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Rice Pest PDF
Rice Pest PDF
Pests
O R F ICE
M. D. Pathak and Z. R. Khan
1994
ICIPE
International Centre of Insect Physiology and Ecology
The International Rice Research Institute (IRRI) was established in 1960 by
the Ford and Rockefeller Foundations with the help and approval of the
Government of the Philippines. Today IRRI is one of 18 nonprofit interna-
tional research centers supported by the Consultative Group on Interna-
tional Agricultural Research (CGIAR). The CGIAR is sponsored by the
Food and Agriculture Organization of the United Nations (FAO), the
International Bank for Reconstruction and Development (World Bank), and
the United Nations Development Programme (UNDP). Its membership
comprises donor countries, international and regional organizations, and
private foundations.
IRRI receives support, through the CGIAR, from a number of donors
including FAO, UNDP, World Bank, European Economic Community,
Asian Development Bank, Rockefeller Foundation, Ford Foundation, and
the international aid agencies of the following governments: Australia,
Belgium, Canada, People's Republic of China, Denmark, Finland, France,
Germany, India, Islamic Republic of Iran, Italy, Japan, Republic of Korea,
The Netherlands, Norway, Philippines, Spain, Sweden, Switzerland,
United Kingdom, and United States.
The responsibility for this publication rests with the International Rice
Research Institute.
ISBN 971-22-0028-0
Contents
The world rice crop is attacked by Reduction in insect pest damage This new edition includes updated
more than 100 species of insects; 20 of should come from incorporating information on biology, damage,
them can cause economic damage. genetic resistance into new genotypes seasonal history and factors of
Insect pests that can cause significant and from the development of suitable abundance, and control measures of
yield losses are stem borers; leafhop- cultural and biological control meth- the major insect pests of rice. IRRI
pers and planthoppers (which cause ods. The first edition of this book, hopes this expanded content will
direct damage by feeding as well as published in 1967, contained basic prove useful to researchers, extension
by transmitting viruses); gall midges, information on the biology and workers, and students everywhere.
a group of defoliating species (main1y factors of abundance of common Many people were involved in the
lepidopterans); and a grain-sucking insect pests of rice. Since then, due to production of this book. N.J. Fernan-
bug complex that feeds on develop- the introduction of high-yielding dez, A.D. Tan, and F.F.D. Villanueva
ing grains. modern varieties, distinct changes helped compile the text, references,
Average yield loss due to various have occurred in the insect pest and tables; A.T. Barrion validates
insect pests in Asia-wheremore complex of rice. Several species, once scientific names of insect pests;
than 90% of the world's rice is pro- considered minor pests, have become E. Panisales provided artwork; and
duced—is about 20%. Any decrease major pests. Also, much information M.L.P. Abenes provided photogra-
in pest damage means a correspond- on various aspects of control, includ- phic services. The volume was edited
ing increase in needed rice produc- ing integrated pest management, has by W.H. Smith and G.S. Argosino.
tion. become available.
Klaus Lampe
Introduction
Rice, the staple diet of over half of the Two major factors are responsible
world's population, is grown on over for low yields: adverse weather
145 million ha in more than (floods, drought, typhoons, etc.) and
110 countries, and occupies almost pest epidemics.
one-fifth of the total world cropland Low temperature is a major factor
under cereals. Classified primarily as limiting rice cultivation. The opti-
a tropical and subtropical crop, rice is mum temperature is about 30°C, and
cultivated as far north as 53° N temperatures lower than 20°C,
latitude on the border between the particularly during the flowering
USSR and China and as far south as stage, induce sterility. In regions of
39° S latitude in Central Argentina, cool winters only one crop a year can
and from sea level to altitudes of be grown, but in warm areas as many
3,000 m. The crop is established either as three crops are common.
by direct sowing (broadcast or Average rice yield varies from less
drilled) or by transplanting. Rice than 1 t/ha in some tropical countries
1. Magnitude of rice crop loss due to
grows under diverse water regimes: to more than 6 t/ha in Japan, Repub- insect pests in the Philippines. The
it is an upland crop where there is no lic of Korea, and the USA. Rice yields average yield from plots protected with
standing water and rains are the sole in South and Southeast Asia, the insecticides was 4.9 t/ha whereas that
source of moisture, or a lowland crop world's rice bowl, fluctuate widely, from unprotected plots was 3.0 t/ha,
suggesting a loss of 40% (modified from
under conditions in which water, averaging around 2 t/ha.
M.D. Pathak and G.S. Dhaliwal, 1981,
derived either from rain or irrigation Most of the world's rice produc- Trends and strategies for rice insect
systems, is impounded in the fields. tion is from irrigated and rainfed problems in tropical Asia, IRRI Res. Pap.
Rice is cultivated on terraces, on lowland ricefields where insect pests Ser. 64, International Rice Research
slopes, and in valleys or other low- are constraints. The warm and humid Institute, P.O. Box 933, Manila 1099,
Philippines).
lying sites. Floating rice may be environment in which rice is grown is
grown in several meters of standing conducive to the proliferation of
water. insects. Heavily fertilized, high-
As many as 80,000 rice accessions tillering MVs and the practice of
(cultivated and wild varieties) have multicropping rice throughout the
been collected at the International year favor the buildup of pest popu-
Rice Germplasm Center of the Inter- lations. The intensity of the insect
national Rice Research Institute problem in such an area can be
(IRRI). The traditional tropical rice illustrated by the experience at IRRI.
varieties are tall and leafy; they often In 117 experiments conducted over
lodge during the later stages of 15 yr, plots protected from insects
growth. Modern varieties (MVs) are yielded almost twice as much as
short—usually about 1 m high—stiff- unprotected plots (Fig. 1). Average
strawed, erect-leafed, and lodging rice yield loss due to various insect
resistant. The plant characters of the pests was estimated to be 31.5% in
MVs are commonly associated with Asia (excluding mainland China) and
high yields. 21 % in North and Central America in
1
1967. 1 Estimates for tropical South Table 1. Insect pests and stages at which they attack the rice crop.
and Southeast Asia are considerably
Insect pests (order:family)
lower. In a 1989 survey of 50 rice en-
tomologists from 11 countries, Vegetative sfage
average yield losses due to insect Seedling maggots (Diptera: Muscidae)
pests were estimated at 18.5%. Yield Rice whorl maggots (Diptera: Ephydridae)
Rice caseworms (Lepidoptera: Pyralidae)
increases of this magnitude fre-
Rice green semiloopers (Lepidoptera: Noctuidae)
quently result from effective insect
Rice leaf beetles (Coleoptera: Chrysomelidae)
control in the different South and Rice thrips (Thysanoptera:Thripidae)
Southeast Asian countries. Rice gall midge (Diptera: Cecidomyiidae)
The rice plant is subject to attack Armyworms and cutworms (Lepidopera: Noctuidae)
by more than 100 species of insects; Grasshoppers, katydids, and field crickets (Orthoptera: Acrididae, Gryllidae, and Tettigoniidae)
20 of them can cause economic Rice leaffolders (Lepidoptera: Pyralidae)
damage. Together they infest all parts Rice stem borers (Lepidoptera: Pyralidae and Noctuidae)
Stalked-eyed flies (Diptera: Diopsidae)
of the plant at all growth stages, and
Black bugs (Hemiptera: Pentatomidae)
a few transmit viral diseases. The Rice hispa (Coleoptera: Chrysomelidae)
major insect pests that cause signifi- Mealybugs (Homoptera: Pseudococcidae)
cant yield losses are leafhoppers and
planthoppers, which cause direct Reproductive stage
damage as well as transmit viruses; Greenhorned caterpillars (Lepidoptera: Satyridae)
stem borers; and a group of defoliator Rice skippers (Lepidoptera: Hesperiidae)
Planthoppers (Homoptera: Delphacidae)
species (Table 1). As in many other
Leafhoppers (Homoptera: Cicadellidae)
agroecosystems, the rice agroecosys-
tem has a few primary pests that may Ripening stage
actually limit production under Ripening seed bugs (Hemiptera: Alydidae)
certain conditions. In addition to the Stink bugs (Hemiptera: Pentatomidae)
primary pests are numerous species
that cause periodic losses, and a few Soil-inhabiting pests
species that may occur in such low Ants (Hymenoptera: Formicidae)
Termites (Isoptera: Termitidae and Rhinotermitidae)
numbers that no damage occurs.
White grubs (Coleoptera: Scarabaeidae)
Since the introduction of high- Field crickets (Orthoptera: Gryllotalpidae)
yielding varieties, distinct changes Mole crickets (Orthoptera: Gryllotalpidae)
have occurred in the insect pest Root weevils (Coleoptera: Curculionidae)
complex of rice in Asia. Several Root aphids (Homoptera: Aphididae)
species, which once were considered Wire worms (Coleoptera: Elateridae)
minor pests, are now considered Root-feeding mealybugs (Homoptera: Pseudococcidae)
major (Table 2). Examples are the
brown planthopper, whitebacked
planthopper, green leafhopper, and
leaffolders. Until the 1960s, the stem
borers were considered the most
serious pests of rice throughout the
tropics. In recent years, however,
damage from them has declined. In
Japan, the population densities of
stem borers have steadily declined
since the mid-1960s (Fig. 2).
Introduction 3
Stem borers
The stem borers, generally consid- Table 3. Stem borers of rice worldwide.
ered the most serious pests of rice
Order Family Species a Distribution
worldwide, occur and infest plants
from seedling stage to maturity. Fifty Lepidoptera Pyralidae Acigona ignefusalis (Hampson)
Adelpherupa flavescens Hampson Africa
species in three families-Pyralidae, Ancylolomia chrysographella (Kollar) Asia
Noctuidae (Lepidoptera), and Diop- Catagela adjurella Walker China
sidae (Diptera)—are known to attack Chilo agamemnon Bleszynski Middle East/North-East Africa
Chilo aleniellus (Strand) Africa
the rice crop (Table 3). Thirty-five
Chilo auricilius Dudgeon Asia
pyralids belonging to 12 genera, Chilo diffusilineus (J. de Joannis) Africa
10 noctuid species belonging to Chilo luniferalis Hampson Africa
Chilo mesoplagalis (Hampson) Africa
3 genera, and 5 diopsid species
Chilo partellus (Swinhoe) West Asia/Africa
belonging to the genus Diopsis have Chilo plejadellus Zincken North America
been recorded as rice stem borers. Chilo polychrysus (Meyrick) Asia
Chilo psammathis (Hampson) Africa
The pyralid borers are the most Asia
Chilo sacchariphagus indicus (Kapur)
common and destructive, and usually Chilo suppressalis (Walker) Europe/Middle East/Asia/
have high host specificity. The Oceania
Chilo zacconius Bleszynski Africa
noctuid borers are polyphagous and
Diatraea lineolata (Walker) Central/South America
only occasionally cause economic Diatraea saccharalis (Fabricius) North/South America
losses to the rice crop. In Asia, the Elasmopalpus lignosellus (Zeller) North/South Amerlca
most destructive and widely distrib- Eldana saccharina Walker Africa
Maliarpha separatella Ragonot Africa/West Asia
uted are yellow stem borer Niphadoses palleucus Common Australia
Scirpophaga incertulas (Walker), Rupela albinella (Cramer) North/South America
striped stem borer Chilo suppressalis Scirpophaga aurivena (Hampson) Asia
Scirpophaga fusciflua Hampson Asia
(Walker), white stem borer Scirpophaga gilviberbis Zeller Asia
Scirpophaga innotata (Walker), dark- Scirpophaga incertulas (Walker) Asia/Australia
headed stem borer, Chilo polychrysus = Schoenobius incertulas (Walker)
= Tryporyza incertulas (Walker)
(Meyrick), and pink borer Sesamia Scirpophaga innotata (Walker) East Asia/Australia
inferens (Walker). In Asia, Scirpophaga = Tryporyza innotata (Walker)
incertulas and Chilo suppressalis are Scirpophaga lineata (Butler) Asia
Scirpophaga nivella (Fabricus) Asia/Australia/Oceania
responsible for a steady annual Scirpophaga occidentella (Walker) Africa
damage of 5-10% of the rice crop, Scirpophaga subumbrosa Meyrick Africa
with occasional localized outbreaks Scirpophaga virginia Schultze Asia
Noctuidae Bathytricha truncata (Walker) Australia
of up to 60%. Busseola fusca Fuller Africa
Scirpophaga incertulas, distributed Sesamia botanephaga Tams &
primarily in the tropics, also occurs in Bowden Afrlca
Sesamia calamistis (Hampson) Africa
the temperate areas where tempera- Sesamia cretica Lederer Africa/Europe/Middle East
ture remains above 10 ºC and annual Sesamia epunctifera Hampson Africa
rainfall is more than 1,000 mm. It is Sesamia inferens (Walker) Asia/Australia/Oceania
Sesamia nonagrioides (Lefebre) Africa
the predominant species in Bangla- Sesamia penniseti Tams & Bowden Africa
desh, India, Malaysia, Pakistan, the Sesamia uniformis Dudgeon Asia
Philippines, Sri Lanka, Thailand, Diptera Diopsidae Diopsis apicalis Dalman Africa
Diopsis circularis Macquart Africa
Vietnam, and parts of Indonesia. Diopsis ichneumonea Linnaeus Africa
Chilo suppressalis and Scirpophaga Diopsis macrophthalma Dalman Africa
innotata follow close behind. In Diopsis servillei Macquart Africa
Bangladesh, Scirpophaga incertulas is a Species printed in boldface are those commonly occurring on rice.
5
of major importance followed by In North and South America, Most borer species can fly
Sesamia inferens. In the Republic of Diatraea saccharalis (Fabricius) is the 5-10 miles, but can cover longer
Korea, Chilo suppressalis is the only most widespread species, followed distances if carried by winds. The
stem borer damaging rice; Scirpophaga by Rupela albinella (Cramer) and distance covered per second has been
incertulas does not occur. In Japan, Elasmopalpus lignosellus (Zeller). reported as 0.6-3.4 m for Chilo
Chilo suppressalis and Scirpophaga Because stem borers are the most suppressalis males, which fly in an
incertulas are the two economically important rice pests in Asia and other irregular or circuitous course, and
important rice borers. Because parts of the world, their bionomics 0.48-2.15 m for females, which usu-
Scirpophaga incertulas is restricted to has been widely studied. Except for ally fly in straight lines.
southern Japan, the maximum area of some investigations in Japan, most Mating in most species generally
ricefields it infests is one-tenth of that studies have been conducted under occurs between 1900 and 2100 h. The
of Chilo suppressalis. Moreover, natural environmental conditions and sex ratio of different species, based on
Scirpophaga incertulas in Japan has any ecological conclusions drawn are light trap catches, has been reported
been steadily decreasing since 1948 more generalized than specific. as generally more females than
and Chilo suppressalis since 1960. males, except for a 1:l ratio for
Scirpophaga incertulas is also a major Maliarpha separatella. In the absence of
pest of deepwater rice in eastern Life history data on phototropism of different
India, Bangladesh, and Thailand, sexes in these experiments, the
causing more than 20% yield loss in Adults validity of light trap catches to
many fields. The adults of lepidopterous stem represent sex ratios in nature is
Chilo suppressalis is highly tolerant borer species (Fig. 3a-3f) are noctur- questionable.
of low temperature. Full-grown nal, positively phototropic, and In experiments at IRRI, field-
larvae exposed to -14°C for 1-3 h do strong fliers; diopsid flies (Fig. 4a, b) collected females of Chilo suppressalis
not exhibit significant mortality. are diurnal and rest in the shade and Sesamia inferens mated many
Scirpophaga innotata, a tropical when not actively flying. Scirpophaga times; those of Scirpophaga incertulas
species, occurs in regions with incertulas moths usually emerge and Scirpophaga innotata mated only
distinct dry and wet seasons. Chilo between 1900 and 2100 h; Chilo once. In laboratory tests using vary-
polychrysus, initially reported as the suppressalis moths emerge from 1500 ing sex ratios of Chilo suppressalis,
most common and destructive in to 2300 h, peaking between 1900 and individual females mated as often as
Asia, has been recorded in several 2000 h, and become active again four times and males as often as eight
other countries in recent years and its toward dawn. During the day, Chilo times. The male moths were strongly
importance is being increasingly suppressalis hides among the grasses attracted to the virgin females.
recognized. while Scirpophaga incertulas and Attraction was maximum on the
In Africa, sorghum stem borer R. albinella remain in nurseries or evenings of female emergence, but
Chilo partellus (Swinhoe), Chilo ricefields. declined on subsequent days. Virgin
diffusilineus J. de Joannis, white stem The strong phototaxis of these females used as baits in field traps
borer Maliarpha separatella Ragonot, species in earlier years was used to attracted several wild males, but no
and African pink borer Sesamia attract them to light traps for moni-
calamistis (Hampson) are serious rice toring and control. In Japan, how-
pests. In eastern Africa, the principal ever, even with one light trap in-
stem borer of upland rice is Chilo stalled in every 80 ha of rice, only
partellus. Maliarpha separatella and 50% of the moth population could be
Sesamia calamistis are more abundant attracted. The moths are most at-
in lowland rice. In Central and West tracted to UV and green fluorescent
Africa, Maliarpha separatella and lights. Light traps are now used only
Sesamia calamistis are dominant stem for studying population fluctuations.
borers of upland rice. Chilo
diffusilineus and Chilo partellus are
important pests in upland savannas.
