer parasitoids into
Africa has been the number of times Cotesia spp. have established as compared
with other parasitoids. Worldwide, C. flavipes has been introduced into more than
40 countries in the tropics and subtropics for biological control of crambid stem
borers, primarily those in the genera Chilo and Diatraea (151). In Mauritius, par-
asitism of 4–50% of larvae of the introduced stem borer, Chilo sacchariphagous,
parasitism of 60% of C. sacchariphagous larvae was reported (28). The success of
(Table 2), where it was introduced against S. calamistis.
Several factors could be responsible for the success of the two Cotesia spp.
In their areas of endemism, the Cotesia spp. parasitoids often attack more than
one species of crambid and/or noctuid stem borers (26,63,114,116,122,173). A
parasitoid that exploits more than one of the hosts in the target habitat may better
colonize a new area than a parasitoid with a narrow host range owing to a more
constant availability of hosts and a lack of population growth depression due to
wasting eggs in attractive but unsuitable hosts.
Another factor that may predispose C. flavipes and C. sesamiae to establish
is gregarious reproduction. Hopper & Roush (75) suggested that an Allee effect
might explain many failures to establish natural enemies for biological control.
KFIR ET AL.
tokus parasitoid, might decrease chances of mate finding, thus leading to possible
extinction. However, C. flavipes and C. sesamiae mate with their siblings before
dispersal (21,144,157), and thus Allee effects will not influence establishment.
A high host-searching ability has been suggested as a factor involved in the
success of C. flavipes. Even at low host densities, C. flavipes successfully located
stem borer hosts (219). The high host-searching success of C. flavipes may in
part be due to its behavior of entering tunnels in plant stems to attack stem borer
larvae (182). Many larval parasitoids of stem borers remain on the outside of the
stem and attack their hosts by drilling, or locating breaches through the stem, with
their ovipositor (182). This strategy may be effective for attacking stem borers in
small-stemmed wild grasses, but in relatively larger-stemmed cultivated grasses,
Thanks to the Illinois Natural History Survey, Department of Entomology, De-
partment of Natural Resources and Environmental Sciences, and the Office of
International Programs of the University of Illinois for support during a sabbat-
ical leave at INHS by the senior author while this article was prepared. Thanks
to Tharina Bird and Kerstin Kr¨ uger (ARC-PPRI) for help in sorting out the refer-
ences. R. Wiedenmann (INHS) constructively commented on an early draft of the
Visit the Annual Reviews home page at www.AnnualReviews.org
1. Abu JF. 1986. Biology and Control of the
Insect Pests of Sorghum in the Southern
Guinea Savanna Zone of Nigeria. Zaria,
Nigeria: Inst. Agric. Res. Samaru. 23 pp.
2. Adenuga AO. 1977. Comparative meth-
ods and the economics of control of
stem borers (Lepidoptera: Noctuidae and
Pyralidae) on local maize, Zea mays.
Nigerian Soc. Plant Prot.
3. Adesiyun AA. 1983. Some effects of in-
infestation by lepidopterous stalk-borers,
particularly Busseola fusca. Insect Sci.
4. Adesiyun AA, Ajayi O. 1980. Control of
the sorghum stem borer, Busseola fusca,
by partial burning of the stalks. Trop. Pest
5. Adeyemi SAO, Donelly J, Odetoyinbo
JA. 1966. Studies on chemical control of
stem borers of maize. Nigerian Agric. J.
6. Ajala SO. 1993. Population cross dial-
lel among maize genotypes with varying
levels of resistance to the spotted stem
7. Ajala SO, Saxena KN, Chiliswa P. 1995.
Selection in maize (Zea mays L.) for re-
sistance to the spotted stem borer (Chilo
partellus (Swinhoe)). Maydica 40:137–
AFRICAN STEM BORERS
Agric. Res., Ahmadu Bello Univ., Zaria,
9. Ajayi O. 1987. Insecticidal control of the
sorghum stem borer. Cereals Res. Pro-
gramme. Crop. Scheme Rep. Inst. Agric.
Res., Samaru, Zaria, Nigeria
10. Ajayi O. 1990. Possibilities for integrated
control of the millet stem borer, Acigona
11. Ajayi O. 1991. Yield reduction in sor-
ghum caused by Busseola fusca (Fuller)
and Sesamia calamistis Hampson. Conf.
Afr. Assoc. Insect Sci., 9th, Accra, Ghana
Ref. 150, pp. 39–45
13. Akinsola EA, Agyen-Sampong M. 1984.
The ecology, bionomics and control of
14. Ampofo JKO, Dabrowski ZT, Omolo
EO. 1986. Registration of germplasm
lines of maize. Crop Sci. 26:650
15. Ampong-Nyarko K, Seshu Reddy KV,
Nyang’or RA, Saxena KN. 1994. Reduc-
tion of pest attack on sorghum and cow-
16. Ampong-Nyarko K, Seshu Reddy KV,
Saxena KN. 1994. Chilo partellus (Swin-
hosts: a mechanism for reduced pest in-
cidence in intercropping. Acta Ecol. 15:
17. Anderson TJ. 1929. Control of maize
stalk borers. Bull. Dep. Agric. Kenya, 7F.
18. Appert J. 1970. Insects Harmful to Maize
in Africa and Madagascar. Madagascar:
Madagascar Inst. Agron. Res. 71 pp.
19. Appert J. 1973. Entomofaune parasitaire
des foreurs des gramin´ ees ` a Madagascar.
20. Appert J, Betbeder-Matibet M, Ranai-
vosoa H. 1969. Vingt ann´ ees de lutte
biologique ` a Madagascar. Agron. Trop.
21. Arakaki N, Gahana Y. 1986. Emergence
pattern and mating behavior of Apanteles
flavipes (Cameron) (Hymenoptera: Bra-
22. Aroga R. 1988. Entomologie des ce-
reales. Rapport d’activities; ann´ ees 1987
23. Atkinson PR. 1979. Distribution and nat-
ural hosts of Eldana saccharina Walker
in Natal, its oviposition sites and feeding
patterns. Proc. S. Afr. Sugar Tech. Assoc.
