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Tropical Pest Management

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Review of the biology and control of the Asian

corn borer, Ostrinia furnacalis (Lep: Pyralidae)
a a
D. M. Nafus & I. H. Schreiner
a
Agricultural Experiment Station , University of Guam , Mangilao, Guam , 96923
Published online: 13 Nov 2008.

To cite this article: D. M. Nafus & I. H. Schreiner (1991) Review of the biology and control of the Asian corn borer,
Ostrinia furnacalis (Lep: Pyralidae), Tropical Pest Management, 37:1, 41-56, DOI: 10.1080/09670879109371535

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 TROPICAL PEST MANAGEMENT. 1991, 37(1), 41-56

Review of the biology and control of the Asian corn borer, Ostrinia
furnacalis (Lep: Pyralidae)

(Keywords: Ostrinia furnacalis, corn borer, biology, control, parasitoids, parasites, biological control)

D. M. NAFUS and I. H. SCHREINER

Agricultural Experiment Station, University of Guam, Mangilao, Guam 96923

Abstract. The biology and control of the Asian corn borer, Ostrinia and Spilodes kodzukalis. Many other variants and combina-
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furnacalis Guenée, is reviewed. This insect is distributed from China tions of these names are present in the literature.
to Australia and the Solomon Islands. In northern parts of its range
the moths have one or a few generations per year, but in the tropics,
generations are continuous and overlapping. The caterpillars 2. Distribution
can cause severe yield losses in corn, both by damage to the kernels
and by feeding on the tassels, leaves, and stalks. Survival and In Asia the range of the ACB extends from India eastward
growth of the caterpillar is highest on the reproductive parts of the through Southeast Asia and north into China, Korea and
plant. Other economic plants attacked include bell pepper, ginger Japan (Mutuura and Monroe, 1970; Lewvanich, 1973). It is
and sorghum. Recently, the Asian corn borer appears to have
become an important pest of cotton. A number of wild grasses are
present in Australia and on a number of islands in the
also used as hosts. Information on predators and parasites is Western Pacific including Java, Sulawesi, the Philippines,
summarized. Egg parasitoids are relatively effective in some Borneo, New Guinea, Solomons, and Western Micronesia
tropical areas, but no significant larval or pupal parasites have been (Mutuura and Monroe, 1970).
found. In temperate parts of the Asian corn borer's range, larval
parasites are more abundant, having their greatest impact during
the diapausing generation. A variety of techniques have been 3. Yield losses
used to control the borer. Insecticides are used in many countries,
but are not totally reliable. Augmentative release of egg parasitoids The ACB has caused severe yield losses to corn in some
is widely used in China and has been tested elsewhere. Mechanical countries, but in others it is a minor pest. Sanchez (1971)
removal of the tassels can reduce borer populations 50%, and this
reported yield losses of 20-80% in the Philippines. High yield
procedure is used in the Philippines. Field corn varieties
which are relatively resistant to the ACB have also been found. losses have also been reported from the Marianas, up to
100% (Schreiner and Nafus, 1987) and Taiwan, 95% (Tseng,
1981). Arunin (1964) listed ACB as one of the two worst pests
of corn in Thailand, but W. Young (1979) stated that it was
sporadic, being relatively uncommon except at lower
In the Asian-Western Pacific region, the Asian corn borer
elevations in the central plains and at Sisaket and Uthong.
(ACB), Ostrinia furnacalis Guenée (Lep: Pyralidae), is prob-
Apparently the borer has declined in importance in Thailand
ably the most serious insect pest of corn, Zea mays, and
as the incidence of growing heavily treated cotton has
only maize downy mildew is potentially more severe in
decreased, and other crops, primarily sugar cane, have
Southeast Asia. The larvae feed on virtually all parts of the
been substituted. The decline of the ACB was suggested to
plant, but the worst damage is caused by boring in the ears
be due to an increase in natural enemies in the less heavily
and stalk. Since O. furnacalis has been recognized as a
treated crops (Anon, 1974). In Papua New Guinea the ACB
distinct species from the European corn borer, Ostrinia
is a severe problem in subsistence gardens In certain
nubilalis (Hubner), research on the ACB has intensified.
locations (O'Sullivan and Bourke, 1975). In plots at the
Unfortunately, much of the research is not easily accessible.
research centre in Papua New Guinea, G. Young (1979)
These factors have inhibited exchange of information, and
found an average of 16 holes per plant. Yield losses of
prompt this review.
10-40% have been reported from Indonesia (Chu, 1979).

1. identification 4. Economic thresholds

Originally, the ACB was described as Botys furnacalis by
Guenée in 1854. Subsequently it was re-described by a
series of authors or repeatedly misidentified as the
European corn borer, O. nubilalis. Much of the literature
prior to 1966 treats it as that species. Mutuura and Monroe
(1970) revised the genus Ostrinia, confirming the status of
O. furnacalis as a valid species separate from the European
corn borer and synonymizing it with Botys (Pyrausta)
damoalls, B. (Pyrausta) salentialis, P. polygoni, P. vastatrix.
Economic thresholds have not been satisfactorily de-
termined for either grain or sweet corn. In the Philippines
yield losses of grain were reported to be 0-95% per tunnel
(Medrano and Raros, 1975), but in Thailand much higher
losses were found by Prachuabmoh (1975a). His data show
a linear increase in yield of grain of approximately 30% as
the number of ACB holes per stalk declined from about 3 to
0-75 per plant. G. Young (1979) thought that corn could
tolerate up to five holes per stalk without significant loss of