Stalk-eyed stem borers Diopsis spp.
are also important rice pests in
Africa.
Stem borers 7
Eggs Egg masses of lepidopterous stem decreases with temperature increase,
Lepidopterous stem borers lay eggs borers usually contain 50-80 eggs, beginning at 30 °C and continuing up
in masses (Fig. 5a-5e); diopsid flies and a single female is capable of to 35 °C. Although embryonic
lay isolated eggs. The eggs of laying 100-200 eggs. Diopsid females development can be completed at 35
Scirpophaga incertulas and Scirpophaga lay about 30 eggs each in a span of °C, the larvae die within the egg shell.
innotata are laid near the tip of the about 2 wk. Pyralids oviposit openly In Chilo suppressalis eggs,
leaf blade, while those of Chilo on the leaf blades, noctuids oviposit cholinesterase activity starts at about
suppressalis and Chilo polychrysus are behind leaf sheaths. The eggs of 60 h after oviposition. This could be
found at the basal half of the leaves Scirpophaga incertulas, Scirpophaga the reason for the ineffectiveness of
or, occasionally, on leaf sheaths. innotata, and R. albinella are covered organophosphate insecticides on
R. albinella and Chilo polychrysus with pale orange-brown hairs from freshly laid eggs. Egg development
oviposit on the lower surface of the the anal tufts of the female moths duration in diopsids is 2-3 d; that in
leaf blade. Several workers in Japan (Fig. 5a, b). Those of Chilo suppressalis lepidopterous moths, 5-9 d.
have reported that first-generation and Chilo polychrysus have no such The optimum egg hatching
Chilo suppressalis moths normally cover (Fig. 5c). Maliarpha separatella temperature is 21-33 °C for
oviposit on the upper surface of the eggs, although devoid of any such Chilo suppressalis and 24-29 °C for
leaves and that moths of subsequent covering, are more ingeniously Scirpophaga incertulas. Both species
generations deposit eggs on the lower protected in that the glue, which the require 90-100% RH; hatching is
surfaces. For several thousand field- females spread on the leaf before severely reduced below 70% RH. The
collected eggs at IRRI, no distinct oviposition, wrinkles the leaves, eggs usually hatch during daytime.
difference in position on either leaf forming a case that encloses the egg In Chilo suppressalis, maximum
surface was recorded, except that the mass. Among eggs of all species, hatching is from 0500 to 0600 h,
eggs on the upper leaf surface of those of Sesamia inferens, laid be- followed by another peak from 1400
hairy varieties were laid in the tween the leaf sheath and the stem, to 1600 h. In R. albinella, hatching
glabrous area along the midrib. are probably the most effectively usually occurs in the evening. Gener-
protected (Fig. 5d). ally, all eggs in a mass hatch simulta-
The threshold temperature for neously (Fig. 5e). Larvae emerged
development of Chilo suppressalis eggs from a large egg mass of Chilo
is reported to be 10-12 °C. Although suppressalis in about 13 min, but those
Scirpophaga incertulas eggs show some from a small egg mass lacked syn-
development at 13 °C, hatching chronization and took longer.
normally occurs at 16 °C or higher. In
both species, the incubation period
5. Eggs of lepidopterous
stem borers: a) Scirpophaga
incertulas, b) Scirpophaga
innotata, c) Chilo
suppressalis, d) Sesamia
inferens, e) hatching of an
egg mass.
6. Larvae of lepidopterous
stem borers: a) Scirpophaga
incertulas, b) Scirpophaga
innotata, c) Chilo auricilius,
d) C. suppressalis,
e) Sesamia inferens.
Stem borers 9
They usually feed individually. Upon developmental period. A more
hatching at dawn, diopsid larvae profound physiological change that
move down the stem and behind the enables stem borers to live for
leaf sheath on a film of dew. The eggs months in suspended development is
are dispersed and, normally, only one called diapause. Diapause can be
larva per tiller occurs. either hibernation (overwintering in
Chilo auricilius Dudgeon is primar- temperate climates) or aestivation
ily a pest of sugarcane and only (dry season dormancy in the tropics).
occasionally infests rice. Generally Scirpophaga incertulas and Scirpophaga
larvae infest grown-up canes only, as innotata hibernate or aestivate.
the mature larvae cannot make exit Depending on the site, Scirpophaga
holes through several leaf layers of incertulas is more prone to diapause
young canes. than Scirpophaga innotata, particularly
The threshold temperature for in the tropics. Stem borers, including
development of Chilo suppressalis diopsids, diapause as last-instar
larvae is 10.5-12 °C, but optimum larvae. Some diopsids diapause as
development is between 22 and 33 adults in swarms.
°C. The threshold temperature for Hibernation is broken by warm
Scirpophaga incertulas larvae, how- weather and longer daylengths;
ever, is a minimum of 16 °C. When aestivation is broken by rainfall or
reared at 12 °C, the second- and flooding. In the Philippines, with
third-instar larvae cannot molt and so multiple rice crops, Scirpophaga
die. The rate of larval development is incertulas is nondiapausing; in
positively correlated with tempera- Pakistan, with only a wet season
ture between 17 and 35 °C. crop, it overwinters in rice stubble. In
In identifying stem borer instars, Indonesia, Scirpophaga innotata does
many workers consider the width of not aestivate in double-cropped
the mandibles a better indicator than irrigated areas.
the width of the head capsule because
the mandibles are contiguous in Pupae
different instars. Scirpophaga incertulas Pupae of lepidopterous stem borers
larvae usually undergo four to seven are shown in Figure 7a-d. Pupation in
larval instar stages to become full lepidopterous rice stem borers
grown. Most larvae undergo five usually takes place in the stem, straw,
instars when reared at 23-29 °C, but or stubble. Diopsids pupate within
only four at 29-35 °C. The number of the stem. Sometimes Sesamia inferens
molts decreases in larvae feeding on also pupate between the leaf sheath
maturing plants compared with those and stem. Before pupating, the full-
feeding on tillering plants. Molts grown larvae cut exit holes in the 7. Pupae of lepidopterous stem borers:
increase where few host plants are internodes through which the emerg- a) Scirpophaga incertulas,
available. ing moths escape. Usually the exter- b) Scirpophaga innotata, c) Chilo
Under optimum conditions, Chilo nal opening of the exit hole is covered suppressalis, d) Sesamia inferens.
suppressalis has five to six larval with fine web and cannot be easily
instars. Under adverse conditions, detected before the moths have
such as those discussed above, as escaped. Chilo suppressalis pupae are
many as nine instars have been without cocoons, but pupae of
recorded. In lepidopterous and Scirpophaga spp., Rupela spp., and
diopsid species, the larval period Maliarpha spp. are covered with
usually lasts from 20 to 30 d. whitish silken cocoons. The anterior
Most stem borer species can pass extremity of the cocoons is tubular
an unfavorable period in dormancy. and attached to the exit holes; often
Drought during the larval period can one or two horizontal septa are
induce a temporary slowing down of webbed by the larvae in this tubular
body metabolism to prolong the area to make the cocoons waterproof.
Stem borers 11
species is governed largely by the essential for population buildup; the often plowed during that time is
number of crops grown in a particu- moths in cool areas are generally equally important.
lar area, especially in the tropics, the smaller and lay fewer eggs. If the The age and variety of the host
role of mature plants in inducing weather stays warm during the plants and the level of soil fertility
diapause is somewhat uncertain. The remaining rice crop seasons, the have an effect on the size of the stem
diapause of R. albinella and larvae develop rapidly and the total borer population. Generally, rice
Scirpophaga innotata terminates with number of generations may increase. plants in the vegetative phase and
higher precipitation. The problem is exacerbated particu- early heading stage receive more eggs
In places having distinct genera- larly in areas of multiple rice crops. than those nearing maturity. The
tions, the first generation usually Larvae suffer high mortality on extended periods of host plants at the
appears when the plants are in the seedlings. Some workers in Japan more attractive stages should there-
nursery or shortly after transplanting; attribute this to high water tempera- fore encourage a population increase.
the population increases in subse- ture. Increased larval mortality is For oviposition, stem borer moths
quent broods and the second or later recorded whenever the average prefer ricefields receiving high rates
generations are often the ones that temperature of floodwater exceeds of nitrogenous fertilizers. Rice plants
cause serious damage. This is why 35 °C for any 5 d in July. Measure- containing higher levels of N are
the borers are more destructive to the ments of the temperature of the more suitable for larval growth.
late-planted crop, or the second crop floodwater and within the rice stem The stem borer problem is more
where double cropping is practiced. suggest that temperature itself is not intense in areas with soils deficient in
Besides the seasonal fluctuations, directly lethal. Rather, high tempera- silica. Both field and laboratory
distinct annual fluctuations also ture might reduce larval vitality, studies have shown that larval
occur in stem borer populations. thereby increasing their vulnerability survival is significantly reduced if
Although the factors responsible for to bacterial diseases or other natural silica is applied to these soils. It has
such fluctuations are not fully under- hazards. also been demonstrated that the soil
stood, some of the possible causes are Larvae on seedlings used for mass itself renders rice plants less attrac-
the following. rearing have high survival, and it is tive to the insect, and the silica
Generally, all borer larvae suffer unlikely that the greater larval particles in the plant interfere with
low mortality during winter. In mortality in the field is due to nutri- larval feeding, often causing exces-
Japan, where the winter temperature tional deficiency. However, because sive mandible wear. A similar effect
is much lower than in most other rice the early-instar larvae feed gregari- of silica on stem borer larvae was
regions, mortality of Chilo suppressalis ously, the food available on the recorded in larvae reared on varieties
and Scirpophaga incertulas has been seedlings is inadequate and the containing different percentages of
low even during severe winters. Chilo larvae are forced to migrate much silica. Silica level also significantly
suppressalis is more tolerant of low earlier, probably resulting in high affects lodging and disease incidence
temperatures than Scirpophaga mortality. In areas of double crop- in the rice plant.
incertulas. In years of high precipita- ping, the seedlings of the second crop
tion during autumn, higher percent- carry a heavy egg load, leading to
ages of larvae hibernate, and if the subsequent high larval mortality.
winter or spring is warm, more of Such regulation of the population
these successfully pupate and emerge may not be operative, however,
as adult moths. These conditions, where planting seasons are not
however, also accelerate pupation distinct.
and emergence. Oviposition then Both in tropical and subtropical
occurs on seedlings, on which the regions, the population has been
larvae suffer high mortality and the reported to decline drastically during
population is reduced. However, if the summer months after the second
late spring is somewhat cooler and crop harvest. The decline has fre-
delays moth emergence, or if the rice quently been attributed to high
is planted slightly earlier, the popula- temperature, but the fact that most
tion builds up rapidly and heavy ricefields have been harvested and
damage may occur. Warm weather is
8. Damage caused by
stem borers: a) deadheart,
b) whitehead.
Stem borers 13
Control methods most effective. Since stubble is the Biological control
major source of the overwintering Most biological control of stem borers
Cultural control stem borer population, proper stub- in tropical Asia and Africa comes
Crop cultural practices have a pro- ble management cannot be overem- from indigenous predators, parasites,
found bearing on the stem borer phasized. and entomopathogens. The conserva-
population. Some methods are In several countries, delayed tion of these valuable organisms is
effective only if carried out through seeding and transplanting have been the key to development of stable and
communitywide cooperation; others effective in evading first-generation successful integrated pest manage-
are effective on a single field. Com- moths. This practice has not been ment (IPM) systems. Over 100 species
munitywide practices act to prevent highly effective against Chilo of stem borer parasitoids have been
colonization and have the greatest suppressalis in Japan since emergence identified. The three most important
potential to minimize infestation. is delayed if planting is delayed. It genera are the egg parasitoids
China and prewar Indonesia devel- has been effective, however, against Telenomus, Tetrastichus, and
oped effective cultural practices, Scirpophaga incertulas, the appearance Trichogramma. Tetrastichus wasps
often in combinations that isolate the of which is not affected by planting have elongated ovipositors and can
rice crop through time and space. dates. The number of generations of lay their eggs in stem borer eggs,
Practices that can be carried out on a this species is determined by the even if the latter are covered with a
single field include using optimal growth duration of the crop. Thus, mat of hair. Telenomus wasps, how-
rates of N fertilizer in split applica- where continuous rice cropping is ever, parasitize stem borer eggs while
tions. Applying slag increases the practiced, a change in planting time the moth is in the act of oviposition-
silica content of the crop, making it has little effect unless practiced over before the eggs are covered with hair.
more resistant. large areas. In such areas, crop The wasp locates the female moth,
Since the eggs of Scirpophaga rotation to include some short- possibly by the sex pheromone,
incertulas are laid near the tip of the duration nongraminaceous crops attaches itself to the tuft of anal hair
leaf blade, the widespread practice of should significantly reduce the borer near the ovipositor, and waits for the
clipping the seedlings before trans- population. moth to lay eggs.
planting greatly reduces the Changing planting time may not Egg masses are also the food of
carryover of eggs from the seedbed to always be feasible because of other several predators. The longhorned
the transplanted fields. However, this agronomic considerations. In Paki- grasshopper Conocephalus longipennis
control method has merit only if stan, the planting date has been (Haan) preys voraciously on eggs of
older seedlings are transplanted. regulated by releasing canal water the yellow stem borer. Other orthop-
Similarly, the height at which a crop only after the first brood Scirpophaga teran predators such as the crickets
is harvested is an important factor in incertulas moths have emerged. This Metioche vittaticollis (Stål) and
determining the percentage of larvae late-planted crop is far less infested Anaxipha longipennis (Serville) feed
that are left in the stubble. At harvest, than fields planted early with private on eggs of Chilo suppressalis. The
Chilo suppressalis larvae are usually irrigation systems. The early planted predatory mirid Cyrtorhinus
about 10 to 15 cm aboveground. fields, however, minimize the full lividipennis Reuter also attacks eggs of
Although Scirpophaga incertulas larvae impact of late planting on the stem Chilo suppressalis.
are located somewhat lower, most of borer population. In Japan, where A wide range of predatory species
them are aboveground as well. highly effective insecticides are attacks the small larvae of stem
Therefore, harvesting at ground level available, early planting has been borers before they enter the stem of
can remove a majority of the larvae of reintroduced at several sites, result- the rice plant. Some important
all species. To destroy those remain- ing in high survival of first-genera- predators are coccinellid beetles
ing in the stubble, burning or remov- tion Scirpophaga incertulas larvae. Micraspis crocea (Mulsant), Harmonia
ing the stubble, decomposing the Also, the first and second broods of octomaculata (Fabricius), and carabid
stubble with low rates of calcium Chilo suppressalis moths appeared beetles such as Ophionea spp. When
cyanide, plowing, and flooding have earlier, possibly introducing a distinct young larvae fall on the water, they
been suggested. Burning is only third generation in the warmer are preyed upon by Microvelia
partially effective because after sections of the country. Light-trap douglasi atrolineata Bergroth and
harvest the larvae generally move catches of moths reveal a change Mesovelia vittigera (Horváth). Ants
below ground level. It is also difficult from a unimodal to a bimodal pattern and a dozen other predators prey
to uniformly burn stubble in a field. in both the first and second broods. upon stem borer larvae.