24. Barrow MR. 1987. The effect of first
generation maize stalkborer, Busseola
fusca (Fuller) (Lepidoptera: Noctuidae),
on yield of different maize genotypes. J.
Entomol. Soc. S. Afr. 50:291–98
25. BeevorPS, DavidH, JonesOT.1990.Fe-
in pest control applications. Insect Sci.
26. Beg MN, Inayatullah C. 1980. Studies on
Apanteles flavipes a parasite of gramina-
ceous borers. Pak. J. Agric. Res. 1:50–53
27. Berger A. 1981. Biological control of
the spotted stalkborer, Chilo partellus
(Swinhoe) in maize by using the bac-
teria Bacillus thuringiensis. Rep. Inst.
Nacional de Investigaáao Agronomica.
Proj. UNDP/FAO MOZ/75/009, Maputo,
28. Betbeder-Matibet M, Malinge P. 1968.
Un succ` es de la lutte biologique: contrˆ ole
de Proceras sacchariphagus Boj. Borer
ponctu´ e de la canne ` a sucre ` a Madagas-
car par un parasite introduit: Apanteles
flavipes Cam. Agron. Trop. 22:1196–220
29. Bleszynsky S. 1970. A revision of the
world species of Chilo Zincken (Lepi-
doptera: Pyralidae). Bull. Br. Mus. (Nat.
Hist.) Entomol. 25:101–95
30. Bonhof MJ. 1998. Predators. See Ref.
150, pp. 295–307
KFIR ET AL.
on maize stem borers in coastal Kenya.
PhD thesis. Wageningen Univ., Neth. 181
32. Bosque-P´ erezNA, MareckJH.1990.Dis-
tribution and species composition of lep-
idopterous maize borers in southern Ni-
geria. Bull. Entomol. Res. 80:363–68
33. Bosque-P´ erez NA, Mareck JH. 1991. Ef-
fect of the stem borer Eldana saccharina
(Lepidoptera: Pyralidae) on the yield of
maize. Bull. Entomol. Res. 81:243–47
34. Bosque-P´ erez NA, Schulthess F. 1998.
Maize: West and Central Africa. See Ref.
150, pp. 11–27
35. Deleted in proof
36. Breni` ere J. 1971. Les probl` emes des
l´ epidopt` eres foreurs des gramin´ ees en
37. Breni` ere J. 1976. Reconnaissance des
principaux l´ epidopt` eres du riz de l’Af-
rique de l’Ouest. Agron. Trop. 1:212–31
38. Breni` ere J, Bordat D. 1982. Int´ erˆ et et dif-
tion d’entomophages—cas des foreurs du
riz en Afrique. Agron. Trop. 37:203–6
39. Breni` ere J, Bordat D,
Hamza H, Renand M. 1985. Les
op´ erations de lutte biologique contre le
foreur du mais Chilo partellus (Swin-
hoe), Lepidoptera, dans l’Ile de Ngazidja.
Agron. Trop. 40:157–66
40. Breni` ere J, Rodriguez H, Ranaivosoa H.
1962. Un ennemi de riz ` a Madagascar,
Maliarpha separatella Rag. ou boreur
blanc. Agron. Trop. 17:223–302
41. CABI. 1989. Chilo partellus (Swinh.).
Distribution Maps of Pests. Series A
(Agriculture). No. 184. London: Int. Inst.
42. Campion DG, Nesbitt BF. 1983. The uti-
lization of sex pheromones for the control
43. Cardwell KF, Schulthess F, Ndemah R,
Ngoko Z. 1997. A systems approach to
assess crop health and maize yield losses
due to pests and diseases in Cameroon.
Agric. Ecosyst. Environ. 65:33–47
44. Carter AO. 1985. An evaluation of the im-
portance of stem borers and their control
on maize grown in south eastern Nigeria
MSc thesis. Univ. Reading, UK. 57 pp.
45. Conlong DE. 1990. A study of pest-
parasitoid relationships in natural habi-
tats: an aid towards the biological con-
trol of Eldana saccharina (Lepidoptera:
Pyralidae) in sugarcane. Proc. S. Afr.
Sugar Tech. Assoc. 64:111–15
46. Deleted in proof
47. Conlong DE. 1997. Biological control
of Eldana saccharina Walker in South
African sugarcane: constraints identified
48. Cook M. 1997. Revision of the genus
Maliarpha Ragonot (Lepidoptera: Pyral-
idae), based on adult morphology with a
tomol. Res. 87:25–36
49. Critchley BR,Hall DR,
McVeigh LJ,Mulaa MA,
1997. Monitoring and mating disruption
of the maize stalkborer, Busseola fusca,
in Kenya with pheromones. Crop Prot.
50. CugalaD, OverholtWA, GigaD, Santos
and Cotesia flavipes (Hymenoptera: Bra-
conidae) as biological control agents
Afr. Crop Sci. J. 7:497–502
51. Delobel A. 1975. Chilo orichalcociliellus
Strand (Lepidoptera: Pyralidae), fˆ oreur
des tiges du sorgho et du mais ` a Mada-
gascar II. Premi` eres donn´ ees biologiques.
ORSTOM S´ er Biol. 10:11–16
52. Dent D. 1991. Insect Pest Management.
Wallingford, UK: CABI. 604 pp.
53. Duerden JC. 1953. Stem borers of cereal
crops at Kongwa, Tanganyika, 1950–52.
East Afr. Agric. For. J. 19:105–19
54. Du Plessis C, Lea HAF. 1943. The maize
stalk borer, Calamistis fusca (Hmps).
Bull. Dep. Agric. For. Union S. Afr. 238.
55. Ebenebe AA, Van den Berg J, Van den
AFRICAN STEM BORERS
Linde TC. 1999. Effect of planting date
of maize on damage and yield loss caused
by the stalk borer, Busseola fusca (Fuller)
(Lepidoptera: Noctuidae) in Lesotho. S.