0143-6147/91 $3.00 © 1991 Taylor and Francis Ltd

 42 D. M. Nafus and I. H. Schreiner

yield. Mekongsee et al. (1979) and Mekongsee and 6. Development and diapause
Kongkanjana (1980) suggested treating corn when the
The length of the life cycle varies with host, diet (Table 1)
number of masses reached 15/100 plants as no yield losses
and weather conditions (Buligan, 1929; Lee etal., 1980; Saito
were found at these or lower densities. Mekongsee et al.
and Oku, 1985). Saito and Oku (1985) reported the
(1979) found an increase in the number of borers per plant
temperature threshold for development in Japan was 6-8°C
as the number of egg masses increased, but the effect was
and a total of 457-3 degree-days were necessary to complete
not linear. At low densities, 0-05 masses per plant resulted
development. They reported a regression to determine
in 0-24 holes per plant, or about 4-8 holes per mass. At
length of the larval period: V= 0005 Τ- 0Ό69, where Vis
higher densities only two holes resulted from each mass,
development velocity (1/days required for development) and
although the overall number of holes per plant increased.
7" is temperature as °C heat unit accumulation above the
In the Philippines, current recommendations are that corn
threshold point. Thresholds and degree-days have not been
should be treated when larval densities are greater than
reported for other areas as most rearing has been done
one per plant (Morallo-Rejesus and Javier, 1985). A sample
under ambient temperature. In tropical areas, and in the
of 100 plants was suggested. No information was presented
warm part of the summer in temperate zones, thé larval
to relate these figures to yield loss.
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stage lasts 16-18 days and has five or six instars. Eggs
We strongly suspect that yield losses do not increase usually develop in 3-5 days (Table 1).
linearly as ACB densities increase when a wide range of
The number of instars is variable, and is affected by host,
borer densities is considered. Data on yield losses in
season, and general conditions for growth. Schreiner et al.
relation to the timing of the infestation, plant density, and
(1989) found that 17% of the larvae reared on corn ears had
other factors are needed.
six instars, with the remainder having only five instars. On
In sweet corn tolerance levels are lower, since even a
Sorghum halepense 25% of the larvae had six instars, and
single borer in an ear can make it unmarketable. Typically
on other hosts higher percentages were found. On less
about half of the borers in a plant will be found in the ear.
preferred hosts, survival was poor and some larvae required
Prachuabmoh (1975b) got no yield at densities of nine or
six instars, suggesting that poor-quality food results in extra
more holes per plant. At lower densities the number of
instars. In Korea, Lee etal. (1980) found seven instars in the
harvested ears increased. At a damage level of four holes
first generation when temperatures were low. At higher
per plant, 1Ό5 ears per plant were harvested.
temperatures there were six instars.
In cotton in China, the economic thresholds for the first,
The pupal period fasts from 6 to 8 days (Table 1). Adults
second and third generations were 2-8, 1-1 and 3-1 egg
emerge between 4 and 8 p.m., and mate 19-24 h later.
masses per 100 plants respectively (Liu and Yuan, 1981).
Oviposition begins 1 or 2 days after mating (Buligan, 1929;
On ginger, no information is available on economic
Camarao, 1976b). Adults are active throughout the night, but
thresholds. However, three or four insecticidai treatments
the peak of activity is in the early evening and in the morning
applied weekly after the first injury reduced the proportion
before sunrise (Calora etal., 1965). Adult females live from
of damaged stalks from 33% to 2%, and increased yield
4 to 11 days in the laboratory (Table 2), but improved
about 14% (Ichihara, 1975).
techniques have now increased longevity (Hirai and
Legacion, 1985).
In the northern part of its range the ACB diapauses, but
5. Life cycle: number of generations not in tropical areas. In northern Japan, larvae in both the
first and the second generation enter diapause. The
In tropical areas the ACB is continuously active (Buligan,
number of first-generation larvae which diapause is
1929; Camarao, 1976a, b; Nafus and Schreiner, 1986a) and
inversely related to heat accumulation, and varies between
can have from 9 to 12 generations per year (Camarao, 1976b;
22% and 88%, depending on weather conditions (Saito and
Buligan, 1929; Hua, 1966; Yunus and Hua, 1969). Generations
Oku, 1985). The incidence of diapause was the most
are overlapping with continuous oviposition throughout the
important factor in determining the population size of the
crop season (Hua, 1966; Meksongsee et al., 1979; Tseng,
second generation. Mortality associated with prédation and
1981). In subtropical and tropical areas with distinct wet and
parasitism was relatively unimportant (Saito and Oku, 1985).
dry seasons, populations and damage levels are highest
In ACBs collected from Qinshui province in China (35-36°N)
during or immediately following the wet season (Tseng,
the critical day length was 14-5 h at 25CC (Gong etal., 1984).
1981; Morallo-Rejesus, 1985). In temperate areas, ACB
At shorter daylengths diapause was induced. Obviously,
populations undergo diapause and there are fewer, more
depending on latitude, there will be variation in the critical
synchronized generations per year. In northern Japan, Saito
temperatures and daylengths necessary to initiate diapause.
and Oku (1985) reported two generations in corn, one in
Research in this area is needed.
June/July and a second in late August and September. In
southern Japan (Ichihara, 1975 and South Korea (Lee et al.,
1980) there are three generations. In northern Japan eggs
are first found in late June and early July and then again in
7. Sex pheromones
mid-August. The first generation takes about 1 month for In 1980 the sex pheromone of O. furnacalis was reported
development and the second slightly longer (Saito and Oku, to be a mixture of (2)- and (£)-12-tetradecen-1-ol acetate by
1985). In Korea the first generation takes about 45 days and several groups of workers (Ando et al., 1980; Cheng ef a/.,
the next two take about 1 month each (Lee ef a/., 1980). 1980; Klun et al., 1980; Cheng et al., 1981). Exact ratios of
 Biology and control of Asian corn borer 43

Table 1. Development times of the Asian corn borer on different hosts or host parts in various countries

Guam1 Korea 2 Malaya 3 Philippines 4 Philippines 5 Thailand 6 Guam 1 Philippines 7 Philippines 4

Host Corn Corn Corn Corn Corn Corn Sorghum cotton Vigna
ears ? stalk stalk stalk halpense pods
Temperature °C Ca. 28 Ca. 21 Ca. 27 N.R." N.R. N.R. N.R. ca. 28 N.R. N.R.
Sex Both Both Both Both Male Female Both Both Both Male Female Male Female

Instar 1 3-4 30 22 3 30 3-6 3-6 3-3 4-7 3-3 3-5 2-4 2-4
Instar 2 30 2-8 20 3 2-2 2-3 3-6 30 20 3-7 3-8 2-3 20
Instar 3 20 3-4 20 2 20 2-1 3.8 3-5 30 3-8 3-9 2-1 2-7
Instar 4 2-8 3-8 2-7 4 2-1 2-3 3-9 3-9 3-3 3-4 3-6 2-3 2-8
Instar 5 4-8 4-8 2-5 6 5-4 60 4-6 3-3 7-1 3-7 3-9 7-6 7-7
Instar 6 50° 50 60 — — — 11-5 — 10Ό 1 0 — — — —
Instar 7 — 7-5 — — — — — — — — — — —

Total larval 16-2 30-4 17-4 18 13-7 16-4 28.9 1 1 17-1 220 17-9 18-7 16-7 17-5
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Pupa 8-6 9-4 6-7 6 6-8 7-2 8-3 5-3 8-4 3-5 5-8 6-8 6-3
Preovi positional — — — 2 — 20 1-8 — — — — —
period
Adult longevity — 10-9 8-2 6 — 70 8-5 4-2 — 3-5 4-1 — —
Total — 45-8 29-3 27 25-5 28-6 41-712 30-4 — 26-9 280 — 28-8
1
Schreiner and Nafus, unpublished data.
2
Lee etal., 1980.
3
Hua, 1966; Yunus and Hua, 1969.
'Camarao, 1976b.
5
Buligan, 1929.
"Areekul el ai, 1964.
7
Valdez and Adalla, 1983,-
* N.R. indicates temperature is not reported and is assumed to be ambient.
8
Ca. 17% of the larvae moulted to sixth Instar. The rest only had five instars.
10
Ca. 25% of the larvae moulted to sixth instar. The rest only had five instars.
11
This is a seasonal average. The larval period varies between 18 and 41 days with the longest period in Oct./Dec. and the shortest in June/July.
12
Total life cycle varies between 29 and 56 days depending on temperature. Temperatures are not given.

Table 2. Fecundity of Asian corn borer and hatching times for eggs

Country Average Number Total eggs Incubation Reference

number masses/ female days
ΙΑΜΑΙΛ

eggs/mass female
Range Average

Thailand 10-25 2-12 300-600 3-4 Lewvanich, 1973

Philippines 36 — 115-1193 465 4-5 Buligan, 1929
Philippines 24 18 up to 579 313 3 Camarao, 1976a
Malaya 27 24 230-1474 638 3 Yunus and Hua, 1969
Malaya 30 14 — 400 3-4 Hua, 1966
Korea (spring) — — 242-810 602 40 Lee etal., 1980
Korea (summer) — — 605-1128 817 3-2 Lee etal., 1980