Plowing and flooding are apparently
Stem borers 15
The rice breeding programs of environments. Granules broadcast bacteria respond to and rapidly
many countries aim at incorporating into irrigation water are particularly consume the insecticide, rendering it
into their improved germplasm genes effective in preventing deadhearts in ineffective. The process can be
for resistance to Chilo suppressalis and a young crop. Gamma BHC has a slowed by using lower dosages in
Scirpophaga incertulas from many fumigant action that kills resting rotation with foliar sprays. The
donors. However, none of the rice moths. The insecticide is partly problem with soil incorporation of
varieties developed so far have more dissolved in the water and moves by insecticides before planting is that the
than a moderate level of resistance. capillary action between the leaf stem borer population cannot be
Some wild rices such as Oryza sheath and stem to come into contact assessed—it might not be large
officinalis and O. ridleyi have very with young larvae: the nonsystemic enough to warrant control.
high levels of resistance to stem insecticide granules act as though A combination of sex attractant
borers. Their resistance needs to be they were systemic. The limitation to (pheromones) and chemosterilant
transferred to cultivated rice, using using granules is cost—they are more could also be a promising control
appropriate distant hybridization expensive to transport. Stable water tactic. High moth populations in
techniques. supply and deep water levels are also overlapping generations, however,
necessary for high levels of control. and the difficulties involved in mass
Chemical control As the water level falls, the capillary rearing some stem borer species are
Stem borers are difficult to control activity progressively declines. If the major limitations to the mass release
with insecticides. After hatching, the field dries out, insecticide efficacy of artificially irradiated sterile male
larvae are exposed only for a few ceases. Flooding from heavy rains moths as a control measure. Explora-
hours before they penetrate a tiller or also washes the insecticide out of the tory experiments on mass rearing
enter the plant. Successful control field. Dosage levels have declined, have shown that, when provided
involves repeated foliar applications consistent with the relatively higher with 1% tepa, apholate, or tretamine,
with spray volumes more than costs of insecticides. or 20% hempa as food, the moths
400 liters/ha. In temperate climates, Systemic granules have an advan- mated normally but deposited 50%
stem borer populations are more tage in that the chemical can enter the fewer eggs. Of the eggs deposited,
synchronized, and well-timed appli- plant even with low water levels. The 20% of those laid by moths exposed
cations have a greater degree of chemical percolates into the soil and to tepa and apholate were sterile.
control than in the tropics where is taken up by the roots. From the
generations overlap. The decline in roots, the chemical is transmitted
stem borer abundance in Japan and through the xylem tissues to the
the Republic of Korea is attributed to stems and eventually to the tips of
the frequent use of insecticides over the leaves. Carbofuran exudes in
many years, even though the stem droplets of water from leaf hyda-
borers have developed insecticide thodes and evaporates into the air. If
resistance. systemic granules are broadcast into
Foliar sprays, which act on the the irrigation water, high dosages are
larvae and on the adult moths and necessary because much of the
eggs, also come into greater contact chemical is absorbed in the soil. The
with natural enemies of the stem dosage needed increases with plant
borer. Cases of stem borer resurgence biomass. If granules are broadcast
are not evident, although secondary during the last harrowing or leveling
pest outbreaks have been reported in operation before planting, dosages
areas of heavy insecticide usage can be cut in half. Effectivity lasts
against stem borers. more than a month because the
Granular formulations, particu- granule is protected from rapid
larly gamma BHC and diazinon, give degradation. Heavy use of granules,
higher control than foliar sprays or however, can lead to microbial
dusts, particularly in high rainfall degradation. Several species of soil
Stem borers 17
Rice leafhoppers and
planthoppers
Several species of leafhoppers and Table 4. Major planthopper and leafhopper pests of rice.
planthoppers are serious pests of rice Name Common name Distribution Vector of
worldwide (Table 4). In many areas,
they frequently occur in numbers Delphacidae (Planthoppers)
Laodelphax striatellus Small brown China, Japan, Rice stripe, rice
large enough to cause complete (Fallen) planthopper Republic of streaked dwarf
drying of the crop, but even sparse Korea, Pale-
populations reduce rice yields. In arctic regions
addition to direct feeding damage, Nilaparvata lugens Brown planthopper South and Grassy stunt,
leafhoppers and planthoppers are (stål) Southeast Asia, ragged stunt
vectors of most of the currently China, Japan
Sogatella furcifera Whitebacked South and
known rice virus diseases. The more (Horvith) planthopper Southeast Asia,
damaging species are green northern Australia,
leafhoppers Nephotettix spp., the China, Japan,
Republic of Korea,
zigzag leafhopper Recilia dorsalis South Pacific
(Motschulsky), the brown Islands
planthopper Nilaparvata lugens (Stål),
Tagosodes orizicolus Rice delphacid Caribbean Islands Hoja blanca
the small brown planthopper (Muir) South America,
Laodelphax striatellus (Fallen), the southern United
whitebacked planthopper Sogatella States
Cicadellidae (Leafhoppers)
furcifera (Horváth), and the rice Cofana spectra White leafhopper South and Southeast
delphacid Tagosodes (=Sogatodes) (Distant) Asia, Australia,
orizicolus (Muir) (Fig. 9a-i). The first Africa, China
five species occur in Asia. Tagosodes Nephotettix cincticeps Rice green China (including Rice dwarf,
orizicolus is found in the southern (Uhler) leafhopper Taiwan), yellow dwarf
USA and in the north central region Japan, Republic
of Korea
of South America. Among several
Nephotettix species, three are Nephotettix virescens Rice green South and Yellow dwarf,
important. Nephotettix cincticeps (Distant) leafhopper Southeast Asia tungro, penya-
kit merah,
(Uhler) is distributed in temperate yellow-orange
areas. Nephotettix virescens (Distant) leaf
and Nephotettix nigropictus (=apicalis)
Nephotettix nigropictus Rice green South and South- Rice dwarf,
(Stål) are distributed in temperate (Stål) leafhopper east Asia, China yellow dwarf,
and tropical Asia. Nephotettix transitory
nigropictus is mainly distributed in yellowing,
tungro, yellow-
tropical Asian rice-growing areas. orange leaf,
In Africa, leafhoppers and rice gall dwarf
planthoppers do not cause serious
Recilia dorsalis Zigzag South and Rice dwarf,
loss, probably because lowland rice is (Motschulsky) leafhopper Southeast Asia; yellow-orange
not widely planted. The only re- Taiwan, China; leaf
ported hopperburn in Nigeria was Japan
19
9. Adults of rice leafhoppers
and planthoppers:
a) Nilaparvata lugens male,
b) Nilaparvata lugens female,
c) Sogatella furcifera male,
d) S. furcifera female,
e) Tagosodes orizicolus male,
f) T. orizicolus female,
g) T. cubanus, h) Nephotettix
virescens, i) Recilia dorsalis.
Photos e, f, and g courtesy of
CIAT.
The rice gall midge Orseolia oryzae infested than upland crops. Both gall generally a 4:l female-to-male ratio,
(Wood-Mason) is a serious pest of midge species prefer lowland to but Orseolia oryzivora has 1:l. The
rice in South and Southeast Asia. It is upland rice. Damage on upland rice adults feed on dewdrops and live for
widely distributed in several parts of in China and on deepwater rice in 2-5 d. Copulation usually takes place
India and in Cambodia, southern India has also been recorded. soon after emergence and lasts for
China, Indonesia, Laos, Myanmar, about 5 min. Oviposition starts a few
Nepal, Pakistan, Sri Lanka, northern hours later. The females mate only
Thailand, and Vietnam. The pest has Life history once and no parthenogenesis has
not been reported in the Philippines, been recorded; unmated females lay
Malaysia, or southern Thailand. Adults sterile eggs.
In Africa, another species of gall The adult flies of both Orseolia oryzae
midge — Orseolia oryzivora Harris and and Orseolia oryzivora are about the Eggs
Gape — damages the rice crop, but it size of a mosquito (wing length is A single female is capable of laying
is not considered a major pest. It is 3.5-4 mm for females and 3.0-3.5 mm 100-200 eggs, either singly or in
reported to occur in Cameroon, for males). They are nocturnal and groups of 3-4 near the base of the
Ghana, Ivory Coast, Liberia, Mali, phototropic (Fig. 12). The females plants, on the ligules or in their
Niger, Nigeria, Senegal, and Sudan. have a bright red, stout abdomen; the vicinity on the leaf blade, or on the
Since the fly requires high humid- abdomens of males are darker. The leaf sheath. In the seedbed as well as
ity, lowland crops are more often field population of Orseolia oryzae has after transplanting, about 60-70% of
the eggs are laid on the leaf blade,
and most of the rest on the leaf
sheaths, except for occasional ovi-
position on the central whorl of the
plants. Eggs laid on the leaf blade are
on the undersurface. In captivity, the
females oviposit on almost any
surface they come in contact with.
The eggs (elongate tubular, 0.5 mm
long and 0.12 mm wide) are shiny
white with pinkish, red, or yellow
shading, and turn shining amber
before hatching. The incubation
period is 3-4 d.
29
Larvae the tillering phase, after which the changes in water levels in the
The newly hatched larvae of both population rapidly declines, primar- ricefields do not seem to have a
species are about 1 mm long and can ily because of limited availability of distinct effect on fly incidence.
live in water up to 3 d without any suitable hosts. Therefore, a late-
harmful effect. They creep down the
leaf sheath to the growing points of
planted field is often severely dam-
aged, but early plantings may evade
Damage
the tillers and reach the interior of the gall midge infestation. In multi- Damage is caused by the transforma-
bud. There they lacerate the tissues cropped rice areas, the fly seldom tion of regular tillers into tubular
and feed until pupation. Their feed- infests the second crop and has not galls, which dry off without bearing
ing stimulates the tillers to grow into been observed infesting the third one. panicles. Early infestation results in
a tubular gall that resembles an onion Usually it occurs in five to eight profuse tillering of the plants, but
leaf. The average larval period of overlapping generations in one these new tillers often become in-
Orseolia oryzae is 15-20 d; that of season. fested and very few, if any, bear
Orseolia oryzivora is 6-22 d. The full- The insect has been reported to panicles. Even these panicles are less
grown larva of Orseolia oryzae is about hibernate in underground dormant vigorous and are often stunted.
3 mm long, pale red, with a pointed buds of grasses serving as alternate The pest starts infesting the plants
anterior end. It feeds at the base of hosts and becomes active again when in the seedbed and continues to do so
the gall where it pupates. Generally the buds start growing after rains. until the booting stage. Because the
only one maggot per tiller is found. Larvae overwinter in a short tube in larvae can develop only on the
The full-grown larva of Orseolia the stubble. The midge also multi- growing primordia, they cannot
oryzivora is whitish pink and 4.4 mm plies on sprouting rice stubble in survive on a crop beyond the vegeta-
long. Under high population pres- unplowed ricefields. In the tive stage.
sure, multiple and simultaneous Cameroons, after rice is harvested, The exact nature of the develop-
infestations occur. the pest migrates to wild rice Oryza ment of galls is not fully understood.
barthii, where it remains active until Either direct feeding in the develop-
Pupae the plants dry up in summer. Several ing primordia or the secretion of
Pupation occurs inside the gall near alternate host plants recorded in some compound by the maggot
the base. The pupa of Orseolia oryzae India and Thailand include wild rice stimulates the growth of the leaf
is about 2-2.5 mm long and Oryza officinalis and several gram- sheath around the larvae into an oval
0.6-0.8 mm wide; that of Orseolia inaceous weed plants such as chamber, which eventually grows
oryzivora is 5.8 mm long. The pupae Ischaemum aristatum, Echinochloa into a long, tubular, onion-leaf-like
are adorned with a series of subequal colona, Paspalum sp., and Leersia sp. gall. It is commonly called silver
spines pointed backwards. These The most serious gall midge shoot or onion leaf, and is usually the
spines enable the pupae to wriggle infestation occurs when early rains same color as the leaf sheath but
their way to the tip of the gall. When make the flies active. Subsequent dry somewhat shiny. Several earlier
the adults are ready to emerge, the periods delay rice planting; in this workers regarded galls as elongated
pupae pierce the tip and project case, the population multiplies on rice stems. But later anatomical
halfway out. The skin of the pupa grasses and the flies migrate in large studies of the galls established that
then bursts and the midge crawls out. numbers to the late-planted rice crop. they are modified leaf sheaths. The
Emergence generally takes place at Cloudy skies and drizzling rains are leaf blade is greatly atrophied and
night. The pupal period is 2-8 d. conducive to the fast buildup of gall remains attached as a leaflet with tiny
midge populations. The favorable ligules and auricles on the tip of the
condition for fly development is gall.
Seasonal occurrence 26-30 °C and 82-88% RH. Heavy rains The galls become apparent 3-7 d
and abundance or storms cause high mortality. The
insects are less abundant in crop
after the larvae enter the growing
point of the plants. A fully developed
The fly becomes active at the onset of years preceded by a warm and dry gall is generally 1-2 cm wide and 10-
the monsoon when it completes one spring. 30 cm long, although galls as long as
to two generations on grasses before Low RH may be the cause of the 50 cm have been recorded. By the
rice is planted, and then transfers to decline in gall midge during the time galls become visible, the adults
the ricefields. It usually takes 9-14 d second rice crop even though there is have generally emerged. However,
to complete one life cycle on grasses, no distinct lack of suitable hosts.
but about 9-26 d on rice. The fly is Because the larvae can live under
capable of infesting the crop only at submergence for several days,
Eight species of pyralid moths, whose Several alternate hosts are re- The freshly hatched larva is
larvae roll or fold leaves of gram- ported for the rice leaffolders. Aside 1.5-2 mm long and 0.2-0.3 mm wide,
inaceous plants, comprise the leaf- from rice, they attack various grasses, has a shiny translucent body and a
folder complex. They are sedges, and cultivated crops such as light-brown head. The body turns
Cnaphalocrocis medinalis (Guenée), maize. green after the larva begins feeding.
Marasmia patnalis Bradley, Marasmia The first-instar larva feeds on the
(=Susumia) exigua (Butler), Marasmia young leaves by scraping the leaf
bilinealis Hampson, Marasmia ruralis Life history surface, but it does not cause the
(Walker), Marasmia suspicalis leaves to fold (Fig. 13d). Its body
(Walker), Marasmia trapezalis The life history of rice leaffolder usually is covered with a silky
(Guenée), and Marasmia venilialis species is more or less similar. material. Five larval instars are
(Guenée). Cnaphalocrocis medinalis is the best completed in an average period of
Leaffolders are widely distributed known leaffolder species in Asia. The 20-30 d. The larvae from the late
in the rice-growing tracts of 29 humid adult moths are 10-12 mm long. They second instar onward can cause the
tropical and temperate countries in are light brown with shiny, brownish leaves to fold (Fig. 13e). The caterpil-
Asia, Oceania, Australia, and Africa yellow wings adorned with dark, lar secretes a series of threads and
between 48° N and 24° S and 0° E to broad margins, and two to three dark uses these to connect the two margins
172°W. Currently, only three species, vertical stripes. Wing expanse is of a leaf blade. The threads contract
Cnaphalocrocis medinalis, Marasmia 13-15 mm. The moths are nocturnal; as they dry and bring the two leaf
patnalis, and Marasmia exigua, have they hide during the day and usually margins together, turning the leaf
attained pest status on rice. emerge at night. They generally mate blade into a tubular structure. The
Cnaphalocrocis medinalis tradition- between dusk and midnight. larva remains within the leaf blade
ally has been accepted as a sole Oviposition starts two-three nights and feeds on it by scraping the leaf
leaffolder pest in the lowland after mating. The female moth usu- surface, thus causing longitudinal,
ricefields of Asia. But the discovery of ally lives for as long as 8-9 d and lays white, transparent streaks on the leaf
Marasmia patnalis in 1981 has compli- 50-300 eggs. The eggs are laid on blade. If a blade is severely damaged,
cated the interpretation of past different nights in batches of 10-12 the larva migrates to other leaves.
results. Marasmia patnalis has often arranged linearly along the midrib on The full-grown larva is yellowish
been confused with Cnaphalocuocis either surface of the leaf blade green with a dark brown head and is
medinalis in South and Southeast Asia (Fig. 13c). The biggest batches are laid 20-25 mm long and 1.5-2 mm wide. It
(Fig. 13a, b). The individuals belong- on the fourth-seventh night after the undergoes pupation inside the leaf
ing to the two genera could be moths emerge. roll in loosely woven strands of silk
differentiated from each other by The individual eggs are translu- thread. The newly formed pupa is
forewing venation. Cnaphalocrocis has cent, yellowish white, oval, 0.90 mm light brown, but turns reddish brown
R2 and R1 (veins 10 and 11) stalked, long and 0.39 mm wide, and almost toward moth emergence in 6-10 d.
with R2 set close to the trunk of R3 flat with a slightly convex surface.
and R4 (veins 8 and 9). Marasmia, on The eggs hatch 3-4 d after oviposi-
the other hand, has R2 and R1 free. tion.