Afr. J. Plant Soil 16:180–85
56. Firempong S. 1988. The effect of maize
lus oviposition in the field. ICIPE 15th
Annu. Rep. 1987
57. Gahukar RT. 1989. Pest and disease inci-
dence in pearl millet under different plant
density and intercropping patterns. Agric.
Ecosyst. Environ. 26:69–74
58. Gebre-Amlak A. 1988. Survival of maize
residues in Ethiopia. Crop Prot. 7:183–
apause in the maize stalk borer, Busse-
ola fusca (Fuller) (Lep., Noctuidae), in
Awassa, southern Ethiopia. J. Appl. En-
60. Gebre-Amlak A, Sigvald R, Petterson
J. 1989. The relationship between sow-
ing date, infestation and damage by the
maize stalkborer, Busseola fusca (Noctu-
Pest Manag. 35:143–45
61. Girling DJ. 1978. The distribution and bi-
other stemborers in Uganda. Bull. Ento-
mol. Res. 68:471–88
62. Girling DJ. 1980. Eldana saccharina as
a crop pest in Ghana. Trop. Pest Manag.
63. Goraya AA, Mushtaque M, Attique MR.
1982. Suitability of Apanteles flavipes
(Cameron) (Hym.: Braconidae) as a bio-
control agent for graminaceous stem-
borers in Pakistan. Proc. Ann. Conv. Pak.
Soc. Sugar Tech., 18th, Rawalpindi, pp.
177–82. Lahore: Pakistan Soc. Sugarcane
64. Gounou S, Schulthess F, Shanower T,
Hammond WNO, Braima H, et al. 1993.
Stem and Ear Borers of Maize in Ghana.
Plant Health Manag. Div., Res. Monogr.
3. Ibadan, Nigeria: Int. Inst. Trop. Agric.
65. Greathead DJ. 1971. A Review of Biologi-
Commun. No. 5. Commonw. Inst. Biol.
Control. Farnham R., Engl: Commonw.
Agric. Bur. 162 pp.
66. Hall RW, Ehler LE. 1979. Rate of es-
tablishment on natural enemies in classi-
67. Hall RW, Ehler LE, Bisabri-Ershadi B.
1980. Rate of success in classical biolog-
ical control of arthropods. Bull. Entomol.
Soc. Am. 26:111–14
68. Harris KM. 1962. Lepidopterous stem
borers of cereals in Nigeria. Bull. Ento-
mol. Res. 53:139–71
69. Harris KM. 1989. Biology of sorghum
stem borers. See Ref. 70, pp. 63–71
70. Harris KM. 1989. Recent advances in
sorghum and pearl millet stem borer
research. In Int. Workshop Sorghum
Stem Borers, Patancheru, 1987, ed. KF
Nwanze, pp. 9–16. Patancheru, India:
71. Harris KM. 1990. Bioecology and Chilo
species. Insect Sci. Appl. 11:467–77
72. Harris KM, Nwanze KF. 1992. Busse-
ola fusca (Fuller), the African maize stem
borer: a handbook of information. Inf.
Bull. 33. ICRISAT. Oxon, UK: CABI. 84
73. Harris KM, Youm O. 1998. Millet: West
Africa. See Ref. 150, pp. 47–48
74. Hawkins BA, Browning HW, Smith JW
Jr. 1987. Field evaluation of Allorhogas
pyralophagus (Hym.: Braconidae) im-
ported into Texas for biological control
of the stalkborer Eoreuma loftini (Lep.:
Pyralidae) in sugarcane. Entomophaga
75. Hopper KR, Roush TR. 1993. Mate find-
ing, dispersal, number released, and the
success of biological control introduc-
tions. Ecol. Entomol. 18:321–31
76. Hutter NJ. 1996. An assessment of the
potential value of ten varieties of Napier
KFIR ET AL.
respect to their use as a trap crop for
tacking maize (Zea mays). MSc thesis.
Univ. Newcastle-upon-Tyne, UK. 120 pp.
77. IITA. 1990. Final Tech. Rep. IITA/
USAID/Zaire Appl. Agric. Res. Outreach
Proj. IITA, Ibadan, Nigeria
78. Ingram WR. 1958. The lepidopterous
Uganda. Bull. Entomol. Res. 49:367–83
79. Ingram WR. 1983. Biological control of
graminaceous stem-borers and legume
pod borers. Insect Sci. Appl. 4:205–9
80. Ingram WR,Irving NS,
cultural Pests in Botswana. Gaborone:
Gov. Printer. 129 pp.
81. Jepson WF. 1954. A Critical Review of
Stalk Borers of Tropical Graminaceous
Crops. London: Commonw. Inst. Ento-
mol. 127 pp.
82. Kaufmann T. 1983. The behavioral biol-
ogy, feeding habits and ecology of three
fusca in Ibadan, Nigeria, West Africa. J.
Ga. Entomol. 18:259–72
83. Kfir R. 1988. Hibernation by the lepi-
dopteran stalk borers, Busseola fusca and
Chilo partellus on grain sorghum. Ento-
mol. Exp. Appl. 48:31–36
84. Kfir R. 1990. Prospects for cultural con-
trol of the stalk borers, Chilo partellus
(Swinhoe) and Busseola fusca (Fuller), in
summer grain crops in South Africa. J.
Entomol. Soc. S. Afr. 53:41–47
85. Kfir R. 1991. Duration of diapause in the
stem borers, Busseola fusca and Chilo
partellus. Entomol. Exp. Appl. 61:265–70
86. Kfir R. 1991. Effect of diapause on de-
velopment and reproduction of the stem
borers Busseola fusca (Lepidoptera: Noc-
tuidae) and Chilo partellus (Lepidoptera:
87. Kfir R. 1991. Non-chemical control of
grain stalk borers. Plant Prot. News 23:4
88. Kfir R. 1992. Alternative, non-chemical
partellus (Swinhoe) and Busseola fusca
(Fuller), in summer grain crops in South
Africa. Tech. Commun. Dep. Agric. Dev.,
Rep. S. Afr. 232:99–103
89. Kfir R. 1993. Diapause termination in
Busseola fusca (Lepidoptera: Noctuidae)
in the laboratory. Ann. Entomol. Soc. Am.