the two enantiomers varied from ca. 1:1 in the Philippines
(Klun etal., 1980), to 53:47 from China (Cheng etal., 1981)
and 3:2 from Japan (Ando er al., 1980). Attraction of moths
to traps using either individual components or mixtures of
the pheromone was reported from China (Klun et al., 1980;
Cheng etal., 1981; Jiang and Klun, 1981; Cheng etal., 1982).
An additional component in ovipositor washes, tetradecyl
acetate, was not demonstrated to possess any attractiveness
(Cheng ei ai., 1981 ; Ren er al., 1982) and was not thought to
be an important component of the pheromone (Klun et al.,
1980; Cheng etal., 1981). In the Philippines (Jackman etal.,
1983,1984) and in Japan (Ando etal., 1980), researchers did
not get attraction to pure synthetic pheromone, and felt that
there was an additional, unidentified component. Jackman
er al. (1984) reported an undescribed attractant soluble in
acetone. They suggested that the acetone-soluble compo-
nent was a long-range attractant, and the previously isolated
heptane-soluble components were short-range attractants.
Further work on the sex pheromone is needed.
8. Oviposition: in relation to growth cycle
The first eggs are normally found when the crop is
between 2 weeks and 1 month old (Yunus and Hua, 1969;
Camarao, 1976b; Nafus and Schreiner, 1987). Oviposition
increases sharply about 1 month after planting, and is
 44 D. M. Nafus and I. H. Schreiner
highest during the mid-whorl to silk stages of the corn with
the peak at the tassel stage (Meksongsee et al., 1979). In
tropical areas, oviposition is continuous with either one
(Meksongsee and Kongkanjana, 1980; Nafus and Schreiner,
1986a) or two main peaks (Areekul era/., 1965; Hussein and
Kamaldeer, 1988). A high proportion of the eggs laid, even
late in the crop cycle, is probably by immigrants (Camarao,
1983; Nafus and Schreiner, 1987), and the number of masses
laid is a function of the attractiveness of the stage of the
plant (Nafus and Schreiner, 1987). Plants from the late whorl
through the silking stage, a period of 15-20 days, are most
attractive for oviposition (Cheu, 1941 ; Nafus and Schreiner,
1987). Plant tissues in these stages support the best ACB
growth and survival (Camarao, 1976b; Saito, 1980). Larger
plants are preferred for oviposition (Cheu, 1941 ; Medrano
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and Raros, 1975).
9. Fecundity
The eggs are laid in white, scale-like masses, which
average from 20 to 40 eggs (Table 2), although means as
high as 62 (Calora et al., 1965) eggs have been recorded.
Sizes of masses range from as few as two eggs to over 200
eggs (Buligan, 1929). Rearing conditions and host plants will
affect fecundity. Females reared on corn normally lay
between 313 and 817 eggs (Table 2), although as many as
1474 eggs can be laid (Yunus and Hua, 1969). In the
laboratory an average of around 600 eggs per female is
possible, but unusual. On other hosts, fecundity is lower.
Females reared on bush sitao laid an average of 187 eggs
(Camarao, 1976b), and on cotton only 63 eggs (Valdez and
Adalla, 1983).
10. Oviposition sites
The oviposition sites on corn shift as the plant grows and
changes phenologically. The majority of the eggs are laid
underneath the leaves (Buligan, 1929; Hua, 1966; Yunus and
Hua, 1969) near the midrib (Camarao, 1976b). On Guam over
80% of the masses are under the leaves with the rest
primarily on the top surface (Nafus and Schreiner, 1987).
However in Malaysia, Hussein and Kamaldeer (1988) found
that nearly 40% of the eggs were on the upper surfaces of
the leaves. Masses are rarely found on the sheath or collar
(Nafus and Schreiner, 1987). Most of the eggs are laid on
leaves intermediate in age. Young, not fully emerged, leaves
from the whorl or very old leaves are less preferred
(Buligan, 1929; Calora et al., 1965; Camarao, 1976b; Nafus
and Schreiner, 1987). In consequence, the position of the
leaves with the largest number of eggs gradually shifts
upward as the plant grows. By the silk stage, most of the
eggs are on the 8th to 12th leaves. Eggs occasionally can
be found on the husk at this stage (Hua, 1966; Yunus and
Hua, 1969).
11. Biology of larvae on corn
Virtually all parts of the plant are eaten, depending on the
phenology of the plant and the larval instar (Buligan, 1929;
Hua, 1966; Areekul ef a/., 1964; Yunus and Hua, 1969;
Camarao, 1976b; Nafus and Schreiner, 1987). The most
important feeding sites are the whorl, tassel, ears, and stalk.
The eggs within a mass hatch synchronously, and the
larvae remain near the mass between 20 min and 5 h
(Buligan, 1929; Calora era/., 1965; Camarao, 1976b). At the
site of the mass the larvae often do not feed, except for
consuming parts of the egg shell, but occasionally they will
eat a little leaf tissue near the mass (Nafus and Schreiner,
1987). After leaving the hatching site the larvae wander until
they find a suitable feeding site. Larvae readily follow silk
trails and will cross silk bridges between plants. Some
larvae disperse on silk threads in the wind (Calora et al.,
1965; Hasse, 1981; Tseng, 1981). The final location depends
on the age of the plant, shifting from inside the whorl to the
tassel and ear, and finally to mostly in the ear, once the
tassel loses its pollen (Nafus and Schreiner, 1987).
As with the European corn borer (Brindley and Dicke,
1963), on young corn the larvae establish in the leaf spaces
in the whorl, where they feed on the developing leaf tissue
(Buligan, 1929; Areekul et al., 1964; Yunus and Hua, 1969;
Camarao, 1976b; Saito and Oku, 1976; Patanakamjorn et al.,
1978a; Nafus and Schreiner, 1987). A few larvae may
establish in folded exposed leaves (Nafus and Schreiner,
1987) or on sheath and collar tissue (Camarao, 1976b; Saito
and Oku, 1976), but this is uncommon.
The second and third instars continue to feed in the whorl
until the tassel develops. The tassel is the primary feeding
site until the pollen has been shed, after which it is less
attractive and the larvae move to other parts. In the tassel,
first and second instars bore into the florets and eat the
developing anthers. Third and fourth instars web florets
together or bore into the ramets or the main stalk. If
populations are very high, the entire tassel and the last two
leaves can be webbed together. A few larvae remain in the
tassel through pupation, but most leave (Nafus and
Schreiner, 1987).
At silking, a few third instars bore into the stalk or leaf
midribs, but most of the first to third instars are in the tassel
or associated with the ear (Saito and Oku, 1976; Nafus and
Schreiner, 1987). In the ear the young larvae are divided
between the silk and husk. On the husk most are found
inside, between the sheath leaves in the upper third of the
ear, although some are on the exposed leaves near the top
of the ear. In the silk the larvae are generally on the strands
outside the ear. Initially the larvae feed on individual
strands, but later the silk is webbed together, creating
clumps which are attractive to new arrivals, and may house
mixed ages. A few fourth-instar larvae remain on the silk,
mostly between the tip of the ear and the outer husk. Most
of the older larvae feed in the ear on developing kernels or
tunnel in the developing cob, but some also eat the husk or
stalk (Nafus and Schreiner, 1987). A few may bore into the
shank or feed in the leaf sheaths (Nafus and Schreiner,
1987).
Older larvae are often found in the stalk. The majority feed
in holes just above the nodes (Camarao, 1976b; Nafus and
Schreiner, 1987), although some feed in the sheath (Saito
and Oku, 1976). Normally only one larva is found per hole,
although more than one can be found if densities are high
 Biology and control of Asian corn borer 45

(Nafus and Schreiner, 1987). Pupation takes place inside the
hole with the cephalic end facing downward (Calóra ef a/.,
1965).
Survivorship (Saito and Oku, 1985) appears to follow the
same pattern as in the European corn borer (Brindley and
Dicke, 1963). Mortality is highest in the first instar and
declines in later instars. Egg mortality can also be high, but
varies between years from about 5% to 40% in Japan (Saito
and Oku, 1985). Camarao (1983) also found high mortality in
the larval stage and relatively less mortality in the egg and
pupal stages. She did not examine survival of the
early-instar larvae. Most mortality was from unknown
causes, and there was low prédation and parasitism.
Survivorship, plant age and ovipositional peaks are
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Which clues trigger movement of larvae from one feeding
site to another is unknown. Saito (1979) found that chemical
composition of the plant was not important, but light and
physical contact were. At low light levels, 150 lux, all instars
were attracted to light. As the light level increased to 9700
lux, first and second instars remained attracted, but third
instars and older became neutral. At 90 000 lux the oldest
larvae were negatively phototactic. Light, however, was less
important than physical contact with the substrate. Second,
third, and fourth instars preferred physical contact with the
plant and were attracted to sheltered areas regardless of
the light levels. First and fifth instars were less responsive
to shelter and continued to be attracted by light at certain
intensities.