These and other morphological as
well as genitalial features are used to
separate the leaffolder species.
33
13. Rice leaffolders: a) Cnaphalocrocis
medinalis adult, b) Marasmia patnalis
adult, c) C. medinalis eggs, d) damage
by a second-instar larva of C. medinalis,
e) rice leaf rolled by a third-instar larva
of C. medinalis, f) a ricefield damaged by
leaffolders.
Rice leaffolders 35
rices— Oryza brachyantha, O. nivara, Selected references
O. rufipogon, and O. perennis -also
Bradley J D (1981) Marasmia patnalis sp. n.
show resistance to Cnaphalocrocis (Lepidoptera: Pyralidae) on rice in S. E.
medinalis and Marasmia patnalis. Asia. Bull. Entomol. Res. 71:323-327.
Research centers with several na-
Chandramohan N, Jayaraj S (1977) Effect
tional rice improvement programs of different levels of nitrogen and age
are also screening rice varieties and of the crop on the incidence of rice
wild germplasm for resistance to leafroller. Madras Agric. J. 64:648-685.
Cnaphalocrocis medinalis; several rice Hanifa M A, Subramanian T R, Ponnaiya
varieties and breeding lines with B W X (1974) Role of silica in resistance
potential for resistance have been to the leafroller, Cnaphalocrocis
identified. Unfortunately, all IR rice rnedinalis Guenée, in rice. Indian J. Exp.
varieties (IR5 to IR72) are susceptible Biol. 12:463-465.
to leaffolders. Improved varieties
Heinrichs E A, Camafiag E, Romena A
with resistance to the leaffolder (1985) Evaluation of rice cultivar for
complex are needed because varietal resistance to Cnaphalocrocis medinalis
resistance is one of the more promis- Guenée (Lepidoptera: Pyralidae). J.
ing tactics in the integrated control of Econ. Entomol. 78:274-278.
this pest in tropical Asia. Hinckley A D (1963a) Insect pests of rice
in Fiji. FAO Plant Prot. Bull. 2(2):31-33.
Chemical control Hinckley A D (1963b) The rice leafroller,
Chemical control is the only practical Susumia exigua (Butler), in Fiji. J. Econ.
method to control increasing leaf- Entomol. 56:112-113.
folder infestation during crop Khan Z R, Joshi R C (1990) Varietal
growth. However, socioeconomic resistance to Cnaphalocrocis medinalis
constraints in tropical developing (Guenée) in rice. Crop Prot. 9:243-251.
countries and insecticide-induced Khan Z R, Barrion A T, Litsinger J A,
resurgence of Nilaparvata lugens Castilla N P, Joshi R C (1988) A
present obstacles to effective chemical bibliography of rice leaffolders (Lepi-
control. Numerous insecticides that doptera: Pyralidae). Insect Sci. Appl.
have been identified for control are 9:129-174.
most effective as foliar sprays. But Lim L L (1962) Biology of the rice leaf-
foliar sprays have to be repeated folder (Cnaphalocrocis medinalis
because they are often washed off by Guenee). BS thesis in Entomology,
frequent rains. Granular insecticides University of the Philippines at Los
broadcast into water are ineffective. Baños, Laguna, Philippines.
Economic thresholds, used to make Reissig W H, Heinrichs E A, Litsinger J A,
decisions on insecticide application, Moody K, Fiedler L, Mew T W, Barrion
are based on egg counts, damaged A T (1986) Illustrated guide to inte-
leaves, or larval densities. Since grated pest management in rice in
tropical Asia. International Rice
leaffolders can attack the crop during
Research Institute, P.0. Box 933,
any growth stage, fields should be
Manila, Philippines. 411 p.
monitored weekly.
to the families Alydidae and Pentato- Common name Family Scientific name Distribution
midae (Table 6) that feed by sucking
the sap of developing spikelets cause Rice bug Alydidae Leptocorisa acuta (Thunberg) Asia,
Australia
serious rice crop losses. Usually, they L. biguttata (Walker) Asia
live either in the ricefields or on L. chinensis (Dallas) Asia
grasses in the vicinity where they L. palawanensis Ahmad Asia
L. oratorius (Fabricius) Asia,
feed and multiply during the vegeta- Australia
tive phase of the rice crop. They then L. solomonensis Ahmad Asia
migrate to flowering ricefields, which Riptortus linearis Fabricius Asia
Stenocoris southwoodi Africa, South
strongly attract them. In Asia, several Ahmad America
species of Leptocorisa are the most S. claviformis Ahmad Africa, South
important grain-sucking pests; in the America
USA, the rice stink bug Oebalus Stink bug Pentatornidae Oebalus pugnax (Fabricius) Southern USA
pugnax (Fabricius) is of major impor- O. poecila (Dallas) Latin America
tance. O. ypsilon-griseus (De Geer) Latin America
O. grisescens (Sailer) Latin America
Eysarcoris (=Stollia) Asia
ventralis (Westwood)
Pygomenida varipennis Asia
Rice bugs
(Westwood)
P. benghalensis (Westwood) Asia
Nezara viridula Linnaeus Asia, Africa
37
mating. A single female lays an reduce it. The population usually
average of 200-300 eggs in batches of increases at the end of the rainy
10-20, usually arranged in two or season and declines rapidly during
three straight rows along the midrib dry months and when temperature is
on the upper surface of the leaf blade. unfavorable. When temperature
The eggs are oval, slightly flattened declines from October onward, the
on the top (Fig. 14b). They are creamy insects hibernate in grasses. In such
white when freshly laid, but become areas, late rice crops escape rice bug
darker as they approach hatching in infestation. The hibernating adults
5-8 d. During hatching, the upper half become active with the onset of
of the egg breaks away, leaving a summer rains. Intermittent rains and
distinct hole. high temperature during summer are
The freshly emerged nymphs are conducive to terminating the dia-
tiny and green, but become brownish pause.
as they grow. They blend with the Factors that cause high rice bug
foliage and are often undetected. populations are nearby woodlands,
They start feeding 3-4 h after hatch- extensive weedy areas near ricefields,
ing, and undergo five nymphal and staggered rice planting.
instars in a total period of 25-30 d to After diapause, the adults feed on
become adults. The feeding habits of weeds and other available alternate
adults and nymphs are similar. hosts on which they pass one to two
A number of alternate host plants, generations before migrating to the
all in the family Gramineae, have rice crop now at flowering stage. In
been recorded. The insects live on Japan, the postdiapause adults of
grasses but prefer flowering rice. L. chinensis migrate from mountain-
Both nymphs and adults are ous areas to upland ricefields during
difficult to see in the ricefields be- flowering. In single-cropped areas,
cause their color resembles that of the insect usually has four overlap- 14. Grain-sucking insect
pests of rice:
rice plants. Infested fields, however, ping generations. But at places where
a) Leptocorisa oratorius
can often be detected, even from a temperatures are optimum and rice is adult, b) L. oratorius eggs,
distance, because they emit a typical grown year-round, the bugs remain c) adult of a pentatomid
rice bug odor produced by scent active throughout the year without a bug.
glands on the abdomen of the insect. distinct diapause. In such areas, fields
When disturbed, adults fly and that mature earlier or later than the
give off the typical rice bug odor. usual crops become heavily infested.
Adults are active in late afternoon Insect abundance becomes especially
and early morning. They rest in high where rice planting is staggered,
grassy areas during periods of bright leading to an extended ripening
sunshine. During the dry season, period over several crops. The strong
adults move to wooded areas where phototropic nature of the insect has
they remain dormant. led some workers to explore light
traps as a control method, but the
results have been erratic. Large rice
Seasonal occurrence bug populations can also be trapped
and abundance in early planted ricefields.
Grain-sucking insects 39
Adults emerging from hibernation
feed on grasses, orchards, and other
usually highest during July, which
coincides with the flowering of the
Control methods
spring and summer crops before they first rice crop. In February, when the Control measures for all grain-
migrate to rice. The attraction to second crop is flowering, the popula- sucking insects are similar.
heading rice is so great that adults tion is comparatively small but still of
migrate to ricefields over several economic significance. The popula- Cultural control
kilometers. tion is at its lowest ebb in May. Various cultural and mechanical
In nature, the population is, to control measures are being adopted
some degree, density dependent. to control grain-sucking bugs. De-
With increased population density in Damage layed, but synchronous, planting of
the second and third generations, the early-maturing varieties is suggested
females exhibit reduced fecundity The damage to the rice crop is caused so that all crops ripen at the same
and the eggs and larvae suffer higher by the feeding of nymphs and adult time. Weed sanitation and eradica-
mortality. Strong winds also reduce bugs on the endosperm of the devel- tion of alternate hosts from ricefields,
the nymphal population. In Japan, oping grains. Growing rice bug levees, and surrounding areas also
several egg parasites play an impor- nymphs are more active feeders than prevent the multiplication of the bug
tant role. Substantial biological adults, but adults cause more damage in rice-free periods. Mechanical
control of this insect has been because they feed for a longer period. control measures such as smoking the
achieved in Hawaii and Australia. Although they also feed on other field, hand-picking of adults and
The adult insects of O. pugnax parts of the rice plant, they prefer nymphs, and the use of sticky traps
overwinter in leaf trash or bunch grains at the milk stage and even have been suggested.
grasses. Emerging from hibernation ripening grain. Both nymphs and
in late April and early May, the adults feed by inserting their probos- Varietal resistance
insects feed on developing seeds of cises at points where palea and All cultivated rice varieties are
grasses adjacent to ricefields and lemma meet. Diffused brown spots susceptible to grain-sucking pests.
move to the ricefields later. A severe caused by the exudation of the sap IRRI attempts to find resistance to
winter reduces the population of mark the points of insertion. Grains Leptocorisa spp. located a rice from
overwintering adults, and high damaged during milk stage remain India in a group of coarse-ground,
temperature during summer causes empty. The panicles in heavily low-yielding rice called Sathi. In this
high nymphal mortality. infested fields contain many rice, the panicle remains enclosed in
Another species, Oebalus (=Solubea) shriveled and unfilled grains and the leaf sheath and offers a sort of
ornata (Sailer), has been reported to usually remain erect. In severe cases mechanical resistance to insect
cause serious losses to rice in the of infestation, most grains in a field sucking. However, the compatibility
Dominican Republic. It frequently are sucked empty and the straw has of this character with yielding ability
occurs in large numbers, causing up an off-flavor, which is unattractive to and ease of threshing is uncertain. In
to 50% crop loss. cattle. When rice bugs feed in soft or India, varieties Mundagakutty from
In general, the bionomics of this hard dough endosperm in a solid Tamil Nadu and Soma from Bihar
pest resembles that of O. pugnax. The state, they inject enzymes to predigest also show similar resistance.
average female-to-male ratio is 1:1. it. Damage during the dough stage Pentatomid bugs bore through the
Most of the eggs are laid on the upper causes discoloration of mature grain lemma or palea, but hairy seeds or
leaf surface and first-instar nymphs and causes weakness in the kernel. seeds with awns that deter bugs draw
congregate around the egg shells. The Such rice has lower milling quality or complaints from threshers. Although
pests are extremely active up to is pecky rice of inferior grade. Par- screening for resistance to rice bug
1000 h when they feed on the panicles tially damaged grains also have an has been limited, moderate levels of
and copulate. They migrate to the off-flavor even after cooking. Injury resistance to the rice stink bug has
base of the plants when the sun during the milk stage causes yield been reported in some rice varieties
becomes brighter. On cloudy or light loss; damage during the dough stage such as Bluebelle, Nortai, PI 9810,
rainy days, the adults and nymphs impairs grain quality. and RU7603069.
remain on the panicles all day and The southern green stink bugs
feed extensively. In the Dominican feeding on rice are generally larger
Republic, they remain active through- than those feeding on grasses and
out the year and occur in seven other hosts.
generations. The population is
Grain-sucking insects 41
Rice hispa
Rice hispa Dicladispa armigera of the leaf blade, generally on the Seasonal occurrence
(Oliver) frequently causes extensive ventral surface, and are partially
damage to lowland rice crops in inserted beneath the epidermis and and abundance
Bangladesh, China (including Tai- covered with a small quantity of a
wan), India, Indonesia, Japan, Myan- dark substance probably secreted by The pest occurs throughout the year
mar, Nepal, Pakistan, West Malaysia, the female beetle. A single female in tropical areas, but is generally
and southern Thailand. The pest is lays an average of 55 eggs; under more numerous during the rainy
not reported on upland rice and heavy infestation, as many as 100 season. In places having cool winters,
prefers more aquatic habitats. In eggs have been recorded from a the insect remains active from May to
Central Africa, a species of rice hispa single plant in the field. The incuba- October and completes four or five
Dicladispa viridicyanea (Kraatz) attacks tion period under natural field generations.
upland rice in the vegetative stage as conditions is 4-5 d. Heavy rains, especially in premon-
well as in lowland seedbeds. Trichispa The pale yellow grubs, dorsoven- soon or earliest monsoon periods,
sericea (Guerin), an African species, trally flattened and 2.4 mm long, followed by abnormally low precipi-
damages upland rice in West Africa hatch and mine immediately into the tation, minimum day-night tempera-
and Madagascar. Three other hispid leaf blades between the epidermal ture differential for a number of days,
insects—Dactylispa dilaticornis, membranes and feed on the green and high RH favor rapid buildup of
Rhadinosa lebongensis Maulik, and tissue. The larval period is 7-12 d; the hispa populations. Incidence is
Leptista pygmaea Baly—damage the pupal stage is 4 or 5 d. Both stages are generally higher in fields treated with
rice crop in several states of India. completed without migration to any higher rates of N and where plants
Dicladispa gestroi (Chapuis) is another other leaf. The adult beetles cut their grow thickly under shade. The leaves
important hispid insect reported from way out of the leaf and begin to lay of semidwarf varieties are more
the Malagasy Republic. eggs 3 or 4 d later. Adults are external heavily infested than are those of
feeders. conventional local varieties.
At initial infestation the adult
Life history beetle prefers the young rice crop. In
India, the adults generally appear in
The adult of Dicladispa armigera is a ricefields in February and the popula-
small (5.5 mm long), shiny, blue- tion increases until June or July when
black beetle with a spiny body the grubs as well as the beetles cause
(Fig. 15). Adult beetles of Didadispa heavy losses to young crops. After
viridicyanea are metallic blue-green, August the population declines and
about 5 mm long, with five lateral the pest is usually of no economic
spines on each side of the thorax and significance. Adult beetles in small
a series of alternately long and short numbers can be collected from
lateral spines on the elytra. The ricefields up to September or October.
females of Dicladispa armigera live an During this period, the pest com-
average of 20 d and the males, 14 d. 15. Adult of rice hispa Dicladispa pletes six overlapping generations.