90. Kfir R. 1993. Diapause termination in the
spotted stem borer, Chilo partellus (Lepi-
Appl. Biol. 123:1–7
91. Kfir R. 1994. Attempts at biological
control of the stem borer Chilo partel-
lus (Swinhoe) (Lepidoptera: Pyralidae) in
South Africa. Afr. Entomol. 2:67–68
92. Kfir R. 1997. Competitive displacement
of Busseola fusca (Lepidoptera: Noctu-
idae) by Chilo partellus (Lepidoptera:
Pyralidae). Ann. Entomol. Soc. Am. 90:
93. Kfir R. 1997. Natural control of the ce-
real stemborers Busseola fusca and Chilo
94. Kfir R. 1998. Maize and grain sorghum:
southern Africa. See Ref. 150, pp. 29–37
95. Kfir R. 2000. Seasonal occurrence, par-
asitoids and pathogens of the African
stem borer, Busseola fusca (Fuller) (Lep-
idoptera: Noctuidae), on cereal crops in
South Africa. Afr. Entomol. :1–14
96. Kfir R, Bell R. 1993. Intraseasonal
changes in populations of the African
maize stem borer, Busseola fusca (Fuller)
(Lepidoptera: Noctuidae), in Natal, South
Africa. J. Afr. Zool. 107:543–53
97. Kfir R, Van Hamburg H, Van Vuuren
R. 1989. Effect of stubble treatment on
the post-diapause emergence of the stalk
borer Chilo partellus (Swinhoe) (Lepi-
doptera: Pyralidae). Crop Prot. 8:289–92
98. Khan ZR, Ampong-Nyarko K, Chiliswa
P, Hassanali A, Kimani S, et al. 1997. In-
tercropping increases parasitism of pests.
AFRICAN STEM BORERS
99. Khan ZR, Chiliswa P, Ampong-Nyarko
K, SmartLE, PolaszekA,etal.1997.Uti-
lization of wild gramineous plants for ce-
100. Khan ZR, Pickett JP, Van den Berg J,
Wadhams LJ, Woodcock CM. 2000. Ex-
ploiting chemical ecology and species di-
versity: stem borer and striga control for
101. Kioko EN, Overholt WA,
1995. Larval development in Chilo
orichacociliellus and Chilo partellus:
a comparative study in the laboratory.
Proc. Meet. Sci. Conf. Afr. Assoc. Insect
Sci., 10th, Mombasa, 1993, pp. 191–98.
Nairobi: Afr. Assoc. Insect Sci.
102. Lawani SM. 1982. A review of the effects
of various agronomic practices on cereal
103. Leslie GW. 1988. The identification and
importance of predators of Eldana sac-
Afr. Sugar Tech. Assoc. 67:122–26
104. MacArthur RH, Wilson EO. 1967. The
Theory of Island Biogeography. Prince-
ton, NJ: Princeton Univ. Press. 208 pp.
105. Maes K. 1998. Pyraloidea: Crambidae,
Pyralidae. See Ref. 150, pp. 87–98
106. Mally CW. 1920. The maize stalk borer
Busseola fusca (Fuller). Bull. Dep. Agric.
S. Afr., 3. 111 pp.
107. Matama-Kauma T. 2000. Yield losses
caused by stem borers on maize and es-
tablishment of Cotesia flavipes Cameron
(Hymenoptera: Braconidae) in Eastern
Uganda. MSc thesis. Makerere Univ.,
Kampala. 98 pp.
C. orichalcociliellus Strand (Lep., Cram-
bidae) and Sesamia calamistis Hmps.
(Lep., Noctuidae) on maize in the Coast
Province, Kenya. Mitt. Schweiz. Entomol.
109. Matthee JJ. 1974. Pests of graminaceous
crops in South Africa. Entomol. Mem. 40,
Dep. Agric. Tech. Serv., Rep. S. Afr., 24
110. McFarlane J. 1990. Damage assessment
and yield losses in sorghum due to the
stem borer Busseola fusca (Fuller) (Lep-
idoptera: Noctuidae) in northern Nigeria.
Trop. Pest Manag. 36:131–37
111. Mchowa JW. 1990. The effect of plant-
ing date on the incidence of the stalk bor-
ers, Busseola fusca (Fuller) (Lepidoptera:
Noctuidae) and Chilo partellus (Swin-
hoe) (Lepidoptera: Pyralidae) in two
varieties of sorghum. Proc. Reg. Work-
Conf., 6th, Bulawayo, 1989, pp. 154–59.
Bulawayo, Zimbabwe: ICRISAT
112. Megenasa T. 1982. Insect pests of
Sorghum Improvement East. Afr., 1982,
Sorghum Improv. Proj.
113. Minja EM. 1990. Management of Chilo
sect Sci. Appl. 11:489–99
114. Mohyuddin AI. 1971. Comparative bi-
ology and ecology of Apanteles flavipes
(Cam.) and A. sesamiae (Cam.) as para-
sites of graminaceous borers. Bull. Ento-
mol. Res. 61:33–39
115. Mohyuddin AI, Greathead DJ. 1970. An
annotated list of the parasites of gramina-
ceous stem borers in East Africa, with a
discussion of their potential in biological
control. Entomophaga 15:241–74
116. Mohyuddin AI, InayatullahC, KingEG.
in Apanteles flavipes (Cameron) (Hy-
biological control of graminaceous stem-
borers (Lepidoptera: Pyralidae). Bull. En-
tomol. Res. 71:575–81
117. MoutiaLA, CourtoisCM.1952.Parasites
of the moth-borers of sugar-cane in Mau-
ritius. Bull. Entomol. Res. 43:325–59
age in relation to maize stand density and
water stress in the Ivory Coast. Int. J. Pest
KFIR ET AL.