generally correlated. Survivorship is poor on vegetative

stage plants, and improves on plants of reproductive age
(Camarao, 1976b; Saito, 1980). Camarao (1976) found that
20% of the larvae survived on plants 27-54 days old while
76% survived on plants older than 55 days. Saito (1980)
found that larvae fed on corn in the tassel stage had the
highest survival rates and the greatest body weights.
The timing of oviposition reflects the survivorship pattern
of the larvae. Few eggs are laid in whorl stage corn. As the
tassel develops, the number of eggs laid increases to a
maximum when pollen and silk are available. After silking,
fewer eggs are laid. Survival rates of larvae feeding on late
stage corn decline, and pupal weights of surviving larvae
also decrease, although both continue to be higher than on
vegetative stage corn (Saito, 1980). The length of the larval
development period also varies depending on the part of the
host the larvae feed on. Yunus and Hua (1969) reported that
larvae fed on ears rather than stalk had a faster development
time.
12. Alternative hosts
The ACB is polyphagous and feeds on plants in several
families. In addition to corn, several other crops are attacked
including ginger, sorghum, millet, bell pepper, cantaloupe,
rice, and rhubarb (Table 3). Recently strains of the ACB have
been reported on cotton in both China and the Philippines
(Liu and Yuan, 1981; Delattre and Pauly, 1982; Valdez and
Adalla, 1983). Around Shanghai, China, the ACB is now
considered to be more damaging on cotton than pink boll
worm, once the major pest (Liu and Yuan, 1981). Early
instars scrape the leaves while third instars and older larvae
bore in the stems, leaf petioles, flowers and squares. Stem
boring can cause wilting of the tips. Damaged bolls may
become secondarily infected and drop (Delattre and Pauly,
1982; Valdez and Adalla, 1983).
In Chiba prefecture, in Japan, the ACB is the most
important pest of ginger (Ichihara, 1975). The eggs are laid

Table 3. Hosts of the Asian corn borer excluding corn. Other hosts have been reported but recent revision of the genus Ostrinia requires
verification of the species of borer

Host Country Reference

Amaranthus sp. Australia D. Waterhouse, personal communication

Brachiaria mutica Guam, Mariana Isl. Schreiner et al., 1989
Capsicum sp. (bell pepper) Guam, Mariana Isl. Schreiner etal., 1989
Cucumis melo Tinian, Mariana Isl. Mutuura and Monroe, 1970
Emex australis Australia D. Waterhouse, personal communication
Gossypium spp. (cotton) Philippines Valdez and Adalla, 1983; Delattre and Pauly, 1982
China Liu and Yuan, 1981
Hibiscus cannabinus Thailand Lewvanich, 1973
Oryza sativa Papua New Guinea Young, G., 1979
Panicum miliaceum Japan Mutuura and Monroe, 1970
Phytolacca octandra Australia O. Waterhouse, personal communication
Polygonum sp. India Mutuura and Monroe, 1970
Polygonum tinctorium Japan Mutuura and Monroe, 1970
Populus deltoïdes angulata Australia D. Waterhouse, personal communication
Rheum rhaponticum Australia D. Waterhouse, personal communication
Saccharum officinarum Papua New Guinea Young, G., 1979
Guam, Marianas Isl. Schreiner et al.. 1989
Saccharum spontaneum Guam, Marianas Isl. Schreiner et al., 1989
Setaria italica Beauver Japan Mutuura and Monroe, 1970
Sorghum sp. Japan Saito, 1981
Sorghum halepense Guam, Marianas Isl. Schreiner et ai., 1989
Sorghum vulgäre Thailand Lewvanich, 1973
Philippines Buligan, 1929
Vigna sinensis Thailand Lewvanich, 1973
Zingiber officinale Japan Mutt ura and Monroe, 1970
 46 D. M. Nafus and I. H. Schreiner
on the underside of the leaves. The larvae bore into the leaf
veins and stalks, causing the shoots to die. In Chiba there
are three generations a year, but only the second generation
attacks ginger in significant numbers.
The ACB has several wild hosts (Table 3), the most
important being grasses (Schreiner et ai, 1989). On Guam,
in the absence of corn, high populations of ACB can be
maintained on wild hosts such as Sorghum halepense
(Schreiner et al., 1989). The moths can then move into corn
fields and completely destroy plantings.
The ACB is listed from many other hosts (Kuwayama,
1930), but these are not included in Table 3 because of the
problems in identifying ACB prior to the revision of Ostrinia.
Many are probably hosts, but need verification.
In India and Australia the ACB is listed from a variety of
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hosts, but does not attack corn (Mutuura and Monroe, 1970;
D. Waterhouse, personal communication).
13. Artificial growth media
Several synthetic media have been developed. Rangdang
(1971) developed a diet based on ground soy beans,
brewers' yeast, and wheat germ. He got about 70% survival
of the larvae. Patanakamjorn et al. (1978b) tested several
diets, but none was entirely satisfactory for mass rearing.
The best diet yielded about 80% larval survival and was
based on agar, corn-leaf powder, and mung beans. Adding
pollen enhanced survival. The ingredients in these diets are
not readily available. Saito and Nakayama (1981) found that
the ACB could be reared on the diet developed for
Spodoptera litura F. This diet is based on wheat germ and
kidney beans. However, only about 70% of the larvae
survived. Some improvements were reported by Yoshio and
Saito (1982) and Hirai and Legacion (1985), who substituted
mung bean for kidney beans and added low amounts of
methyl paraben, sorbic acid, and propionic acid (0-1-0-5%
dry weight of diet) to retard fungal growth. These additives
did not affect ACB growth or survival. Aureomycin was
added as a bactéricide. They obtained survival rates of 90%
for the larvae and 95% for the pupae. Egg production was
over 300 eggs per moth. The survival of ACBs on this diet
was better than on a corn-based diet used by Camarao
(1976b) and Ceballo and Morallo-Rejesus (1983). The mung
Table 4. Natural enemies known to attack immature stages of the Asian corn borer
Parasite, parasitoid, predator species Stage
attacked
Parasitoids
Hymenoptera: Trichogammatidae
Trichogramma australicum Girault (probably Egg
actually T. chilotreae)
Trichogramma chilonis Ishii Egg
Trichogramma chilotreae Nagaraja and Nagarkatti Egg
Trichogramma evanescens Westwood Egg
Trichogramma forcipiformis Zhang and Wang Egg
bean diet also avoids problems with high tryptophan, which
lowers survival, or low lysine.
Much of the improvement in egg production obtained by
Hirai and Legacion (1985) was due to improvements in the
design of the cage. Humidity is critical for good adult survival
and to induce oviposition. By using foam pads above and
below the cage, and plastic rather than screen sides, nearly
twice as many eggs were obtained.
14. Natural enemies
14.1. Parasitoids
Many species of parasitoids and predators are associated
with the ACB (Table 4), and there are probably many more.
In the 1930s American scientists, working under the
impression that the ACB was O. nubilalis, conducted
extensive surveys to find larval and pupal parasites in
Japan, Korea and northern China. Cartwright (1933) found
that the overall parasitization rate was low. The aggregate
parasitization by all species rarely exceeded 10% and
generally was less than 5%. On one occasion late in the
season, a sample of corn in Hokkaido had a parasitization
level of 35%. There were differences among crops. In
Hokkaido he found that less than 1 % of ACBs in hemp were
parasitized, but in corn and millet, parasitization was 4%.
Millet and corn have recently been re-confirmed as host
plants for O. furnacalis in Japan (Saito, 1981), but the
species of Ostrinia in hemp may not have been furnacalis. In
Kyushu, Cartwright (1933) found seasonal variation in the
species of parasitoids present. Trathala flavo-orbitalis
(Cameron) was dominant in June, Macrocentrus gifuensis
Ashmead in July and Lydella thompsoni Herting in August.
Saito and Oku (1985), working in Northern Japan, found that
M. gifuensis parasitized 6-7% of fifth-instar larvae in the first
generation on corn, 3-3% in the second generation, and
21-32% of overwintering larvae. In Korea, Lee er al. (1980)
reported about 6% of the first-generation, 14% of the
second-generation, and none of the third-generation larvae
were parasitized by a tachinid. Macrocentrus grandii
Goidanich killed about 1 % of the larvae of each generation.
Parasitization rates in the tropics are also low. In the
Philippines (Camarao, 1976a) and Papua New Guinea (G.
Country Percentage Reference
attacked
Thailand 15-100 Areekul era/., 1965
Taiwan Chiu and Chen, 1986
Guam 13-19 Nafus and Schreiner, 1986b
Philippines Alba, 1988
Thailand D. K. Reed, personal communication
Korea 0-63 Lee era/., 1980
China Zhang and Wang, 1982
 Biology and control of Asian corn borer 47