They mate 3 or 4 d after emergence. armigera.
The eggs are laid singly near the tip
43
Damage Biological control
Selected references
The role of natural enemies in con-
Alam M Z (1967) Insect pests of rice in
Both adults and grubs feed on and trolling hispa has not been fully
East Pakistan. Pages 643-655 in The
damage rice plants. The adults scrape assessed. However, several braconid major insect pests of the rice plant.
the upper surface of the leaf blade, wasps such as Bracon hispae (Viereck), Proceedings of a symposium at The
often leaving only the lower epider- Bracon sp., Campyloneurus sp., and International Rice Research Institute,
mis. The damaged areas appear as Macrocentrus sp. are known to para- September 1964. The Johns Hopkins
white streaks parallel to the midrib. sitize the larvae. Likewise, the Press, Baltimore, Maryland.
The tunnelling of the grubs between ichneumonid wasp Isotima sp. and Banerjee S N (1964) Paddy pests. Pages
the two epidermal layers results in two unidentified species of eulophid 292-296 in Entomology in India.
irregular translucent white patches are also larval parasites. Pupal Entomological Society of India.
starting from ovipositional sites near parasites include three pteromalid Chand P, Tomar J B (1984) Screening for
the leaf tip and extending toward the species: Eupteromalus sp., resistance to rice hispa. Int. Rice Res.
base of the leaf blades. The affected Trichomalopsis apanteloctena Newsl. 9(2):6.
parts of the leaves usually wither off. (Crawford), and Serotenus sp. An Dhaliwal G S, Singh J (1979) Effects of
In severe infestations, the leaves turn unidentified trichogrammatid species spacing on rice hispa incidence. Int.
whitish and membranous and finally parasitizes the eggs of this pest. A Rice Res. Newsl. 4(2):19.
dry off. In Kenya, Trichispa sericea can reduviid Rkinocoris fuscipes Feakin S D (1970) Pest conrol in rice.
transmit rice yellow mottle virus. (Fabricius) is a recorded predator of PANS Manual 3. Centre for Overseas
Infested plants have reduced leaf hispa adults. Pest Research, London. 270 p.
area, become less vigorous, and are Krishnaiah N V, Kalode M B (1983)
often stunted. In recent years, hispa Varietal resistance Effectiveness of some granular and
has been a perpetual problem in At present there are no varieties with spray formulations against rice hispa.
Bangladesh, infesting several thou- proven resistance to hispa, but Pesticides 17(7):25-28.
sand hectares annually and causing OR165-94-1 and KAU1945 have been Krishnaiah P V, Sanjeava Ras P, Ras
affected areas to suffer a significant reported as moderately resistant. In- N H P, Seshaguri Ras P, Narasimham
yield loss. The insect also attacks dications are that some cultivars such V (1987) Control of rice hispa. Indian
sugarcane and some wild grasses. as MTU6637, RNR1446, IET26889, Farming 37(7):37-39.
Norin B, IR579, and Manoharsali are Prakasa Rao P S, Israel P, Rao Y S (1971)
less preferred than others. Epidemiology and control of the rice
Control methods hispa, Dicladispa armigera Oliver.
Chemical control Oryza 8(2) (suppl.):345-359.
Cultural control Chemicals play an important role in Razzaque Q M A, Rezaul Karim A N M,
Close plant spacing results in greater rice hispa control. Adults are more Kamal N Q (1986) Studies on some
leaf densities that can tolerate higher exposed and susceptible to insecticide bio-ecological aspects of rice hispa,
hispa numbers. Bunds and vicinities than are the larvae, which are pro- Dicladispa armigera (Oliver)
should be kept free from grassy tected in leaf mines. Sprays and dusts (Hispidae:Coleoptera). A paper
weeds on which these beetles can are more effective than granular presented at the SAARC Workshop on
maintain their population. Stubble formulations. Systemic insecticides Rice Hispa, 28-29 December 1986.
Bangladesh Rice Research Institute,
should be uprooted after harvest to give longer residual protection and
Joydebpur. 12 p.
avoid ratooning. In some countries, are more effective against larvae than
Reissig W H, Heinrichs E A, Litsinger J A,
leaf tips are clipped off before trans- are nonsystemic chemicals.
Moody K, Fiedler L, Mew T W, Barrion
planting to eliminate early stages of
A T (1986) Illustrated guide to inte-
the pest. Excess N application should grated pest management in rice in
be avoided. Planting early at the tropical Asia. International Rice
beginning of the monsoon allows a Research Institute, P.O. Box 933,
field to escape hispa buildup. Hand- Manila, Philippines. 41 1 p.
picking damaged leaves removes Singh O P (1985) New record of
larvae from the field and prevents Rhinocoris fuscipes Fabricius, as a
hispa buildup. Damaged leaves can predator of Dicladispa armigera
be removed up until booting. (Oliver). Agric. Sci. Digest 5(3):179-180.
The rice water weevil Lissorhoptrus water surface. The adults feed and under deep snow and at very low
oryzophilus Kuschel, which was copulate on the aerial plant parts, but temperature. In Japan, 60% of the
originally distributed in the Missis- oviposit on submerged plant parts. adults survive until April under
sippi River basin in the USA, is now Cylindrical, pearly white eggs, about -18.6 °C (minimum) air temperature
one of the most destructive rice pests 0.8 mm long, are individually in- and -6 °C (minimum) soil tempera-
in Japan. The weevil was recorded for serted in the basal half of the sub- ture. Laboratory experiments on
the first time in 1959 in California. merged portion of the leaf sheath, overwintering adults exposed to
Only females were found and the and only occasionally on the sub- various low temperatures for 3 mo
weevil reproduced parthenogeneti- merged upper portion of the plant or showed that the critical minimum
cally. The pest was found in Cuba in the roots. Under field conditions, temperature ranges from -5 to -10 °C.
and the Dominican Republic in 1972. the eggs hatch in an average of 8 d. The supercooling point of overwin-
The Dominican Republic strain also The larvae hatching from eggs laid in tering adults under wet conditions is
consists only of parthenogenic the leaf sheath mine the leaf sheath from -6.2 to -17.6 °C. The adults
females. for a short period and then crawl move into the fields with young rice
In Japan, the weevil was first down to the roots. The larvae hatch- plants and feed on the leaves. In
recorded in 1976 in Aichi Prefecture. ing in the roots remain there to feed. unflooded fields, they hide in the soil
Only parthenogenic females with There are four larval instars, and it during the day and feed at night, but
triploid chromosomes are found. takes 30-40 d for the larvae to reach in flooded fields they usually feed
They are probably derived from the adulthood. The larger third-instar day and night. Oviposition usually
California population. The Nayoga larvae feed externally among the occurs after the field is irrigated.
Plant Quarantine Station suggests roots and sometimes up into the Newly emerged adults usually fly at
that the insect arrived in Japan with crown. Several larvae are often found night to adjacent fields of younger
hay imported from the USA. By the in the roots of a single plant. The rice.
end of the year, the insect was con- larvae can migrate up to 15 cm Larval population density varies
firmed to have occurred in 730 ha of through the soil to other roots. considerably with time of transplant-
ricefields in three cities and two Pupation occurs in a cocoon attached ing and with time when rice is first
towns in the Prefecture. to the roots. The pupa is white and of flooded. When rice seedlings are
Since 1976 the rice water weevil the same size as the adult weevil. transplanted in mid-May in Japan,
rapidly spread. By the end of 1986, adult females are most abundant and
more than 1 million ha, or 46% of the yield loss is highest. In the USA,
total ricefields in Japan were infested. Seasonal occurrence ricefields flooded very early or late
The insect is now regarded as the
number one pest of rice in Japan and
and abundance are less infested than those flooded in
between. In addition, young rice
the most difficult one to control. The adult weevil overwinters in plants are more heavily infested than
Spanish moss, rice stubble, or fine older plants. Larval, pupal, and adult
matted grass and becomes active populations are generally higher in
Life history again as the weather warms up from fields with standing water through-
March to June. The survival rate of out the cropping season than in fields
The adult weevil is about 3 mm long the overwintering adults is high in flooded 1 mo after transplanting.
and is greyish brown with a darker favorable hibernating sites such as
area on the dorsum. It is semiaquatic groves, but low in dry leaves. Over-
and can fly or swim just beneath the wintering adults can survive even
45
Damage are known. Varieties with moderate Matsui M (1987) Expansion of distribu-
tion area of rice water weevil and
levels of resistance are Ae Guk Ai
Adults feed on leaves of young rice Koku, Iljin, and Mit Dari from the methods of controlling the pest in
plants. The resulting longitudinal Republic of Korea; Toyokuni and Japan. JARQ 20:166-173.
strips on the leaf surface usually are Mogami mochi from Japan; IR269-1- Rolston L H, Rouse P (1964) Some factors
not of much economic significance. 1-3, IR404-1-3-1-1, IR404-3-2-7, IR404- influencing larval infestations of the
The major damage is caused by the 6-3-10-1, and IR455-5-5-1-2 from the rice water weevil. J. Kansas Entomol.
maggots, which feed within and Philippines; and CI 9903 and CI 8900 Soc. 37.
upon the roots, pruning them se- from the USA. Smith C M, Robinson J F (1982) Evalu-
verely in heavy infestations and ation of rice cultivars grown in North
producing loss of vigor, lodging, and Chemical control America for resistance to the rice water
weevil. Environ. Entomol. 11:334-336.
reduced yields. The most important Granular insecticides applied at the
factor for yield loss is the decrease in proper time control the rice water Smith K A, Grigarick A A, Oraze M J
number of tillers and panicles. weevil. (1988) Field evaluations of de-
flubenzuron and triflumuron for
control of the rice water weevil in
Control methods Selected references California rice fields. J. Agric. Entomol.
5(2):121-126.
Cultural control Bang Y H, Tugwell N P (1976) Adult rice Takagi Y (1989) Benfuracarb (OncolR), a
Intermittent draining and flooding of water weevil feeding references for new broad-spectrum carbamate
ricefields at 15-d intervals considera- rice plants and leaves of different ages. insecticide. New pesticides. Jpn. Pestic.
bly reduce the damage caused by rice Arkansas Agric. Exp. Stn. Rep. Ser. Inf. 54:23-27.
231.12 p. Tugwell N P, Stephen F M (1981) Rice
water weevil. However, this proce-
dure is prohibitive in many areas Bowling C C (1967) Insect pests of rice in water weevil seasonal abundance,
because of limited water supply and the United States. Pages 551-570 in The economic levels, and sequential
major insects pests of the rice plant. sampling plans. Arkansas Agric. Exp.
loss of fertilizers. In Japan, trans-
Proceedings of a symposium at the Stn. Bull. 849. 16 p.
planting rice seedlings early in April
International Rice Research Institute, Way M 0, Wallace R C (1988) Susceptibil-
significantly reduces yield loss. September 1964. The Johns Hopkins ity of selected Texas rice genotypes of
Damage can also be reduced by Press, Baltimore, Maryland. the rice water weevil 1986-1987. Tex.
transplanting mature or middle-age
Bunyarat M, Tugwell P, Riggs R D (1977) Agric. Exp. Stn. Prog. Rep. 4563.7 p.
seedlings. Plots receiving greater Seasonal incidence and effect of a
amounts of fertilizers are more mermithid nematode parasite on the
severely infested. Removal of aquatic mortality and egg production of rice
grasses, which are alternate hosts, water weevil, Lissorhoptrus oryzophilus.
reduces the pest population. Environ. Entomol. 6:712-714.
Cave G L (1983) Biological, ecological,
Biological control and morphological investigations of
An undescribed mermithid almost rice water weevil, Lissorhoptrus
exclusively parasitized female wee- oryzophilus Kuschel, on two rice
vils in the USA. Several bird species genotypes. Ph D thesis, Louisiana State
and frogs are known to ingest the University, Louisiana, USA. 78 p.
weevils. Tettigoniid grasshoppers Cave G L, Smith C M (1983) Number of
such as Conocephalus fasciatus (De instars of the rice water weevil,
Geer), Neoconocephalus triops Lissorhoptrus oryzophilus (Coleoptera:
(Linnaeus), and Orchelimum agile Curculionidae). Ann. Entomol. Soc.
Am. 76:293-294.
(De Geer) prey upon adult weevils.
The fungus Beauveria bassiana Everett T (1965) Rice insects. Rice J. 68:
(Balsamo) Vuillemin is known to 28-30.
infect the rice water weevil in Japan. Grigarick A A, Beards G W (1965)
Ovipositional habits of the rice water
Varietal resistance weevil in California as related to a
High levels of resistance to rice water greenhouse evaluation of seed treat-
weevil are not found in any rice ments. J. Econ. Entomol. 58:1053-1056.
genotype. Several rice varieties with
moderate or low levels of resistance
Thrips have been recorded as pests of S. biformis is also known as rice Life history
rice from most of the rice-growing leaf thrip, paddy thrip, or oriental
countries of the world in all rice rice thrip. The first specimens were S. biformis is a minute, fragile-looking
environments. The two most com- collected in 1913 in Oxford, England, insect, usually 1-2 mm long, with
mon species associated with rice are from sedge stacks, accumulations of well-pronounced five- to eight-
Stenchaetothrips (=Baliothrips, =Thrips) plant materials consisting mainly of segmented antennae (Fig. 16a). The
biformis (Bagnall) (Thripidae) and Phragmites and Phalaris, removed adults can be winged or wingless.
Haplothrips aculeatus (Fabricius) from water courses. The species is Both pairs of wings are elongated,
(Phlaeothripidae). In Bangladesh, one of the very few thrips that live narrow, and fringed with long hairs.
China, India, Indonesia, Japan, and only on plants growing in water. The It is this latter character that gives the
Sri Lanka, S. biformis is now consid- abundance, host plant preference, order the name thrips. These hairs
ered a major rice pest. It attacks rice and distribution of other thrip species arise from sockets, except in the
seedlings in nurseries as well as associated with rice are given in family Phlaeothripidae (such as
small, newly transplanted plants. Table 7. H. aculeatus), as extensions of the
Unlike other thrips, it apparently wing membrane. Parthenogenetic re-
does not attack more mature plants production, with haplodiploidy
and is not found in the panicles. process of sex determination, is
common in thrips.
Thrips have a relatively short life
Table 7. Common thrips pests of rice. cycle and can multiply rapidly. The
life history of each species features an
Species Abundance Host plant Distribution
egg, two active larval instars that
Haplothrips aculeatus Very common Polyphagous: grasses, Taiwan and northeastern feed, followed by two or three (as in
(Fabricius) cereals, many kinds China, India, Indonesia, the case of Phlaeothripidae) relatively
of flowers Japan, Republic of Korea,
Philippines, USA, northern inactive pupal instars that probably
USSR, Thailand, Africa, do not feed, and adult. Adults are
Europe found inside rolled leaves on the
Stenchaetothrips biformis Very common Primarily rice, maize, Bangladesh, Cambodia,
(Bagnall) sugarcane; other India, Indonesia, Japan, upper parts of the plant.
graminaceous crops Malaysia, Myanmar, Female thrips lay an average of 93
and weeds Philippines, Sri Lanka, and a maximum of 147 eggs at 25 °C.
Thailand, Vietnam
Anaphothrips obscurus Less common Tobacco, rice, other Japan, Europe The eggs of S. biformis are tiny,
(Muller) graminaceous plants usually 0.25 mm long by 0.1 mm
Chirothrips manicatus Less common Polyphagous: oat, Japan, USSR, Europe,
wide, and pale yellow. They are laid
(Haliday) barley, wheat North America
Frankliniella intonsa Less common Clover, alfalfa, flowers Taiwan, China; Japan; singly in slits cut in the leaf blade
(Trybom) of graminaceous and Europe tissue by the saw-like ovipositors of
other plants
Frankliniella tenuicornis Less common Flowers of grami- Germany, England,
the female. The upper half of the egg
(Uzel) naceous plants, e.g, Finland, Japan, is exposed on the leaf surface. The
wheat and rye Scandinavia, Sweden incubation period is about 3 d. The
Aelothrips fasciatus a Less common Polyphagous Northeastern China, India,
(Linnaeus)
optimum temperature for incubation
Japan, Republic of Korea,
USA, northern USSR, is 25 to 30 °C, and for nymphs, 26.5 °C.