119. Moyal P. 1998. Infestation patterns and
ola fusca (Fuller) (Lepidoptera: Noctu-
idae), in the Ivory Coast. Afr. Entomol.
120. Moyal P, Tran M. 1992. Chilo aleniel-
lus (Lepidoptera: Pyralidae), a stem borer
of maize in Cˆ ote d’Ivoire. Bull. Entomol.
121. Muyango S. 1987. Les chenilles foreuses
des tiges du ma¨ ıs et du sorgho: ´ ecologie
et lutte. In S´ eminaire sur les maladies
et les ravageurs des principles cultures
vivri` eres d’Afrique Centrale, pp. 117–
26. Ede/Wageningen: Centre Tech. Coop.
Agric. Rurale (CTA)
122. Nagarkatti S, Nair KR. 1973. The influ-
Cyperaceae on populations of sugarcane
borers and their parasites in North India.
123. Ngi-SongAJ, OverholtWA, AyerteyJN.
1995. Suitability of African gramineous
stemborers for the development of Cote-
sia flavipes and Cotesia sesamiae (Hy-
menoptera: Braconidae). Environ. Ento-
124. Ngi-Song AJ, Overholt WA, Njagi PGN,
Dicke M, Ayertey JN, Lwande W. 1996.
Volatile infochemicals used in host
and host habitat location by Cotesia
flavipes Cameron and Cotesia sesamiae
(Cameron) (Hymenoptera: Braconidae),
larval parasitoids of stemborers on Gram-
inae. J. Chem. Ecol. 22:307–22
125. Nunes E, Sousa D, Sataric I. 1985. Re-
tion in Mozambique. To feed ourselves.
Proc. First East., Cent. South. Afr. Reg.
Maize Workshop, Lusaka, Zambia
126. Nwanze KF, Mueller RAE. 1989. Man-
agement options for sorghum stem bor-
ers for farmers in the semi-arid trop-
ics. Int. Workshop Sorghum Stem Borers,
Patancheru, 1987, ed. KF Nwanze. pp.
105–14. Patancheru, India: ICRISAT
inaceous crops in East Africa. Colonial
tionary Off. 48 pp.
128. Odindo MO. 1990. Potential of Nosema
spp. (Microspora, Nosematidae) and
viruses in the management of Chilo spp.
129. Ofomata VC. 1997. Ecological inter-
actions between Chilo orichalcociliellus
Strand and Chilo partellus (Swinhoe)
(Lepidoptera: Pyralidae) on the Kenya
Anambra State, Nigeria. 206 pp.
130. OfomataVC, OverholtWA, EgwuatuRI.
1999. Diapause termination of Chilo
partellus (Swinhoe) and Chilo orichal-
cociliellus Strand (Lepidoptera: Pyrali-
dae). Insect Sci. Appl. 19:187–91
131. Ofomata VC, Overholt WA, Lux SA,
tive studies on the fecundity, egg survival,
larval feeding and development of Chilo
partellus (Swinhoe) and Chilo orichal-
cociliellus Strand (Lepidoptera: Cram-
132. Ofomata VC, Overholt WA Van Huis A,
Egwuatu RI, Ngi-Song AJ. 1999. Niche
overlap and interspecific association be-
tween Chilo partellus and Chilo orichal-
cociliellus on the Kenya coast. Entomol.
Exp. Appl. 93:141–48
133. Ogedah KO. 1999. Evaluation of the im-
pact of Cotesia flavipes Cameron (Hy-
menoptera: Braconidae) and indigenous
parasitoids on stemborer populations in
southwestern Kenya. MSc thesis. Ken-
yatta Univ., Nairobi. 71 pp.
134. Ogunwolu EO, Nwosu K, Ogunyebi SO.
1981. Stem borer damage in maize as af-
fected by host plant density. J. Agric. Sci.
135. Okuda T. 1988. Effect of artificial wet-
ting and rainfall on the larval diapause of
a stem borer, Busseola fusca in western
Kenya. Entomol. Exp. Appl. 48:263–67
AFRICAN STEM BORERS
in a stem borer, Busseola fusca. Entomol.
Exp. Appl. 57:151–55
137. Okuda T. 1991. Factors inducing and ter-
minating larval diapause in a stem borer,
Busseola fusca in western Kenya. Jpn.
Agric. Res. Q. 25:47–54
138. Oloo GW, Ogeda K. 1990. The incidence
of Chilo partellus (Swinh.) (Pyralidae)
and the contribution of natural enemies to
its mortality under intercropping systems
in Kenya. Insect Sci. Appl. 10:243–51
139. OmwegaCO, KimaniSW, OverholtWA,
Ogol CKPO. 1995. Evidence of the es-
tablishment of Cotesia flavipes Cameron
(Hymenoptera: Braconidae) in continen-
tal Africa. Bull. Entomol. Res. 85:525–30
140. OmwegaCO, OverholtWA, MbapilaJC,
Kimani-Njogu SW. 1997. Establishment
doparasitoid of Chilo partellus Swinhoe
(Lepidoptera: Pyralidae) in northern Tan-
zania. Afr. Entomol. 5:71–75
141. Overholt WA. 1998. Biological control.
See Ref. 150, pp. 349–62
142. OverholtWA, Ngi-SongAJ, KimaniSK,
Mbapila J, Lammers P, Kioko E. 1994.
Ecological considerations of the intro-
ductions of Cotesia flavipes Cameron
(Hymenoptera: Braconidae) for biologi-
cal control of Chilo partellus (Swinhoe)
(Lepidoptera: Pyralidae), in Africa. Bio-
control News Inf. 15:19–24
143. Overholt WA, Ngi-Song AJ, Omwega
et al. 1997. A review of the introduc-
tion and establishment of Cotesia flavipes
Cameron (Hymenoptera: Braconidae) in
stemborers. Insect Sci. Appl. 17:19–35
144. Overholt WA, Ochieng JO, Lammers P,
methods for Cotesia flavipes Cameron
(Hymenoptera: Braconidae), a parasitoid
of tropical gramineous stemborers. Insect
Sci. Appl. 15:253–59
145. Overholt WA, Songa JM, Ofomata V,
Jeske J. 2000. The spread and ecologi-
cal consequences of the invasion of Chilo
partellus (Swinhoe) (Lepidoptera: Cram-
bidae) in Africa. In Invasive Species in
Eastern Africa: Proc. Workshop ICIPE,
2000, ed. EE Lyons, SE Miller, pp. 52–
58. Nairobi: ICIPE Sci. Press. 108 pp.
146. P¨ ats P. 1992. Reproductive biology of the
cereal stemborers Chilo partellus. PhD
thesis. Swed. Univ. Agric. Sci., Uppsala,
Swed. 97 pp.