Table 4. (continued)

Parasite, parasitoid, predator species Stage Country Percentage Reference

attacked attacked

Trichogramma nr. plasseyensis Egg New Guinea 98 G. Young.1979

Trichogramma ostriniae Peng and Chang Egg China 0-θΟ Ν AS, 1977
Taiwan Chiu and Chen, 1986
Trichogramma tielingensis Zhang and Wang Egg China Zhang and Wang, 1982
Trichogramma nr. papilionis Nagarkatti Egg Malaysia 98 M. Y. Hussein, personal communication
Trichogramma sp. Egg Japan 31-73 Saito and Oku, 1985
Trichogrammatoidea armigera Nagaraja Egg Philippines Alba, 1988
Trichogrammatoidea sp. Egg Taiwan Chiu and Chen, 1986

Hymenoptera: Braconidae
Apanteles sp. Larval Korea Cartwright, 1933
Apanteles thompsoni Lyle Larval Manchuria, Low Clarke, 1934
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Japan
Bracon atricornis (Smith) Larval Japan Low Clarke, 1934, Baker er al., 1949
Chelonus communis Baker Larval Philippines Buligan, 1929
• Macrocentrus gifuensis Ashmead Larval Japan, Korea 1-32 Cartwright, 1933; Saito and Oku, 1985
Macrocentrus grandii Goidanich Larval Korea 1-2 Lee etal., 1980
Microgaster tibialis Nees Larval Manchuria, 1-5 Clarke, 1934; Cartwright, 1933
Japan

Hymenoptera: Ichneumonidae
Campoplex alkae (Ellinger and Schachtleben) Larval North China, 7-12 Clarke, 1934; Cartwright, 1933; Baker er
Korea al., 1949
Echthromorpha sp. Guam 0-2 Unpublished data
Eriborus sinicus (Holmgren) Larval Japan Kusigemati, 1981
Eriborus terebrans (Gravenhorst) Larval Hokkaido rare Kuwayama, 1930
North China, 10-20 Clarke, 1934; Baker etal., 1949
Korea,
Hokkaido
Exeristes roborator (F.) ? Pupal Hokkaido Kuwayama, 1930
Itamoplex sp. Hokkaido rare Kuwayama, 1930
Phaeogenes nigridens Wesm. Pupal Japan, Korea Cartwright, 1933
Phaeogenes sp. Pupal Japan 11 Clarke, 1934
Trathala flavo-orbitalis (Cameron) Larval Japan <1-30 Cartwright, 1933; Clarke, 1934
Larval, Philippines 2-1 Camarao, 1976a
Trichomma sp. pupal
Larval, Taiwan 2 Clarke, 1934; Baker etal.. 1949
Trichomma cnaphalocrocis Uchida pupal
Larval, Taiwan Cartwright, 1933
Xanthopimpla punctata (F.) pupal Guam, Tinian <1 Nafus and Schreiner, 1986b
Larval, Taiwan 2-80 Clarke, 1934; Cartwright, 1933
Xanthopimpla stemmator (Thunberg) pupal
Philippines 1-2 Camarao, 1976a
Malaysia 1 Yunus and Hua, 1969; Hussein, 1986

Hymenoptera: Chalcididae
Brachymeria albotibialis (Ashmead) Pupal Guam <1 Nafus and Schreiner, 1986b
Brachymeria euploeae (Westwood) Pupal Japan Clarke, 1934
Brachymeria lasus Walker Pupal PNG 4-5 G. Young,1979
Brachymeria lasus Walker Pupal Malaysia 15 Hussein, 1986
Brachymeria obscurata (Walker) Philippines <1 Camarao, 1976a
Malaysia Yunus and Hua, 1969
Brachymeria sp. Pupal Taiwan Cartwright. 1933
Chalcis euploeae Hope Pupal Philippines Buligan, 1929

Hymenoptera: Eulophidae
Tetrastichus Tmferens Yoshimoto Pupal Guam <1 Nafus and Schreiner, 1986b

Diptera: Tachinidae
Exorista tritaeniata Rondani Hokkaido 10 Kuwayama, 1930
Exorista sp. Hokkaido 2-4 Kuwayama, 1930
Lydella thompsoni Herting Larval China, Korea 0-1-17 Cartwright, 1933; Baker etal., 1949
Japan
Nemorilla floralis (Fallen) Larval Japan, Korea Baker etal., 1949
Paradrino laeviculus (Mesnil) Larval Philippines Shima, 1984
Pseudoperichaeta erecta (Coq.) Larval Japan Baker etal., 1949
Tachinid sp. Larval Korea 0-15 Lee etal., 1980
Tachinid sp. Pupal Papua 1-2 G. Young,1979
New Guinea
 48 D. M. Nafus and I. H. Schreiner

Table 4. (continued)

Parasite, parasitoid, predator species Stage Country Percentage Reference

attacked attacked

Tachlnid sp. Larval Taiwan 40-60 Tseng,1981

Parasites
Nematoda (Mermithidae)
Hexameris meridionalis Steiner Japan, 0-30 Cartwright, 1933
Taiwan