Europe The freshly hatched larvae are
a Status as a rice pest doubtful; the species has been recorded to feed on other thrips, aphids, mites, and various colorless but turn pale yellow in the
other small insects. second instar. First- and second-
47
instar larvae feed actively on the soft
tissue of the unopened young leaves
in spring and multiply on these
weeds or on other graminaceous
Damage
or within leaves rolled by thrip crops such as wheat, barley, and oats. Larvae and adults have rasping type
adults. The entire larval, prepupal, From these crops they move to rice mouthparts and only one mandible,
and pupal periods are completed at nurseries or transplanted fields. the left one. Leaf-feeding thrips
these sites. Most of the time, mobile Infestation in the nursery is fre- species exhibit a punch-and-suck
adults are also found hidden within quently carried over to transplanted feeding technique: the single man-
the rolled leaves; in some instances fields. Occasionally, overwintering dible punches a hole in the plant
both nymphs and adults may be H. aculeatus adults have been found surface through which the paired
found feeding on the outer surfaces in stored unhulled rice. With multir- maxillary stylets are then inserted to
of these areas. The duration of the ice cropping in the tropics, thrips imbibe the plant sap. Typical symp-
various life stages is greatly influ- have been reported to be more toms of thrip damage include inward
enced by existing environmental abundant in ricefields from July to rolling of the leaves along the mar-
conditions, especially temperature. September and in January. These gins, wilting, and stunting (Fig. 16b).
Duration from the newly hatched periods coincide with the seedling There are also fine, yellowish or
nymph to the adult was 9 d at 23.3 ºC stage of the rice crop. silvery streaks developing from the
and 4 d at 36 ºC. Both S. biformis and H. aculeutus margin to the midrib. In severe
have been recorded throughout crop infestation, seedlings may die result-
growth but are more abundant ing in a low number of plants per
Seasonal occurrence during the seedling and flowering unit area (Fig. 16c). The damage is
and abundance stages. Thrips are frequently reported most evident if there is no standing
water in the ricefields.
as more serious during dry periods;
Despite their small size and fragile this could be due to a reduced toler-
appearance, thrips can travel long ance of rice plants under drought
distances. They migrate during the stress. Control methods
day and seek out newly planted In India, thrips infestation is
ricefields. They are day-flying insects generally recorded during the first Cultural control
and are not attracted to light traps. In week of August, with peak infesta- Flooding the field to submerge plants
areas of cool winters, S. biformis tion during the second and third for 2 d effectively controls thrips.
adults migrate to graminaceous weeks. The population declines in
weeds after the flowering of rice and early September and disappears by
hibernate. They become active again mid-September.
16. Rice thrips: a) adult,
b) damaged leaf,
c) damaged plants in a
field.
Rice thrips 49
Rice caseworm
The rice caseworm Nymphula which occurs in China, India, Indone- Life history
depunctalis (Guenée) (Pyralidae- sia, Japan, Philippines, Sri Lanka, and
Lepidoptera) is an important insect Thailand. N. depunctalis occurs The adult moths are snowy white,
pest of rice. It occurs in Australia, regularly in low populations, except about 6 mm long, and have a wing
many tropical countries (India, for some occasional buildup in small expanse of about 15 mm. The white
Indonesia, Malaysia, Philippines, Sri areas where it may severely defoliate wings are marked with a few light
Lanka), and in Africa, and South the plants. Its common name, case- brown to black specks and two or
America. N. depunctalis was previ- worm, refers to the larval habit of three submarginal fulvous bands
ously known as N. stagnalis (Zeller) forming the leaves of rice plants or (Fig. 17a). The adults are nocturnal,
1852, Hydrocampa depunctalis grasses into tubes or cases and hiding in ricefields during the day
(Guenee) 1854, and Zebronia decussalis enclosing itself within them during and laying eggs at night. Females
(Walker) 1954. Among related species feeding. The leaf cases protect the oviposit an average of 2.5 d after
are Nymphula vittalis Bremer and larvae from natural enemies and act emergence, usually starting within
Nymphula fengwhanalis (Pryer), which as floats to carry the larvae from one 1-4 d. Eggs, 10-20 in a batch, are laid
occur in China; Paraponyx plant to another. Rice at the seedling in one or two adjacent rows on the
(=Nymphula) fluctuosalis Zeller, which and vegetative stages is its preferred undersides of leaves floating on
occurs in some African countries and host. It also infests millet and various water (Fig. 1%). A single female lays
in Australia, China, India, Japan, grasses such as Panicum, Eragrostis, an average of about 50 eggs in her
Malaysia, Philippines, Sri Lanka, and and Paspalurn. lifetime. Mated females die 1 d after
Thailand; and P. diminutalis (Snellen),
51
oviposition; unmated females can live
up to 1 wk. The egg is light yellow,
Seasonal occurrence Control methods
circular, about 0.5 mm in diameter, and abundance Cultural control
and somewhat flattened with a Cultural methods involving water
smooth surface. Before it hatches, the The caseworm occurs only in management are effective in control-
egg turns darker with two purplish ricefields with standing water. It is ling the rice caseworm larvae. A
dots, which represent the eyes of the found in irrigated and rainfed low- nonflooded seedbed is protected
developing larva. Average incubation land environments and is more from caseworm attack. Draining the
period is 4 d. prevalent in the rainy season. The field for at least 3 d will kill most of
The freshly hatched larva is pale pest is usually abundant July to the larvae because they are highly
cream, about 1.2 mm long and November, when it infests rice plants dependent on water for oxygen.
0.2 mm across the head. The head is and other grasses in the ricefields. However, this practice favors weed
light yellow. The second instar larva After November, it migrates to grassy growth. Transplanting older seed-
is somewhat greenish. Five larval areas. No hibernation is recorded in lings may also help in limiting the
instars are completed in an average of the Philippines and the insect occurs period of caseworm larvae attack.
20 d. The full-grown larva is about in overlapping generations. Irriga-
14 mm long and 1.6 mm in diameter. tion, which ensures prolonged Biological control
It is pale green with a semitranspar- standing water in the vegetative Trichogramma minutum Ashmead is
ent skin and light brown prothoracic stage, increases the pest’s abundance. reported as a parasite of caseworm
shield and head. A characteristic eggs. While foraging for algae, snails
feature that becomes visible only at such as Pila sp. and Radix sp.
the start of the second instar is six Damage (Lymnaeidae) may dislodge case-
rows of gills with tubes connected to Larvae begin to feed shortly after worm eggs from rice leaves. A
the main trachea (Fig. 17c). The five hatching and start making leaf cases braconid wasp Dacnusa sp. and a
pairs of spiracles on the larva’s body 2 d later. The freshly hatched larvae tabanid fly Tabanus sp. parasitize the
are considered nonfunctional and the feed on the surface of the tender caseworm larvae. The larvae of
larva breathes through its gills. leaves, but later instars feed from hydrophilids [e.g., Hydrophilus affinis
During the day, larvae float on the within the case or on the surface of (Sharp), Sternolophus rufipes
water; at night they climb plants to even the older leaves. Damage is (Fabricius), and Berosus sp.] are
cut off leaves to make new cases, or caused by larvae feeding and cutting reported predators of the caseworm
feed on severed leaves on the water off the leaf tips for making leaf cases. larvae, as are dystiscids Laccophilus
surface. In making a case, the larva Damage is characterized by difficilis (Sharp) and Cybister
moves to the tip of a young leaf, cuts ladder-like appearance of the re- tripunctatus orientalis Gschwendther.
the leaf blade on one side of the moved leaf tissue, leaving the upper The common red ants Solenopsis
midrib, then secretes a silken thread epidermis somewhat papery. Dam- geminata (Fabricius) attack the larvae
to bind the two margins of the leaf aged plants occur in patches in the and pupae of the pest, especially
blade into a tubular structure. The field. Heavily infested plants may when the infested ricefields become
larva detaches this tube from the leaf still recover in 1 mo, but are already dry. Several spiders prey on the
by severing the other part of the leaf stunted. They produce fewer tillers moth. They include three species of
blade. The larva replaces these leaf and smaller panicles, and maturity is Araneidae—Neoscona theisi
casings with new ones after each molt delayed. Yield loss occurs if, during (Walckenaer), Argiope catenulata
and undergoes pupation within the the first 30 d after transplanting, (Doleschall), and Araneus inustus
leaf case. other pests such as whorl maggot or (L. Koch); one species of
The pupa, about 5.5 mm long and stem borer infest the crop, reducing Oxyopidae—Oxyopes javanus
1.5 mm wide, is cream colored when the plants’ ability to recover. (Thorell); one species of Lycosidae
freshly formed, but turns silvery Pardosa pseudoannulata (Boesenberg
white toward moth emergence and Strand); one species of
(Fig. 17d). The pupal period lasts for Tetragnathidae—Tetragnatha nitens
about 1 wk. The adult usually (Audouin); and one species of
emerges during the night through an Clubionidae—Clubiona japonicola
opening at the upper end of the case. Boesenberg and Strand.
The life cycle is completed in 1 mo.
Rice caseworm 53
Rice mealybugs
55
Damage pygrnaeus Lamb and Mepachymerus Prakasa Rao P S (1989) Bioecology and
management of insect pests of rainfed
ensifer (Thompson)] and one droso-
Nymphs and adult females infesting philid [Gitona perspicax (Knab)]. upland rice ecosystem. Nutr. Ecol. Inst.
the leaf sheath damage the plant by Environ. 93-106.
sucking sap from the rice stem. That Varietal resistance Prakasa Rao P S, Das A P K (1971) New
results in smaller leaves, yellowing, No varieties resistant to B. rehi are records of some natural enemies of the
abnormal tillering, and stunted commercially available. rice mealy bug Ripersia oryzae Green
plants. Under heavy infestation, (Hemiptera, Pseudococcidae). Oryza
81111-112.
either no panicles are formed or they Chemical control
do not fully exsert from the boot; the The waxy secretion covering the Reissig W H, Heinrichs E A, Litsinger J A,
Moody K, Fiedler L, Mew T W, Barrion
plants may even dry off. The damage mealybugs and their habit of living
A T (1986) Illustrated guide to inte-
is in patches since the young nymphs behind leaf sheaths protect them
grated pest management in rice in
have rather limited migrating ability. from insecticide. Foliar sprays are tropical Asia. International Rice
Likewise, mealybug numbers vary effective, however, if the nozzle is Research Institute, P.O. Box 933,
greatly between hills. This causes the directed to the bases of plants. Granu- Manila, Philippines. 411 p.
field to have several spots of de- lar insecticides are effective in fields Williams D J (1970) The mealybugs
pressed growth, which are known by with standing water. If damaged (Homoptera, Coccoidea, Pseudococ-
various names such as chakdhora, fields have no standing water, broad- cidae) of sugarcane, rice, and sorghum.
soorai disease, etc. Damage is intense casting granules is impractical. Bull. Entomol. Res. 60:109-188.
during drought conditions when rice Williams D J, Watson G W (1988) The
plants can least tolerate sap loss. scale insects of the tropical South
Selected references Pacific region. Part 2. The mealybugs
Alam M Z (1967) Insect pests of rice in (Pseudococcidae). CAB International
Control methods East Pakistan. Pages 643-655 in The Institute of Entomology, London, UK.
major insect pests of the rice plant. 260 p.
Cultural control Proceedings of a symposium at the
International Rice Research Institute,
Cultural control measures include
September 1964. The Johns Hopkins
timing of planting dates to escape
Press, Baltimore, Maryland.
peak infestation, continuous flooding
Alam S, Alam M S, Rezaul Karim A N M
of the field at 5-cm depth throughout
(1979) Rice mealybug outbreak in
the crop growth period, and removal
Bangladesh. Int. Rice Res. Newsl.
and destruction of infested plants at 4(5):20-21.
the first sign of mealybug damage.
Basu A C, Banerjee S N (1965) The control
of Ripersia oryzae Green, a mealy bug of
Biological control the paddy plant in West Bengal. J.
Lady beetles such as Coccinella Econ. Entomol. 58:621-623.
repanda Thunberg, Menochilus Gopalan M, Coumararadja N,
sexmaculatus Fabricius, and Harmonia
Balasubramanian G (1987) Effect of
octomaculata (Fabricius) are the main
different planting dates and irrigation
natural enemies of the mealybug. regimes on the incidence of rice mealy
Hymenopterous parasites of B. rehi bug, Brevennia rehi Lindinger. Madras
recorded from India include Ceraph- Agric. J. 74:226-227.
ronidae (Cerapkron sp.), Encyrtidae Manjunath T M (1968) New records of
(Adelencyrtus sp., Cheiloneurus sp., some parasites and predators of the
Doliphocerus sp., Gyranusa sp., rice mealy bug, Ripersia oryzae Green
Mayeridia sp., Parasyrphophagus sp., (Hemiptera: Pseudococcidae). Curr.
and Xanthoencyrtus sp.), Eulophidae Sci. 37:354-356.
(Aprostocetus sp., Chrysocharis sp., Pai P S, Mahabala Gowda T (1981)
Desostenus sp., and Tetrastichus sp.), Preliminary studies on the control of
Mymaridae (Lymaemon sp.), Pter- the rice mealy bug Heterococcus rehi
omalidae (Callitula sp. and Diparini (Ldgr.) (Hemiptera: Pseudococcidae).
sp.), and Thysanidae (Thysanus sp.). Curr. Res. 10:88-89.
Recorded dipterous predators of
B. rehi are two chloropids [Anatrichus
Whorl maggots constitute a complex vegetation. During the early and later
of several species belonging to the parts of the day, they float on the
genus Hydrellia in the family Ephy- water or perch on floating vegetation.
dridae and order Diptera. All mem- During midday, they are sedentary,
bers of the genus are stem or leaf clinging on upright vegetation. The
miners. They prefer an aquatic average egg-laying capacity is 20
habitat, usually living near water or eggs/female. The egg-laying period
in damp areas. Rice whorl maggots lasts for 1-8 d.
are difficult to identify because they The eggs are laid singly on either
resemble other flies in the field such surface of the leaves. They are cylin-
as Psilopa and Paralimna whose larvae drical, whitish, and are 0.65-0.85 mm
feed on rice, and Notiphila spp. that long and 0.15-0.20 mm wide. When
live on decomposing organic matter freshly laid, the entire egg yolk mass
in ricefields (Fig. 19a, b). Three is opaque, but later the anterior half
important species of Hydrellia are shows the black cephalopharyngeal
known to infest rice plants. skeleton of the developing embryo.
The incubation period is 2-6 d.
The freshly hatched larvae are 19. Rice whorl maggot: a) Hydrellia
transparent to light cream colored, philippina adult, b) Notiphila sp. adult.
57
Damage suffer thermal mortality above 40 °C. Pupation occurs either within the
Mating starts 3 d after emergence, original mines or, more frequently,
The fly maggots feed on the inner and within the temperature range of the full-grown larvae form a new
margins of unfurled leaves. The 13-37 °C, emergence and mating mine for pupation. The pupae are
newly hatched larvae migrate to the occur at any hour of the day. Both visible within the translucent mines.
central whorl and feed on the meso- males and females multimate. The Pupae from soils have also been
phyll tissue. preoviposition period is about 5 d. recorded.