147. P¨ ats P. 1996. Management of crop resi-
dues to reduce the aestivating population
148. P¨ ats P, Ekbom B, Scovg˚ ard H. 1997.
Influence of intercropping on the abun-
dance, distribution and parasitism of
Chilo spp. (Lepidoptera: Pyralidae) eggs.
Bull. Entomol. Res. 87:507–13
149. PoinarGO, PolaszekA.1998.Nematoda,
fungi, protozoa, bacteria and viruses. See
Ref. 150, pp. 283–93
150. Polaszek A. 1998. African Cereal Stem
Borers: Economic Importance, Taxon-
omy, Natural Enemies and Control.
Wallingford, UK: CABI. 530 pp.
151. PolaszekA, WalkerAK.1991.TheCote-
sia flavipes species complex: parasitoids
of cereal stemborers in the tropics. Redia
152. Rajabalee MA, Govendasamy M. 1988.
Host specificity and efficacy of Apanteles
flavipes (Cam.) and A. sesamiae (Cam.)
(Hymenoptera: Braconidae) parasites of
sugar cane moth borers in Mauritius. Rev.
Agric. Sucr. Ile Maurice 67:78–80
153. Revington J. 1986. This borer spreads
rapidly through crops of maize and
sorghum on the Highveld, but it can
be controlled. Farmer’s Weekly, Oct. 24.
154. Revington J. 1987. Use of a synthetic
sex pheromone in forecasting attacks on
maize by maize stalkborer Busseola fusca
(Fuller) (Lepidoptera: Noctuidae). MSc
thesis. Univ. Witwatersrand, Johannes-
burg. 204 pp.
155. Risch SJ, Andow D, Altieri MA. 1983.
KFIR ET AL.
Agrosystems diversity and pest con-
trol: data, tentative conclusions and new
research directions. Environ. Entomol.
156. Sagnia SB. 1983. Possible integrated pest
management tools for the effective con-
trol of cereal stem borers in the Gambia.
Insect Sci. Appl. 4:217–19
157. Sallam MN,Overholt WA,
2001. Dispersal of the exotic parasitoid
Cotesia flavipes in a new ecosystem. En-
tomol. Exp. Appl. 98:211–17
sures. Experiment Station Information
Sheet: SASA. Mount Edgecombe, Natal:
S. Afr. Sugar Assoc.
borers (Lepidoptera: Pyralidae and Noc-
tuidae) in Ghana. Z. Angew. Entomol.
160. Scheltes P. 1978. Ecological and phys-
iological aspect of aestivation-diapause
in the larvae of two Pyralid stalk bor-
ers of maize in Kenya. PhD thesis. Land-
bouwhogeschool, Wageningen. 110 pp.
161. Scheltes P. 1978. The condition of the
host plant during aestivation-diapause of
the stalk borers Chilo partellus and Chilo
in Kenya. Entomol. Exp. Appl. 24:679–88
162. Scovg˚ ard H, P¨ ats P. 1996. Effects of
intercropping on maize stemborers and
their natural enemies. Bull. Entomol. Res.
163. Seshu Reddy KV. 1983. Sorghum stem
borers in eastern Africa. Insect Sci. Appl.
164. Seshu Reddy KV. 1985. Integrated ap-
ers. Proc. Int. Sorghum Entomol. Work-
shop, Tex., 1984, pp. 205–15. Patancheru,
165. Seshu Reddy KV. 1988. Assessment of
on-farm yield losses in sorghum due to
insect pests. Insect Sci. Appl. 9:679–85
166. Seshu Reddy KV. 1990. Cultural control
of Chilo spp. in graminaceous crops. In-
sect Sci. Appl. 11:703–12
167. Seshu Reddy KV. 1998. Integrated pest
management. See Ref. 150, pp. 39–45
168. Seshu Reddy KV. 1998. Maize and
sorghum: East Africa. See Ref. 150, pp.
169. Seshu Reddy KV, Sum KOS. 1991. De-
termination of economic injury level of
the stem borer, Chilo partellus (Swinhoe)
in maize, Zea mays L. Insect Sci. Appl.
170. Seshu Reddy KV, Sum KOS, Lubega
MC. 1989. Empirical models for predict-
ing yield loss in sorghum caused by Chilo
partellus (Swinhoe). Discov. Innov. 1:90–
171. S´ etamou M, Schulthess F, Bosque-P´ erez
NA, Thomas-Odjo A. 1993. Effect of
of Sesamia calamistis Hampson (Lepi-
doptera: Noctuidae). Bull. Entomol. Res.
172. S´ etamou M, Schulthess F, Bosque-P´ erez
NA, Thomas-Odjo A. 1995. The effect
of stem and cob borers on maize sub-
ments, with special reference to Sesamia
calamistis Hampson (Lepidoptera: Noc-
tuidae). Entomol. Exp. Appl. 77:205–10
173. Shami S, Mohyuddin AI. 1987. Host
selection by Indonesian strain of Apan-
teles flavipes (Cam.) and its suitability
for various graminaceous borers in Pak-
istan. Proc. Annu. Conv. Pak. Soc. Sugar
Technol., 23rd, Rawalpindi, pp. 286–91.
Rawalpindi: Pak. Soc. Sugar Technol.