Predators
Hemiptera: Harpactoridae
Phemius tibialis Westwood Larval Philippines Buligan, 1929
Sphodronyttus erythropterus Burmeister Larval, Philippines Buligan, 1929
pupal
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Hemiptera: Anthocoridae
Orius sp. Egg Philippines rare Hasse, 1981
Onus niobe Herring Egg Guam Unpublished
Hemiptera: Pentatomidae
Eocanthecona furcellata (Wolff) Larval Thailand D. K. Reed, personal communication
Menida formosa Westwood Egg Philippines rarely Hasse, 1981
Dermaptera: Chelisochidae
Chelisoches morio F. New Guinea common G. Young, 1979
Proreus slmulans Stål. All stages Philippines sometimes Buligan, 1929, Camarao, 1976a
Thailand common Patanakamjorn et al., 1978c
Dermaptera: Carcinophoridae
Euborelia stali (Dohrn) Larval, Philippines Hasse, 1981
pupal
Dermaptera: Labiduridae
Labidura riparia (Pallas) All stages Philippines Situmorang and Gabriel, 1988
Coleoptera: Histeridae
Carphophyllus foveicollis Murr. Larval, Philippines Buligan, 1929
pupal
Coleoptera: Coccinellidae
Menochilus sexmaculatus (F.) Egg Philippines Camarao, 1976a
Micraspis crocea Mulsant Egg Philippines common Hasse, 1981
Coleoptera: Carabidae
Plochnius sp. Larval Japan Clarke, 1934
Chaenius sp. Hokkaido Kuwayama, 1930
Hymenoptera: Formicidae
Solenopsis geminata (F.) All stages Philippines Hasse, 1981
Guam Schreiner and Nafus, 1988
Monomorium minutum Mayr Egg Guam
Hymenoptera: Sphecldae Schreiner and Nafus, 1988
Sceliphron madraspanatum cospicillatum Philippines
Areneida: Argiopidae Camarao, 1976a
Araneus inustus (L. Koch) Larval Philippines
Areneida: Oxyopidae Hasse, 1981
Oxyopes javanus Thorell Larval Philippines
Areneida: Theriidae Hasse, 1981
Therlld sp. Larval Philippines
Areneida Hasse, 1981
Anystls sp. Egg Japan
Saito and Oku, 1985
Diseases
Beauveria bassiniana (Bals.) Vuillemans Larval China, Japan Clarke, 1934
Korea
Nosema furnacalis Wenn Larval China 0-92% Wenn, 1986
Nosema sp. Guam 0-10% Unpublished data
Metarrhizum anisopliae (Metchnikoff) Larval Taiwan Chiu and Chen, 1985
 Biology and control of Asian corn borer
Young, 1979), 7% or less of the larvae and pupae were
reported to be parasitized. In the Mariana Islands, less than
0-1 % of the larvae and pupae were parasitized (Nafus and
Schreiner, 1986b). The highest parasitization rates reported
were in Taiwan, where an unidentified tachinid seasonally
parasitized as much as 40-60% of the larvae (Tseng, 1981).
Normally it was present at much lower levels.
Egg parasitoids are present throughout the region, and
may play an important role in reducing numbers of O.
furnacalis. In China the predominant parasitoid in corn fields
is Trichogramma ostriniae Pang and Chen (this may be a
junior synonym for T. chilotreae Nagaraja and Nagarkatti).
In Beijing province, infestation rates are initially low, but rise
to about 20-30% in July, and 80% towards the end of the
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growing season (USDA, 1982). In Korea a Trichogramma
species (reported as evanescens) was absent in the
first-generation eggs, but parasitized 62-8% of the eggs in
the second generation (Lee et ai, 1980). In Taiwan three
species of egg parasitoids were found, 7". ostriniae, T.
chilonis Ishii and Trichogrammatoidea sp. (Chiu and Chen,
1986). T. ostriniae was the predominant parasitoid, and
appears to be better adapted as it is able to more fully utilize
masses when it discovers them. In laboratory tests, T.
ostriniae parasitized a greater proportion of the eggs in a
mass (over 85%), whereas T. chilonis parasitized less than
5%. T. ostriniae was also more tolerant of the pesticides
used in corn fields. In the Mariana Islands, 13-20% of the
eggs were parasitized by T. chilonis (Nafus and Schreiner,
1986b). This percentage increased late in the growing
season, after the number of ACB eggs laid on the plants fell
due to unsuitability of the host plant. In Papua New Guinea,
7". sp. nr. plasseyensis parasitized up to 98% of the egg
masses (G. Young, 1979). In Thailand, T. australicum Girault,
was found to parasitize about 8% of the egg masses when
moth oviposition began, but this increased to 80% the
following week and remained at that level or higher
throughout the growing season (Areekul era/., 1965). Recent
information from Thailand identifies T. chilotreae as the most
important parasitoid of O. furnacalis, and it is probable that
the name Γ. australicum was incorrectly assigned to the
parasitoid. No data were available for either PNG or Thailand
as to what percentage of the eggs within a mass were
parasitized.
A summary suggests that egg parasitoids are relatively
more effective in tropical than in temperate areas. No larval
and pupal parasitoids appear to have a significant effect on
ACB populations. In temperate parts of the ACBs range,
larval parasitoids are more abundant, having their greatest
impact during the diapausing generation.
14.2. Predators
Predators have been studied in the Philippines. Camarao
(1976a) observed Proreus simulans Stål (Chelisochidae),
Menochilus sexmaculatus F. (Coccinellidae) and several ant
species preying on the eggs. She pinned eggs onto plants
and found that over 60% of the eggs were lost, presumably
to predators. Islam (1981) reported that M. sexmaculatus
populations in corn fields were responding primarily to
aphid populations, and only ate ACB eggs when aphid
49
populations were low. In Thailand, undetermined predators
were found to remove 33-50% of the egg masses before
hatching (Mekongsee et al., 1979). In northern Japan only
0-3% of eggs were lost to predators and another 1-3%
became dislodged or failed to hatch (Saito and Oku, 1985).
On Guam only 4% of marked egg masses were lost (Nafus
and Schreiner, 1986a). The predators observed feeding on
the egg masses were the ant Monomorium minutum Mayr,
the fire ant Solenopsis gemlnata (F.) and a species of
anthocorid bug. Hasse (1981) found 5. geminata and several
species of spiders preying on small, exposed ACB larvae.
Only earwigs were observed to follow the ACB into their
tunnels. The ladybeetle Micraspis crocea Mulsant was the
most common egg predator.
Several species of earwigs are reported as predators. In
the Philippines, Islam (1981) reported the earwigs P.
simulans, Euboriella stali, and Labidura riparia (Pallas). P.
simulans and Labidura riparia were voracious feeders on
ACB larvae and pupae, but they were rare in the fields
studied. Euboriella stall (Dohrn) was the more common,
increasing in numbers when ACB populations were high.
Though it fed on ACB larvae, and particularly pupae, it was
also phytophagous. He was unable to correlate earwig
numbers with the number of consumed ACB larvae and
pupae in the tunnels. Situmorang and Gabriel (1988) found
that L. riparia would also consume eggs as well as all instars
of larvae. In the laboratory, individual L riparia killed many
more larvae than they ate. In the field L. riparia colonized
corn fields after the plant reached the whorl stage, if the soil
was moist. The earwigs would climb to the top of plants and
attack both eggs and larvae.
P. simulans has been reported to be an efficient predator
of ACBs. Camarao (1976a) found that, in a cage, P. simulans
and ants would eliminate an ACB population within two
generations. In the laboratory in Thailand, P. simulans
consumed large numbers of ACB. In the field, however,
increases in P. simulans populations were correlated with
aphid rather than ACB numbers. Still, Patanakamjorn er al.
(1978c) thought prédation by P. simulans was efficient
enough to hamper the establishment of sufficiently high ACB
populations for host resistance work.
In Papua New Guinea, Chelisoches morio (F.) has been
observed feeding on ACB (G. Young, 1979). This species has
also been observed in ACB tunnels in Guam, and
occasionally damaged pupae were observed.
14.3 Diseases
Native diseases of the ACB have been investigated to a
limited extent. Camarao (1983), in the Philippines, found
about 1 % of the larvae and 4% of the pupae died from
bacterial diseases. Fungi killed less than 1%. She also noted
a linear increase in the number of larvae in corn with
the incidence of bacterial diseases (Camarao, 1976a). Saito
and Oku (1985) did not find any disease in larvae during the
summer months in northern Japan, but 12-13% of
overwintering larvae were killed by disease.
Several diseases have been identified in China. A strain
of Beauveria bassiana was isolated from ACBs in Jilin
Province (USDA, 1982). The fungus is cultured by farmers
 50 D. M. Nafus and t. H. Schreiner
and used for control. It is formulated with sand or cinders
and applied to piled, infested corn stalks, where it kills
overwintering larvae. By treating stalks and stubble piled
near villages, damage to crops in the following year can be
reduced by about 70% (Cooperative group of ACB control
by Beauveria bassiana of Kirin Province, 1977). The fungus
can also be applied to field populations of moths. Depending
on the method of application, 60-80% kills are reported
(NAS, 1977).
Wenn (1986) found a species of microsporidian, Nosema
furnacalis Wenn, Infecting the ACB in China. Wenn et al.