Damage is characterized by small, Optimum temperature for oviposi-
chewed-up, discolored areas on the tion is 10-32 °C when a single female
innermost margin of the central lays about 50-100 eggs. High winds Seasonal occurrence
whorl. These areas eventually dry up
and damaged leaves usually droop.
and abundant algal growth cause
several of the lower leaves to remain
and abundance
Heavy infestation causes a marked on the water surface; these leaves are In California, the adult flies occur
stunting of the plant, fewer tillers, preferred for oviposition and are throughout the year, but their popu-
and delayed panicle initiation and suitable for larval growth. Eggs are lation is usually highest from March
maturity. laid singly on the leaves, but a single to June. No oviposition has been
leaf may contain several eggs. Ovi- recorded from November to Febru-
position usually occurs on horizontal ary. The pest occurs in 11 genera-
leaves near the water surface, but tions, and beyond the first generation
Rice leaf miner with dense foliage and high RH, eggs
have been recorded at 15-20 cm
the broods usually overlap. In north-
ern Japan, eight generations have
above water level. The eggs hatch been recorded. The first generation
Hydrellia griseola (Fallén) is not a true between 6 and 32 °C and above 50% occurs in the second half of April.
whorl maggot but a leaf miner. It is RH; hatching is highest at 98% RH. Adults of the second generation
an important pest in California, USA, The average incubation period of oviposit on rice leaves in May. The
and northern Japan. It has also been about 5 d between 21 and 32 °C is flies overwinter in the pupal stage.
recorded in northern USA, Europe considerably prolonged at higher Optimum temperature, adequate
(France, Italy, and Germany), and temperature. humidity, and availability of host
South America. This species is pri- The freshly hatched larvae are plants regulate fly populations.
marily a pest of rice, but is also tiny, usually 0.10-0.17 mm wide and Temperatures below 0 °C for a short
known to infest barley, oat, timothy, 0.33-1.13 mm long, and nearly trans- duration cause no harm. The insect's
and wheat, in addition to aquatic parent to light cream colored. They activity increases from the beginning
plants. It is also referred to by various have well-developed mouth hooks, of spring because of warmer tem-
common names such as rice leaf which they use to mine inside the leaf perature and availability of standing
miner, smaller rice leaf miner, gray tissue. The larvae start mining soon water.
barley fly, and gray mining fly. after hatching, either with their Low temperatures in the preceding
caudal segments still within the egg summer provide favorable conditions
Life history shell or after a short migration from for insect survival and multiplication,
the oviposition sites. Mining usually and result in an increased fly popula-
The adult fly, about 2 mm long and lasts for 2-3 h. Under limited food tion in the fall. Higher temperatures
0.7-0.8 mm wide, is light grey and supply, crowding, or submergence of than usual during winter, accompa-
looks like a small housefly. It usually the affected leaves, the larvae may nied by an early thaw, cause low
lives for 3-4 mo and prefers high- leave the mines and migrate to newer mortality of the overwintering
moisture areas. The flies are hydro- leaves. They can crawl on a sub- population. These conditions, aug-
fuge, i.e., they can float on the water merged surface; later instar larvae mented by synchronism of the
surface if submerged and also walk usually take 0.5-1.5 h to bore into the oviposition period with transplant-
on the water surface. It has been leaves. The maggots undergo three ing, deep water in the fields, and
recorded feeding on dead organisms. larval instars. The total larval period slow growth of the plants because of
The adults are positively pho- is 7-10 d at 21-32 °C, but is considera- low temperature, increase the dam-
totropic. They exhibit optimum bly prolonged at lower temperature. age. A combination of these factors
activity at 11-31 °C, and are fairly The entire larval and pupal periods resulted in a severe outbreak of the
tolerant of cold weather (showing are completed within the mines. fly in northern Japan in 1954.
some activity even at -2 °C), but
Whorl maggots 59
(Boesenberg and Strand), and araneid Selected references Reissig W H, Heinrichs E A, Litsinger J A,
Moody K, Fiedler L, Mew T W, Barrion
Neoscona theisi (Walckenaer). Fungus
Ferino M P (1968) The biology and control A T (1986) Illustrated guide to inte-
of the genus Entomophthora is also of the rice leaf whorl maggot, Hydrellia
recorded as parasite of whorl maggot. grated pest management in rice in
philippina Ferino (Ephydridae, tropical Asia. International Rice
Diptera). Philipp. Agric. 52:332-383. Research Institute, P.O. Box 933,
Varietal resistance Ferino M P, Saavedra N T, Magallona E D Manila, Philippines. 43 1 p.
Only one rice cultivar (IR40) and two (1983) Major insect pests of rice and Rubia E G, Shepard B M (1987) Biology of
wild rices (Oryza brachyantha and their control. Pages 232-265 in Rice Metioche vittaticolis (Stål) (Orthoptera:
O. ridleyi) have been identified as re- production manual, Philippines. Gryllidae), a predator of rice pests.
sistant to whorl maggots. University of the Philippines at Los Bull. Entomol. Res. 77:669-676.
Baños, College, Laguna.
Tomioka T (1983) Ecological note on two
Chemical control Grigarick A A (1959) Bionomics of the rice species of Hydrellia (Diptera, Ephydri-
Broadcasting of nonsystemic granules leaf miner, Hydrellia griseola (Fallen) in dae). Hokkaido Cent. Agric. Exp. Stn.
on standing water in the field, or soil California (Diptera: Ephydridae). Annu. Rep. Soc. Plant Prot. North Jpn,
incorporation of systemic granules Hilgardia 29:1-80. 34:13-16.
during the last harrowing before Heinrichs E A, Medrano F G, Rapusas H Viajante V D (1982) Yield assessment of
transplanting is usually more effec- R (1985) Genetic evaluation for insect rice damaged by the whorl maggot,
tive than foliar sprays. Foliar sprays resistance in rice. International Rice Hydrellia sasakii Yausa and Isitani.
can, however, be applied 1 or 2 wk Research Institute, P.O. Box 933, Unpublished MS thesis, Gregorio
Manila, Philippines. 356 p.
after transplanting. Araneta University Foundation,
Kuwayama S (1958) The smaller rice leaf Malabon, Metro Manila, Philippines.
miner, Hydrellia griseola Fallen in 130 p.
Japan. Pages 17-25 in Proceedings of Viajante V D, Heinrichs E A (1985a)
the Xth International Congress of Oviposition of rice whorl maggot as
Entomology, August 1956. Montreal, influenced by azolla. Int. Rice Res.
Canada. Newsl. 10(1):23.
Meneses R, Cordero V (1989) Response of Viajante V D, Heinrichs E A (1985b)
lines and varieties of rice to Hydrellia Influence of water regime on rice
sp. under field conditions. Rice Caribb. whorl maggot oviposition. Int. Rice
3:1-5. Res. Newsl. 10(1):24-25.
Natarajan K, Mathur K C (1980) New
record of Chaenusa sp. (Braconidae -
Hymenoptera) as a parasite of the rice
whorl maggot. Sci. Cult. 46:337-338.
Nurullah C M (1979) Component analysis
for the integrated control of the rice
whorl maggot, Hydrellia sasakii Yuasa
and Isitani. Unpublished MS thesis,
University of the Philippines at LOS
Baiios, Philippines. 120 p.
Okamoto D, Koshihara T (1962) Studies
on paddy stem maggot, a pest of the
late transplanted rice plants. Bull.
Chugoku Agric. Exp. Stn. A(8):235-267.
Patanakamjorn S (1964) Morphology and
bionomics of rice leaf whorl maggot.
International Rice Research Institute,
P.O. Box 933, Manila, Philippines. 11 p.
Philippine-German Crop Protection
Programme (1986) Integrated pest
management rice pocket reference
manual. Bureau of Plant Industry,
Ministry of Agriculture and Food,
Manila, Philippines. 137 p.
The ladybird beetles (Coleoptera: Table 8. Ladybird beetles recorded feeding on rice.
Coccinellidae) are brightly colored, Name Feeding on Distribution
small, oval, convex insects, and
Coccinella repanda Rice pollen, Australia; Bangladesh; Bhutan; Taiwan, China;
comprise about 5,000 species. The
(Thunberg) panicle India; Melanesia, Micronesia; Nepal; New
greater number of these species are [ = Coccinella transversalis Zealand; Sri Lanka; Tasmania; Thailand
predaceous, both as larvae and Fabricius]
adults, and feed on a wide variety of Exochomus nigromaculatus Leaf Malaysia
(Goeze)
soft-bodied insects such as leafhop- Micraspis afflicta Panicle Indonesia, Malaysia
pers, planthoppers, mealybugs, and (Mulsant)
aphids, and on the eggs of various Micraspis crocea Rice pollen, Philippines
(Mulsant) leaf, panicle
insects. A comparatively small Micraspis discolor Rice pollen, South and Southeast Asia, China, Japan
number of species are of economic (Fabricius) leaf (including Ryukyu Islands)
significance as plant pests. Most Micraspis lineata Indonesia
Micraspis vincta Rice pollen India, Malaysia, Myanmar, Pakistan, Thailand
predaceous species can be broadly (Gorham)
distinguished by their mandibles,
which have simple or bifid apices,
each with a basal tooth, lacking in
most phytophagous species. In
phytophagous insects, the apices of presence of hosts, and are numerous
the mandibles are multidentate. at the bases of plants and on the
Some ladybird beetles recorded as upper leaves and panicles. After
feeding on rice plants are listed in harvest, they congregate in the
Table 8. The most common and stubble. Adults live 25 to 40 d.
widely distributed are Micraspis
discolor (Fabricius) and M. crocea Eggs
(Mulsant). The preoviposition period is 6-10 d.
Females lay 14-26 eggs/d in a cluster
on the upper and lower leaf surfaces.
Life history Eggs within a cluster are not attached 20. Ladybird beetle Micraspis discolor.
to each other. The egg is about 0.99-
Adults 1.05 mm long and 0.41-0.44 mm wide,
Beetles of M. discolor are about 3.2- elongate-oval, and shiny yellow- day. Larvae are soft-bodied,
3.8 mm long and 2.8-3.2 mm wide orange, which gradually changes to brownish black, elongate, somewhat
(Fig. 20). They have oval convex pale brown and later to brown. flattened, and covered with minute
bodies with bright orange to red- Reddish eye spots develop just before tubercles or spines. They have the
orange elytra. The males are usually hatching. Egg incubation is 2-4 d. general body shape of an alligator.
slightly smaller than the females. The They are very active and are seen
beetles are cannibalistic, feeding on Larvae moving about the whole plant,
their own eggs, larvae, and pupae, The hatching larva cuts an irregular preying on various insects or feeding
but usually prefer soft-bodied insects. hole through the upper end of the on the exposed pollen. They undergo
They are diurnal and live on various egg shell. Eggs from a single batch four larval instars in an average of 15-
plant parts as predators. They select usually hatch simultaneously and the 20 d. The full-grown larva is 5-6 mm
their feeding location according to the larvae remain clustered for almost a long. It becomes sedentary before
61
pupation, attaching itself to the rice
Control method Selected references
plant by means of a suction disc
Agarwala B K, Das S, Senchowdhuri M
located at the abdominal tip. Micraspis spp. can be readily con- (1988) Biology and food relations of
trolled by organophosphate insecti- Micraspis discolor F. on aphidophagous
Pupae cides used as sprays. However, since coccinellid in India. J. Aphidol. 2:7-17.
The newly formed pupa is pinkish, these beetles are predators of numer- Patnaik N C (1983) Toxicity of some or-
but later turns orange to reddish ous harmful insects, controlling the ganophosphorus insecticides to
orange. The pupation period is 3-8 d. prey population helps to control the predatory coccinellids of rice fields.
beetles. Oryza 20:237-239.
Samal P, Misra B C (1985) Morphology
Seasonal occurrence and biology of the coccinellid beetle
and abundance Verania discolor Fab. (Coccinellidae:
Coleoptera), a predator on rice brown
Occurrence of the beetles depends on planthopper Nilaparvata lugens (Stål).
prey. At IRRI, studies on the life cycle Oryza 22:58-60.
and occurrence of the beetles re- Shepard B M, Rapusas H R (1989) Life
vealed that M. crocea is most abun- cycle of Micraspis sp. on brown
dant during rice flowering and planthopper (BPH) and rice pollen. Int.
during outbreaks of its prey Rice Res. Newsl. 14(3):40.
Nilaparvata lugens (Stål). In India, Yunus A (1967) Insect pests of Malaya.
adults of M. discolor occur throughout Pages 617-633 in The major insect pests
the crop year in aphid-infested fields of the rice plant. Proceedings of a
symposium at the International Rice
and then disperse to weeds. By the
Research Institute, September 1964.
end of March, the beetles undergo
The Johns Hopkins Press, Baltimore,
reproductive diapause, which ends Maryland.
within 6-8 d of aphids' appearance on
the crop.
Temperatures of 16-26 °C and RH
of 60-80% favor faster larval
development.
Damage
Ladybird beetle pests of rice have
mixed feeding habits. Adults and
nymphs prefer to prey upon various
aphid, leafhopper, and planthopper
nymphs and adults; stem borer eggs;
and other soft-bodied, small insects
such as thrips and mealybugs. In the
absence of prey, however, they feed
on leaf blades (leaving small chewed
areas), pollen, and frequently damage
developing grains. They gnaw a hole
through the rice hull to feed on the
soft grain during the dough stage.
The two most common species of Black bugs are known to attack the to light, but not to yellow electric
black bugs (Hemiptera: Pentatomi- rice crop at various growth stages bulbs. Their flight activity to light
dae) attacking rice plants are the from seedling to maturity. Heavy traps coincides with the lunar cycle.
Malayan rice black bug Scotinophara damage is usually seen after heading Adult females lay their eggs in
coarctata (Fabricius) and the Japanese or maturing, especially when irriga- clusters, each containing 29-34 eggs,
rice black bug Scotinophara lurida tion is stopped during the maturation on the basal parts of the rice plants
(Burmeister). They are also com- period. Large populations of the pest near the water surface. The egg is
monly known as rice pentatomid are more common on lowland than about 1.0 mm long and 0.65 mm
bugs. Scotinophara coarctata is an on upland rices, and on continuously wide; shiny, pale greenish grey;
important pest of rice in Cambodia, irrigated or continuously rice- cylindrical; and finely reticulated.
China (including Taiwan), India, cropped fields than on rainfed or The top is greyish white-indicating
Indonesia, Malaysia, Philippines, single-cropped fields. They are also the cap, which splits to allow the
Thailand, and Vietnam. Scotinophara more common in direct-seeded nymph to emerge. The female guards
lurida occurs in China (including paddies than in transplanted fields the eggs by staying on top of the
Taiwan), India, Japan, and Sri Lanka. because of their preference for mass. Average incubation period is
The occurrence of Scotinophara densely planted fields. 4-7 d. There are five nymphal instars,
coarctata on rice was first recorded in Aside from rice, other hosts are Zea which last for 26-34 d. Adult bugs
Indonesia in 1903. In West Malaysia mays (L.), Alocasia indica (Roxb.) survive for 4-7 mo.
the pest was first recorded on rice in Schott, Colocasia esculenta (L.) Schott,
1918. Since 1983 its numbers have Lecrsia hexandra (L.) Sw., Echinochloa Seasonal occurrence
increased because of the staggered crus-galli (L.) P. Beauv., Rottboellia
planting of the rice crop. In the cochinchinensis (Lour.) W. D. Clayton, and abundance
Philippines, the pest was first re- Echinochloa colona (L.) Link.,
corded in September 1979 in southern Fimbristylis miliacea (L.) Vahl, Cyperus Eggs hatch and nymphs develop
Palawan; it later spread to central and rotundus L., Cyeperus iria L., Sacciolepis optimally at temperatures of 25-28 °C
northern Palawan. The black bug lnyurus (Lam.) A. Chase, Scirpus and 75% RH.
problem in Palawan is partly related grossus L.f., Scleria sumatrensis Retz., In Sri Lanka, Scotinophara lurida
to double cropping of irrigated rice and Hymenachne acutigluma (Steud.) has three or four overlapping genera-
since the introduction of high-yield- Gilliland. tions in a year. The first generation
ing cultivars and high N levels. Black and part of the second generation
bug infestation is most serious in occur on the first crop; the remainder
poorly drained ricefields around Life history of the second, the third, and part of
marshes. Except for Palawan, no the fourth generation occur on the
other parts of the Philippines have The adult of Scotinophara coarctata has second crop. There are two overlap-
reported severe black bug infestation. a black head, collar, and cicatrices; ping generations in aestivation at
Scotinophara lurida is a major pest yellowish brown antennae; reddish to each period between crops. At
of rice in China and Japan. Before dark brown abdomen; and reddish aestivation sites, fourth- and fifth-
1940 the pest was recorded as an brown legs with yellowish tibiae and instar nymphs and adults are most
occasional rice pest in Sri Lanka. tarsi. Average size is 9 mm from the commonly found.