174. Shanower TG, Schulthess F, Bosque-
P´ erez NA. 1994. The effect of larval
diet on the growth and development
of Sesamia calamistis Hampson (Lepi-
doptera: Noctuidae) and Eldana saccha-
rina Walker (Lepidoptera: Pyralidae). In-
sect Sci. Appl. 14:681–85
175. Shanower TG, Schulthess F, Gounou S.
1991. Distribution and abundance of
some stem and cob borers in Benin. Plant
176. Sibanda S. 1985. The use of sorghum and
AFRICAN STEM BORERS
millets for feeding livestock. Proc. Reg.
Workshop Sorghum Millets for S. Afr.,
2nd, Gaborone, pp. 228–47. Gaborone,
177. Sithole SZ. 1987. Maize insect pests
in Zimbabwe. Toward Insect Resistant
Maize for the Third World. Proc. Int.
Symp. Methodol. Developing Host Plant
Resistance Maize Insects, Mexico, pp.
286–88. Mexico City: Centro Int. Majo-
ramiento Maiz y Trigo
southern Africa. Int. Workshop Sorghum
Stem Borers, Patancheru, 1987, ed. KF
Nwanze, pp. 41–47. Patancheru, India:
179. Sithole SZ. 1989. The effect of date of
planting on shootfly and stem borer in-
festations on sorghum. Proc. Reg. Work-
1986, pp. 174–83. Patancheru: ICRISAT
180. Sithole SZ. 1990. Status and control of
the stem borer, Chilo partellus (Swin-
hoe) (Lepidoptera: Pyralidae) in southern
Africa. Insect Sci. Appl. 11:481–88
181. Skoroszewski RW,
1987. The release of Apanteles flavipes
(Cameron) (Hymenoptera: Braconidae)
against stalk-borers of maize and grain-
S. Afr. 50:249–55
182. Smith JW Jr, Wiedenmann RN, Over-
holt WA. 1993. Parasites of Lepidop-
teran Stemborers of Tropical Gramineous
Plants. Nairobi: ICIPE Sci. Press. 89 pp.
183. Songa JM. 1999. Distribution, impor-
tance and management of stemborers
(Lepidoptera) in maize production sys-
tems of semi-arid eastern Kenya with em-
phasis on biological control. PhD thesis.
Kenyatta Univ., Nairobi. 251 pp.
184. Starks KJ.1969. Some cereal crop insects
in east Africa. East Afr. Agric. For. Res.
Organ., Serere Res. Stn., Uganda
185. Swaine G. 1957. The maize and sorghum
stalkborer, Busseola fusca (Fuller), in
peasant agriculture in Tanganyika terri-
tory. Bull. Entomol. Res. 48:711–22
Van Hamburg H.
Heterocera. Entomologist 65:1241–49
187. Tams WHT, Bowden J. 1953. A revision
of the African species of Sesamia Gue-
nee and related genera (Agrotidae—
Lepidoptera). Bull. Entomol. Res. 43:
188. Taneja SL, Nwanze KF. 1989. Assess-
ment of yield loss of sorghum and
pearl millet due to stem borer damage.
Int. Workshop Sorghum Stem Borers,
Patancheru, 1987, ed. KF Nwanze, pp.
95–104. Patancheru, India: ICRISAT
189. Tavakilian G. 1977. Le genre Chilo en
Cˆ ote d’Ivoire. Cah. ORSTOM S´ er. Biol.
190. Tran M. 1977. Morphologie des pi` eces
g´ enitales et nervation alaire des prin-
cipales pyrales foreurs du riz en Cˆ ote
d’Ivoire. Description de quelques Hym´ e-
nopt` eres parasites. Cah. ORSTOM S´ er.
191. Ukwungwu MN. 1985. Effect of nitrogen
and carbofuran on gall midge (GM) and
ria. Int. Rice Res. Newsl. 10:19–20
192. Unnithan GC, Seshu Reddy KV. 1989.
Incidence, diapause and carryover of the
cereal stem borers in Rusinga Island,
Kenya. Trop. Pest Manag. 35:414–19
193. Usua EJ. 1973. Induction of diapause in
the maize stemborer, Busseola fusca. En-
tomol. Exp. Appl. 16:322–28
194. Usua EJ. 1974. Observations on the phys-
iology of diapause and non-diapause lar-
tuidae). Bull. Entomol. Res. 63:513–18
195. Van den Berg J, Nur AF. 1998. Chemical
control. See Ref. 150, pp. 319–32
196. Van den Berg J, Nur AF, Polaszek A.
1998. Cultural control. See Ref. 150, pp.
197. Van den Berg J, Van Rensburg JBJ. 1991.
Infestation and injury levels of stem bor-
ers in relation to yield potential of grain
sorghum. S. Afr. J. Plant Soil 8:127–
198. Van den Berg J, Van Rensburg JBJ. 1991.
KFIR ET AL.
The effect of plant density on the injuri-
ousness of Busseola fusca (Fuller) (Lepi-
doptera: Noctuidae) in grain sorghum. S.
Afr. J. Plant Soil 8:85–87
199. Van den Berg J,
Pringle KL. 1990. Damage caused by
Chilo partellus (Swinhoe) (Lepidoptera:
Pyralidae) to various cultivars of grain
sorghum, Sorghum bicolor (L.) Moench.
S. Afr. J. Plant Soil 7:192–96
200. Van den Berg J,
tuidae) and Chilo partellus (Lepidoptera:
Pyralidae) on grain sorghum. Bull. Ento-
mol. Res. 81:137–42
201. Van den Berg J, Van Rensburg JBJ, Van
der Westhuizen MC. 1991. The effect of
single and mixed populations of Busse-
ola fusca (Lepidoptera: Noctuidae) and
Chilo partellus (Lepidoptera: Pyralidae)
on damage to grain sorghum. J. Entomol.