(1986) surveyed 12 provinces and found the incidence of
infection by N. furnacalis varied from 0% to 92%. Heavy
infection rates occurred in areas with one ACB generation
per year, and lower rates in areas with two and three
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generations. In Guam, a survey showed larvae to be infected
with a microsporidian which appeared to be Nosema, but
the incidence of infection in any field never exceeded 10%
(unpublished data). Wen (1984) found that Infection could be
increased by artificial infestation, the most efficient
transmission occurring at the silking stage of corn. Mortality
of larvae increased if moderately resistant corn varieties
were used (Wen, 1985).
15. Classical biological control
Limited attempts have been made to use classical
biological control against O. furnacalis. The most extensive
efforts were in the Mariana Islands. In the 1920s and 1930s,
Exerlstes roborator (F.) (Ichneumonidae) was introduced to
Guam from the USA (Vandenberg, 1930), and Campoplex
alkae (Ellinger and Sachtleben) (Ichneumonidae), Lydella
thompsoni Herting (Tachinidae) and Trathala flavoorbitalis
(Cameron) (Ichneumonidae) were brought in from Japan and
Manchuria (Vandenberg, 1933). The latter two established:
T. flavoorbitalis only on leaf rollers attacking grasses
(Swezey, 1946) and L. thompsonlon the ACB. Swezey (1946)
reported that by 1936 Lydella was parasitizing a high
proportion of the ACB and that there was little damage to
corn. However, in 1951, Peterson (1955) collected several
thousand ACB larvae from which only one tachinid emerged.
Peterson (1955) imported and released L thompsoni,
Macrocentris grandii Goidanich (Braconidae), Eriborus
terebrans (Gravenhorst) (Ichneumonidae), Chelonus annu-
lipes Wesmael (Braconidae) and Agathis agilis (Cressman)
(Braconidae) from the USA, but none became established.
Trichomma cnaphalocrocis Uchida was introduced from
Taiwan in 1986; however, ants consumed all the parasitized
ACB pupae in the field preventing establishment (Nafus and
Schreiner, 1989). L. thompsoni, E. terebrans, and C.
annulipes were released on Saipan in 1956, but did not
establish (Gardner, 1958; Nafus and Schreiner, 1986b).
In the Philippines, Macrocentrus gifuensis Ashmead,
Horogenes punctorius Roman and L. thompsoni were
introduced from the USA In the 1950s. The former did not
establish. One male of each of the latter two species was
reared from the field several months to several years later,
but no further sign of establishment was observed (Baltazár,
1963). T. australicum (reported as a parasitoid of corn borers
in Thailand) was imported from Taiwan for control of
sugarcane borers. They were mass-reared for several years
and became established on the islands of Luzon, Negros
Occidental and Panay (Baltazár, 1981). However, according
to Alba (1988) the species actually introduced from Taiwan
was T. chilonis, which was already present In the
Philippines. In China, Trichogramma nubilale Ertle and
Davis (imported from the USA where it parasitizes O.
nubilalls) has been field-tested, and was found to parasitize
eggs of O. furnacalis and other insects (Li et al., 1984).
The microsporidian Nosema pyraustae was imported to
the Philippines from the USA and tested in the laboratory
against the ACB, but no infection developed (Baltazár, 1963).
This organism was also imported from the USA and applied
to rice and cane borer eggs and larvae in West Java,
Indonesia (Dressner, 1958).
16. Augmentative biological control
In China, augmentative biological control with a number
of species of Trichogramma is successful. Two widely used
species are Trichogramma dendrolimi Matsumura and T.
ostriniae. T. dendrolimi kills the ACB eggs but is unable to
complete development and cannot recycle (NAS, 1977;
USDA, 1982). 7". ostriniae does recycle and is widely used,
but is harder to mass-rear economically. T. dendrolimi is
reared on the eggs of the giant silkworm moth Antheraea
pernyi (Guérin-Ménéville), which is raised on oak in the
forest regions of northwest China and sold to biological
control stations. T. ostriniae will not accept this host and
must be reared on Corcyra cephalonica (Stainton). C.
cephalonica is fed rice bran. The efficiency of production is
much less for T. ostriniae than for T. dendrolimi (Cock,
1985).
T. evanescens Westwood, T. japonicum, T. chilonis and T.
closterae Pang & Chen have also been used (Cock, 1985).
Care is taken to maintain quality stocks, regardless of
species. Stocks are renewed every year and only active,
healthy wasps are reared (Institute of Agricultural Science
of He-j'ang District, Heilkungkiang Province, 1976; USDA,
1982; Cock, 1985).
The Chinese have developed a management system using
early releases and later spot follow-ups as needed.
Emphasis is on reducing the first generation as much as
possible. To do this the Chinese forecast the expected time
when the moths will begin ovipositing. Three releases of
Trichogramma are then made. The first is made several days
prior to expected beginning of oviposition by ACB, and the
next two releases at 4-day intervals. Where there are two or
more generations per year, additional releases are made,
timed to coincide with the beginning of oviposition of the
second and third generations if ACB populations are high
(USDA, 1982). Successful control of the later generations
may not be as good, possibly due to the effects of high
temperatures and rainfall on the parasitoids (Cock, 1985).
Releases are made by a variety of methods. Parasitized
eggs may be placed in the fields on cards pinned to the
plants, or by other methods if the stalks will be used as cattle
feed (NAS, 1977). Alternatively, plastic capsules may be
 Biology and control of Asian corn borer
used. Fifteen release stations are used per hectare.
Depending on the wasp species and other factors, from 12
to 38 wasps/m2 are released at each station (Cock, 1985).
Originally, a larger number of release stations, 75 per ha,
were used, but the number was reduced after the Chinese
discovered there was an optimum height for release. Using
these procedures, AC6 numbers are estimated to be
reduced 60-80% (Institute of Agricultural Science of He-jiang
District, Heilkungkiang Province, 1976; Feng et al., 1977;
NAS, 1977; USDA, 1982).
Augmentative releases of Trichogramma have also been
tried in Taiwan, where up to 95% of the ACB eggs were
parasitized (Tseng, 1981). Chiu and Chen (1986) found that
release of T. chilonis at rates of 40 000-80 000 did not affect
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yield or appreciably increase parasitization rates. T. chilonis
parasitized 19% of the eggs in the non-augmentation plots
and 27-36% of the eggs in the augmentation plots. Grain
yields were similar, although infestation rates were slightly
lower in the augmentation plots. They concluded that T.
chilonis was much less effective than T. ostriniae.
T. chilonis is also much more susceptible to insecticides
than T. ostriniae (Chiu and Chen, 1986). Furadan, commonly
used to control the ACB in Taiwan, is extremely toxic to both
species, as is Dursban. Carbaryl, EPN, and Lannate are also
moderately toxic. Maneb and Zineb were the least toxic.
Because of its inefficient parasitization and susceptibility to
insecticides, T. chilonis is probably of limited use for
augmentation purposes. Taiwan has recently completed a
facility to mass-rear Trichogramma for use in ACB control.
In the Philippines, Trichogramma evanescens has been
tested for use in augmentative releases (Hassan, 1984).
17. Cultural controls
17.1. Intercropping
Augmentation of Trichogramma without rearing has been
tried. In Guam an attempt was made to increase field
populations of T. chilonis by intercropping corn with sweet
potatoes. Sweet potatoes are a host for Agrius convolvuli
(L.) (Sphingidae), a highly suitable host for T. chilonis. The
moths colonize the potatoes earlier than the ACBs colonize
the corn, providing time for one to three generations of 7.
chilonis before ACB eggs are abundant. The percentage of
parasitized ACB eggs was higher in the intercrop, but not
significantly, and there was no reduction in damage to the
corn (Nafus and Schreiner, 1986a).
Intercropping corn with peanuts or soy beans reduced
populations of the ACB in the Philippines (IRRI, 1973, 1974;
Hasse, 1981). Hasse (1981) found that ACB larvae were more
successful at colonizing corn plants in monoculture rather
than as an intercrop. He hypothesized that the intercrop
acted as a barrier which slowed dispersal of the first-instar
larvae and exposed them to predators, of which there were
more in the intercrop.
77.2. Planting dates
Manipulating planting dates to break links between crop
and pest cycles is commonly used to reduce pest problems.
51
The range of the ACB extends from temperate, seasonal
environments to both non-seasonal and seasonal tropical
areas. Where the ACB diapauses, or where generations are
synchronized by wet or dry seasons, planting dates can be
effectively manipulated. In central Shensi province, in China,
corn is planted in the early summer after winter wheat. ACB
populations are minimal because the corn is planted too late
for the overwintering ACB. In contrast, in northern Shensi,
where corn is planted in the spring, ACB can be a problem
(NAS, 1977).
In the tropical parts of its range, generations are
continuous and overlapping, but population levels vary
seasonally if there is a distinct wet and dry season. In the
Philippines, ACB populations are higher during the wet than
the dry season (Camarao, 1979). In the wet season, Viado
et al. (1958) reported that infestations were higher in the
early part than in the middle, when rainfall was heaviest. In
Guam the likelihood of severe infestations is highest at the
end of the rainy season, and lowest if the corn is planted at
the end of the dry season (unpublished data). This probably
relates to the biology of the ACB in its alternative hosts,
many of which die back during the dry season. For the
Philippines, Bato etal. (1983a) recommend that corn should
be planted as early as possible in the cropping season.
77.3. Other cultural controls
Overfertilization of corn can result in higher ACB
populations early in its growth. Saito (1985) found that corn
fertilized at double the normal rate in Japan had more ACB,
and the larvae had slightly faster growth rates. The higher
number of ACB in the heavily fertilized corn was probably
a function of its size. The fertilized corn grew faster and was
taller than less fertilized corn. Cheu (1941) has shown that
the ACB prefers taller corn. Later in the season, when the
second generation emerged, the less-fertilized corn was in
a growth stage which was more attractive for oviposition
and it received more eggs. Survival was about the same
regardless of fertilizer level, probably due to the increased
suitability of the stage of the unfertilized corn. By
the end of the season, because of the shifts in the
attractiveness of the plants, fertilizer levels had no effect on
the final densities of ACB.
18. Mechanical controls
In many areas of China, light traps are used to catch ACB.
In Honan province, 8000 permanent blacklight traps were
built on 20 000 acres. The traps caught about 200 ACBs per
night, corresponding to a total catch of 33 000 kg in 1974.
The number of infested plants was claimed to be reduced
by 50%, leading to a reduction of 67-90% in insecticide cost
and 77-80% in labour costs (NAS, 1977). However, in the
Philippines, ACB adults were not found to be attracted to
light traps (Camarao, 1979; Hasse, 1981).
In both the Philippines and Guam a folk method of
controlling ACB is to remove the tassels before or at pollen
shed. Felkl (1988a) found that two-thirds of all first-,
second- and third-instar larvae and one-fourth of the
 52 D. M. Nafus and I. H. Schreiner
fourth-instar larvae are in the tassel at the early tasselling
stage. In the Philippines, Bato et al. (1983b) removed the
tassels from four out of every six rows of field corn and found
that it reduced the numbers of cavities in the stalk and
increased yield in some instances. Morallo-Rejesus and
Javier (1985) report that the effect of 75% detasselling on
yield was comparable to two or three insecticide sprays
applied before the tasselling period. Numbers of borers at
harvest and yield were comparable whether the tassels left
in the field for pollination were sprayed or not. Felkl (1988b)
reported that removing the remaining tassels after pollen
shed did not increase yield. In her experiments the
application of carbofuran after detasselling did increase
yields compared to detasselling alone. However, under
Philippine conditions, detasselling alone provided a higher
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marginal return than did detasselling plus the use of
insecticide.
In Guam, detasselling reduced the number of ACBs
significantly on sweet corn, but marketable yield was
obtained only if effective insecticides were also used at
weekly intervals. Insecticides alone were not as effective as
the combination of 75% detasselling and insecticide use. If
ACB numbers were very high, no combination of treatments
resulted in any marketable yield (Schreiner and Nafus, 1987,
1988).
19. Chemical controls
Numerous insecticides have been tested against ACB on
corn. In com still in the whorl stage the larvae can be
effectively controlled by the application of granular
insecticides into the whorl (Kongkajana et al., 1972;
O'Sullivan and Bourke, 1975; Young, W., 1979). However,
Nickle (1966) found that control at the whorl stage did not
significantly reduce subsequent stalk tunnelling or yield
loss. In the Philippines, Felkl (1988b) reported the application
of carbofuran at the whorl stage with no further treatment
resulted in a negative net-marginal return.
In older corn, insecticides are less reliable (Litsinger ef
al., 1980). Methomyl and decamethrin were among the most
effective non-chlorinated-hydrocarbon sprays tested, and
Bacillus thuringiensis also provided relatively good control
(Litsinger and Dela Cruz, 1980, 1981; Nafus and Schreiner,
1985). Third-instar larvae were the most resistant to
insecticides in toxicity tests (Rodriguez and Sanchez, 1975).
Chitin inhibitors may prove to be effective as ovicidal agents
(Chen et al., 1982). On cotton no effective insecticides have
been reported (De I attre and Pauly, 1982).
In the rainy season it is often not possible to control the
ACB with insecticides. Areekul (1965) found that endrin,
carbaryl and endrin + malathion provided some reduction
of the ACB if rainfall was around 1-5 mm, but not at 3·5-4·9
mm.
Insecticide use varies. In the Philippines, small-scale
farmers do not use insecticides to control the ACB (Litsinger
et al., 1980). In China, several insecticides are widely used
in areas where there is more than one generation of ACB
per year. These include BHC, Dipterex (trichlorfon), granular
carbaryl and Phoxim (NAS, 1977). Insecticidal materials are
sometimes used together with B. bassiana. A synergistic
effect of about 10% between B. bassiana and carbaryl was
found (NAS, 1977).
20. Breeding for resistance
Many Asian countries have programmes to develop
varieties of corn resistant to the ACB. Resistance to ACB in
vegetative stage corn has been demonstrated in several
breeding lines (Guthrie era/., 1982; Charoenying era/., 1979;
Mekongsee et al., 1979; Adaila, 1987) and several varieties
have been released (Kim ef al., 1978a, b). In comparative
trials between selected inbreds in Taiwan and Iowa, in
pre-reproductive corn, inbreds with resistance to the
European corn borer scored similar ratings against the ACB
(Guthrie et al., 1982). Some resistance was present in 14 of
104 breeding lines tested. In the Philippines, corn varieties
with high DIMBOA contents were demonstrated to be more
resistant to the ACB (Santiago and Mendoza, 1983).
In reproductive stage corn, little resistance has been
reported. Two inbreds which showed resistance to the
European corn borer did not show resistance to the ACB
(Guthrie ef al., 1982). Feeding habits between the two
species differ to some extent at this stage (Nafus and
Schreiner, 1987), so this is not surprising, Antigua Group 1
has been identified as a good source of general resistance
to the corn borer in the Philippines, and current breeding
work there has concentrated on a number of elite lines of
IPB VAR 1 x Antigua Group 1 (Adalla, 1987). These lines
have been found to be tolerant of corn borer damage to the
leaves, stalks and tassels.
21. Conclusion
Since the revision of the genus Ostrinla, research on the
ACB has increased markedly. Progress is being made in a
number of areas including general biology, host-plant
resistance, and biological control. Some disturbing trends,
including switches to new hosts such as cotton, are
apparent. Host-plant resistance has been found, and
prospects are good for developing better resistance in
vegetative stage corn. Detasselling is an aid to controlling
the ACB in tropical areas, and is inexpensive for small-scale
farmers. It is also compatible with other strategies in a
management programme. Inundative releases of Tricho-
gramma and of diseases have been demonstrated to work
in China.
There are many gaps. Better understanding of the natural
enemy complexes where the ACB is not a severe pest is
needed. Interactions between alternative, wild hosts,
immigration into farm fields, and the population dynamics of
the ACB need to be better understood. The relationship
between yield losses, ACB infestation levels, and timing of
ACB infestation needs more research. In tropical areas,
more information on the contribution of early infestations to
the development of reproductive stage infestations is
needed.
 Biology and control of Asian corn borer
Acknowledgements
We thank K. Hsiao, C. T. Lee and M. Marutani who
assisted us in translating documents in Chinese and
Japanese. The retrieval of documents was funded by USDA
Special Grants in Tropical and Sub-Tropical Agriculture
83-CSRS-2-2167. This is publication 83 of the Guam
Agricultural Experiment Station.
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