Soc. S. Afr. 54:231–42
202. Van den Berg J, Van Rensburg JBJ, Van
der Westhuizen MC. 1993. Status of
resistance to Chilo partellus (Lepido-
ptera: Pyralidae) in South African grain
sorghum hybrids. S. Afr. J. Plant Soil 10:
203. Van Hamburg H. 1976. Die bionomie
en ekonomiese belangrikheid van die
graansorghum-stamruspe, Chilo partel-
lus (Swinhoe) (Lepidoptera: Pyralidae).
DSc thesis. Univ. Pretoria, Pretoria. 299
204. Van Hamburg H. 1979. The grain-
sorghum stalk-borer, Chilo partellus
(Swinhoe) (Lepidoptera: Pyralidae): sea-
S. Afr. 42:1–9
205. Van Hamburg H. 1987. A biological con-
crops with special reference to the con-
trol of stalkborers. Tech. Commun., Dep.
Agric. Water Supply, Rep. S. Afr. 212:52–
206. Van Rensburg GDJ, Bate R. 1987. Pre-
Van Rensburg JBJ,
Van Rensburg JBJ,
and distribution of the stalk borers Busse-
ola fusca and Chilo partellus Tech. Com-
mun., Dep. Agric. Water Supply, Rep. S.
207. Van Rensburg JBJ. 1992. Evaluation of
pheromone trapping systems in rela-
tion to light trap captures of the maize
stalk borer Busseola fusca (Fuller) (Lep-
idoptera: Noctuidae). S. Afr. J. Plant Soil
208. Van Rensburg JBJ. 1997. Seasonal moth
flight activity of the maize stem borer,
Busseola fusca (Fuller) (Lepidoptera:
Noctuidae) in small farming areas of
South Africa. Appl. Plant Sci. 11:20–23
209. Van Rensburg JBJ,
Carnegie AJM, Eulitz EG, Rust D. 1989.
Stemborers. In Crop Pests in Southern
Africa, ed. AC Myburgh, 4:64–73, Bull.
416. Pretoria: Plant Prot. Res. Inst., Dep.
Agric. Water Supply. 156 pp.
210. Van Rensburg JBJ, Walters MC, Gilio-
mee JH. 1985. Geographical variation in
the seasonal moth flight activity of the
211. Van Rensburg JBJ, Walters MC, Gilio-
mee JH. 1988. Plant population and cul-
tivar effects on yield losses caused by the
maize stalk borer, Busseola fusca (Lepi-
doptera: Noctuidae). S. Afr. J. Plant Soil
212. Van Rensburg JBJ, Walters MC, Gilio-
J. Entomol. Soc. S. Afr. 52:105–8
213. VanRensburgNJ, VanHamburgH.1975.
Grain sorghum pests: an integrated con-
Entomol. Soc. S. Afr.
214. Waage JK. 1990. Ecological theory and
the selection of biological control agents.
In Critical Issues in Biological Control,
ed. M Mackauer, LE Ehler, J Roland, pp.
135–58. Andover, UK: Intercept. 330 pp.
AFRICAN STEM BORERS Download full-text
215. Walker PT. 1960. The relation between
infestation by the stalk borer Busseola
fusca, and yield of maize in East Africa.
Ann. Appl. Biol. 48:780–86
216. Walker PT, Hodson MJ. 1976. Devel-
opments in maize stem-borer control in
East Africa, including the use of insecti-
cide granules. Ann. Appl. Biol. 84:111–
217. Warui CM, Kuria JN. 1983. Population
incidence and the control of maize stalk-
borers Chilo partellus (Swinh.) and Chilo
orichalcociliellus Strand and Sesamia
calamistis Hmps. in Coast Province,
Kenya. Insect Sci. Appl. 4:11–18
218. Way MJ, Kfir R. 1997. The stem borer
Chilo partellus (Swinhoe) in sugarcane
in southern Africa. Afr. Entomol. 5:170–
219. Wiedenmann RN, Smith JW Jr. 1993.
Functional response of the parasite Cote-
sia flavipes (Hymenoptera: Braconidae)
at low densities of the host Diatraea sac-
charalis (Lepidoptera: Pyralidae). Envi-
ron. Entomol. 22:848–58
220. Wiedenmann RN, Smith JW Jr. 1997.
Novel associations and importation bio-
logical control: the need for ecological
and physiological equivalencies. Insect
Sci. Appl. 17:51–60
221. Wilkinson H. 1939. Entomol. Sect. Annu.
Rep. 2. Dep. Agric., Kenya
222. Williams JR. 1983. The sugar cane stem
borer (Chilo sacchariphagus) in Mauri-
tius. Rev. Agric. Sucr. Ile Maurice 62:5–
223. Yitaferu K, Gebre-Amlak A. 1994. Phe-
nology and survival of the maize stalk
borer, Busseola fusca in eastern Ethiopia.
Insect Sci. Appl. 15:177–84
of food and industrial crops of Africa and
their economic importance. In Biological
Control: A Sustainable Solution to Crop
Pest Problems in Africa, ed. JS Yanninek,
HR Herren, pp. 51–60. Ibadan, Nigeria:
Int. Inst. Trop. Agric. 210 pp.
225. Youm O, Beevor PS. 1995. Field evalua-
tion of pheromone-baited traps for Coni-
esta ignefusalis (Lepidoptera: Pyralidae)
in Niger. J. Econ. Entomol. 88:65–69
226. Youm O, Gilstrap FE. 1994. Habitat site
selection, crop damage and oviposition
preference by Coniesta (=Haimbachia)
ignefusalis (Lepidoptera: Pyralidae). Int.
J. Pest Manag. 40:231–36
227. Youm O, Mamalo AK,
1993. Bio-ecology and integrated man-
agement of the millet stem borer (Lep-
idoptera: Pyralidae): a review and high-
lights of current research at the ICRISAT
Sahelian Center. Proc. Reg. Pearl Millet
Workshop, Niamey, 1990, pp. 55–63. Ni-
amey, Niger: ICRISAT Sahelian Cent.
228. Zhou G, Baumgartner J, Overholt WA.
2001. Impact of an exotic parasitoid on
stemborer (Lepidoptera) populations dy-
namics in Kenya. Ecol. Appl. 11:1554–62