Since then it has become economi- anterior margin of the head to the In Japan, Scotinophara lurida devel-
cally important, occurring periodi- apex of the abdomen and 4.5 mm ops only one generation a year. The
cally in large numbers and causing across the prothorax. Black bugs are first generation bugs that develop on
extensive damage to rice crops in regarded as noisome insects with a the rice crop hibernate in the winter
most parts of the country. bad smell. They are strongly attracted in adult stage and invade the next
year's crop in the seedling stage.
63
Damage the carabid predator Agonum daimio Hirao J, Ho N K (1987) Status of rice pests
Bates; the nabid bug Stenonabis and measures of control in the double
Black bug nymphs and adults are tagalicus (Stål); the spider predators cropping area of the Muda Irrigation
usually found at the basal part of the Pardosa pseudoannulata (Boesenberg Scheme, Malaysia. Trop. Agric. Res.
rice plant and damage is caused by and Strand), Oxyopes javanus Ser. 20: 107-115.
their feeding on the plant sap. At the (Thorell), Tetragnatha virescens Kiritani K (1986) Changes in rice pest
early hours of the day, the bugs are Okuma, and Camaricus formosus status with cultivation systems in
found feeding on the upper parts of Thorell; and the entomopathogenic Japan. Pages 1-15 in Seminar on Rice
the rice seedlings and on the upper Insect Pest Control, 18 September 1986.
fungi (Deuteromycotina) Beauveria
Tsukuba, Japan.
surfaces of the leaves. Later, when bassiana (Balsamo) Vuillemin,
sunlight intensity increases, they Metarhizium anisopliae (Metschnikoff) Miyamoto S, Torres N A, Habibuddin
move to the undersides of the leaves Sorokin, and Paecilomyces lilacinus B H, Montri R, Fujimura T, Thieng P,
and stems and continue to feed there. (Thompson) Samson. Mochida O (1983) Emerging problems
They are very sluggish in the day, of pests and diseases—the black bug in
rarely taking wing. At night they Southeast Asia. Paper presented at the
Varietal resistance
International Rice Research Confer-
become very active, are often seen in Two rice cultivars, IR10781-75-3-2-2
ence, 18-22 April 1983. International
flight, and feed continuously and IR13149-71-3-2, are resistant to Rice Research Institute, Los Baños,
throughout the night. Older nymphs rice black bugs. Laguna, Philippines. 33 p.
and adults are sometimes found on
Mochida O, de Sagun S, Parducho M,
cracks in the soil during the day. Chemical control Gapasin D, Orolaza N P (1986) General
Heavy infestation during the Directly spraying insecticides at the information on black bugs, insecticide
tillering stage kills the central shoots, base of the plants where the black evaluation, economic injury levels, and
resulting in deadheart. Rice plants bugs stay provides the most effective principles of chemical control. Paper
show slight stunting, yellowing, control. presented at the PCARRD/MAF/IRRI
chlorotic lesions, and fewer tillers. Workshop on Black Bug, 25-29 October
During the booting stage, plants 1986. Palawan, Philippines. pp. 1-17,
show stunted panicles, no panicles, or Selected references Perez V A, Shepard B M, Arida G S (1989)
incompletely exserted panicles, and Arida G S, Shepard B M, Perez V A (1988) Indigenous natural enemies of the
unfilled spikelets or whiteheads. Parasitization of the Malayan black Malayan black bug Scotinophara
During crop maturation, incomplete bug (MBB) by five species of egg para- coarctata (Fab.) on Palawan Island,
and unfilled spikelets increase. sitoids. Int. Rice Res. Newsl. 13(6):42. Philippines. Philipp. Entomol. 7:485-
Barrion A T, Mochida O, Litsinger J A, 490.
Heavy damage at any stage causes
the plants to wilt and dry, a condition de la Cruz N (1982) The Malayan black Reissig W H, Heinrichs E A, Litsinger J A,
known as bug burn. bug Scotinophara coarctata (F.) [Hem- Moody K, Fiedler L, Mew T W, Barrion
iptera: Pentatomidae]: a new rice pest A T (1986) Illustrated guide to inte-
in the Philippines. Int. Rice Res. grated pest management in rice in
Control methods Newsl. 7(6):6-7. tropical Asia. International Rice
Research Institute, P.O. Box 933,
de Sagun S B (1986) Insecticide screening Manila, Philippines. 411 p.
Cultural control for black bug, Scotinophara coarctata
Since black bugs remain in the rice (Fabricius) (Heteroptera: Pentatomi- Rombach M C, Aguda R M, Shepard B M,
stubble after harvest, plowing the dae) control in Palawan, Philippines. Roberts D W (1986) Entomopathogenic
field helps to kill the pests and MS thesis, Gregorio Araneta Univer- fungi (Deuteromycotina) in the control
of the black bug of rice, Scotinophara
destroy their host plants. Removing sity Foundation, Malabon, Metro
Manila. 59 p. coarctata (Hemiptera: Pentatomidae). J.
weeds from the field to allow more
Inverteb. Pathol. 48:174-179.
sunlight to reach the bases of the rice Fernando H E (1960) A biological and
Shepard B M, Perez V A (1987) Influence
plants is another cultural control ecological study of the rice pentatomid
bug, Scotinophara lurida (Burm.) in of cultivation on survival of the
method.
Ceylon. Bull. Entomol. Res. 51:559-576. Malayan black bug in ricefields. Int.
Rice Res. Newsl. 12(3):35.
Biological control Heinrichs E A, Domingo I T, Castillo E H
(1987) Resistance and yield responses Subramanian A, Murugesan S, Rajendran
Natural enemies of the rice black bug
of rice cultivars to the black bug R, Babu P C S (1986) Occurrence and
include the scelionid egg parasitoid control of rice black bug at Coimba-
Telenomus triptus Nixon; the gryllid Scotinophara coarctata (F.) (Hemiptera:
Pentatomidae). J. Plant Prot. Tropics tore. Int. Rice Res. Newsl. 11(3):24.
predators Metioche vittaticollis (Stål)
4:55-64.
and Anaxipha sp.; the coccinellid
predator Micraspis crocea (Mulsant);
65
Life history
Adults
The adult moths are pale and brick-
red to pale brown, and have a very
hairy body covered with dark specks
and patches (Fig. 21a). They are about
2-3 cm long and have a 3-5 cm wing-
spread. The adult moths are noctur-
21. Armyworm and
nal, strongly phototropic, and are cutworm pests of rice:
quiescent during the day. The moths a) Mythimna separata
feed on dewdrops and other sugary adult, b) armyworm larva,
substances, and start mating 1-3 d c) Spodoptera mauritia
egg mass, d) S. litura
after emergence (DE). Mating and
adult, e) S. litura larva.
oviposition usually occur after dark
and during early evening hours. The
male moths live an average of 3 d; the
females live for 7 d. Generally, the
males die shortly after mating. The
females have an oviposition period of
about 5 d.
73
Control methods
Cultural control
Placing crop residues in the field at
planting time can divert the pest from
the growing crop.
Chemical control
Chemical control includes treating
seeds with insecticides at planting or
22. Soil-inhabiting pests
applying granular insecticide in the of rice: a) queens of
seed furrows or hills. The decision to termites, b) Gryllotalpa
use insecticides should be based on orientalis adult, c) adult of
the history of termite damage in a field cricket Plebeiogryllus
plebejus, d) adult of white
particular field.
grub, e) white grub larva,
f) root aphid.
Crickets
that gives them the common name of mole cricket populations.
Several species of mole crickets mole cricket. The insect uses these The mole cricket cannot live in
(Gryllotalpidae) and field crickets modified legs to burrow into the soil flooded fields, so they swim across
(Gryllidae) feed on the underground where it feeds on tender roots of the water to the levees. They burrow
parts of all upland crops including growing plants. Mole crickets gener- into the levees and lay eggs. When
rice (Fig. 22b, c). Four species of mole ally live in burrows 8-10 cm below the water level recedes, they migrate
crickets are reported to attack rice: the soil surface, but during unfavo- to the field to feed.
Gryllotalpa orientalis (=africana) rable weather they burrow deeper. Field cricket nymphs and adults
Burmeister in Asia, Gyllotalpa Adults aestivate or hibernate. They have similar nocturnal habits and
africana Palisot de Beauvois in Africa, are strong fliers. They are attracted to damage rice as much as do mole
and Neocurtilla hexadactyla (Perty) and light and are often recorded near crickets. Piles of weeds remaining on
Scapteriscus didactylus (Latreille) in street lights or in light traps. fields attract them. They also make
Latin America. Several species of The eggs are white, oval, and are subterranean nests and tunnel
field crickets are reported as rice laid in hardened cells or chambers through the soil to feed on roots.
pests: Gryllus assimilis (=bimaculatus) constructed in the soil by the females. Some species prefer seed to roots;
(Fabricius), Gryllus (=Liogyllus) One cell generally contains others feed at the base of stems.
bimaculatus de Geer, Brachytrupes 30-50 eggs. Depending on the tem- Like mole crickets, field crickets
portentosus (Lichtenstein), perature, the incubation period is cannot survive in standing water and
Plebeiogryllus plebejus (Saussure), 15-40 d. The hatching nymphs feed are therefore more prevalent in
Teleogryllus testaceous (Walker), on the roots, damaging the crop in upland fields.
Teleogryllus occipitalis (Serville), patches. The nymphs have limited
Loxoblemmus haani Saussure, and migrating ability and generally suffer
Velarifictorus aspersus membranaceus heavy mortality. The nymphal period Damage
(Drury) in Africa. lasts 3-4 mo. The insect has only one
generation per year in temperate Both mole crickets and field crickets
areas. live in ramifying underground
Life history After ricefields have been flooded, burrows, generally 8-10 cm below the
mole crickets are usually seen swim- surface. One mole cricket can burrow
Mole crickets are brownish and very ming on the water, particularly when a distance of several meters in one
plump insects, usually 2.5-3 mm long, the flooded fields are being plowed night. Under upland conditions the
with short antennae. Their front legs and puddled. In areas where mole burrows are more abundant in moist
are classical examples of modification crickets are eaten, people may be seen patches, but in lowland fields they
for digging. The foretibia are modi- collecting them while a field is being are common near the levees and in
fied into spade-like structures resem- plowed. Cannibalism is apparently areas that are not submerged. These
bling the feet of a mole, the character the most important factor regulating burrows run close to the soil surface
Life history
The beetle is reddish, blackish red, or
and the crickets feed on the roots of (Ashmead), Motes subtessellatus dark tan, about 2 cm long and 1 cm
the plants or even the base of the (Smith), and Motes loboriosus wide (Fig. 22d). Antennae have 10
seedling, resulting either in complete (Smith)—parasitize field crickets. segments. Female beetles are usually
severing of the aerial parts from the A nematode, Mermis nigrescens bigger than males. Females prefer to
root or heavy root pruning. Mole (Dujardin), parasitizes adults and lay eggs in moist sandy soil, to a
crickets cannot kill older plants with nymphs of mole crickets. depth of 7-12 cm. A single female lays
large root systems. In China, the fungus Beauveria a maximum of 14 eggs/d and about
Some species of field crickets bassiana (Balsamo) Vuillemin parasit- 50-60 eggs throughout its adult life.
defoliate rice plants by removing the izes nymphs and adults of Gryllotalpa The eggs are laid singly and are
central portion of leaves. africana. The earwig Labidura sp. covered with a jelly-like white sub-
preys on both nymphs and adults of stance. The freshly laid egg is smooth,
the pest. milky white, oval, and is about
Control methods 3.7 mm long and 2.2 mm wide. As the
Chemical control egg grows, a part of it becomes
Cultural control Poisoned bait made from moistened rice hyaline white and other parts remain
Maintaining standing water in the bran and liquid or powdered insecticide milky white. The developing embryo
field prevents mole crickets and field can be placed in the field or on rice becomes visible about 7 d after
crickets from tunneling into the soil bunds to kill night-foraging mole oviposition. Before hatching, the egg
and damaging the crop. crickets and field crickets. In deepwa- is smooth, elliptical, and is about
ter areas, the main control measure is 5.3 mm long and 4.05 mm wide. The
Biological control chemical treatment of soil or seeds at incubation period is 8-10 d.
Mole crickets are cannibalistic, thus planting. Immediately after hatching, the
regulating their own numbers. grub eats the egg shell then remains
Pheropsophus jessoensis Morawitz inactive for 2-3 h. It is milky white
(Carabidae) larvae prey on the eggs with a dirty-white head, and is about
of Gryllotalpa africana. Sphecid wasps
Larra carbonaria (Smith), Larva White grubs 9.8 mm long and 2.3 mm wide at the
head. After 3-4 h, the body expands,
luzonensis (Rohwer) and Larva the head becomes brownish, and the
sanguinea Williams parasitize The term white grub refers to the large grub starts feeding on organic matter.
nymphs and adults by paralyzing larva of a scarab beetle. White grub At this stage, the grub requires a
them and dragging them into wasp species are known either as the higher quantity of organic matter
nests as food for the young. chafers, in which only the larvae feed than of roots.
Other sphecid species — Liris on roots, or the black beetles, in The second-instar grub has mor-
aurulenta (Fabricius), Motes manilae which only the adults feed on roots. phological characters and color
White grubs are widely distributed pattern similar to those of the first
and are considered important pests
Soil-inhabiting pests 75
instar, but the last abdominal seg- the roots cause the plants to become America, and the Caribbean. The pest
ment is enlarged. It is 36.5 mm long stunted and then wilt. Damage to rice is also known as the grass root aphid
and 3.85 mm wide at the head. The crops is higher during drought because it infests the roots of other
second instar is more curved, active, conditions because plants are less graminaceous plants aside from rice.
and cannibalistic. At the first half of able to produce new roots. Damage is These other hosts include species of
the second instar, the food require- in patches since white grubs are Agropyon, Cenchrus, Chloris, Cynodon,
ment is the same as that of the first unevenly distributed in the soil Echinochloa, Eleusine, Eragrostis,
instar, but at the later part, the grub because of their strict moisture Panicum, Saccharum, and Setaria. The
requires a higher quantity of roots requirement. pest has been recorded under differ-
than of organic matter. The second- ent names in several countries. In the
instar period lasts 14-62 d, with an Philippines and Taiwan, China, it
average of 37 d. Control methods was previously known as Dryopeia
The third-instar grub is creamy hirsuta Baker.
white and is about 46.5 mm long and Cultural control Other root aphids known to infest
6.60 mm wide at the head. This Cultural control methods that help rice plants are Rhopalosiphum
period lasts for 143 d (Fig. 22e). reduce field population of the pest rufiabdominalis (Sasaki), Anoecia
Pupation takes place in earthen include delaying land preparation fulviabdominalis (Sasaki), Paracletus
cells in the soil 20-40 cm deep. The until most chafer adults pass their cimiciformis Heyden, and Geoica
freshly formed pupa is creamy white egg-laying phase or die. Weeding lucifuga (Zehntner).
and later turns brown. The pupae are also reduces egg laying by females
exarate and the pupal period lasts since they are attracted to thick
20-24 d. When adults emerge, they vegetation. Life history
remain in the soil until the onset of
the monsoon. Biological control The rice root aphid Tetraneura
The life cycle is completed within Several specialized scoliid wasps nigriabdominalis is small, greenish or
1 yr in the tropics. In temperate such as Campsomeris marginella brownish white, plump, and oval-
regions the life cycle is 2 yr. modesta (Smith) can parasitize white bodied (Fig. 22f). It is not so globose
grubs in the soil. Mermithid nema- compared with most Tetraneura
todes such as Psammomermis sp. also species. Rice root aphids usually
Seasonal occurrence parasitize the larvae. cluster on roots of rice plants. There
and abundance Chemical control
are two adult forms: winged and
nonwinged. Winged adults are
In India, the beetles emerge from the The only practical insecticidal control 1.5-2.3 mm long and nonwinged
soil after about 35 mm of rainfall in measure against white grubs is the forms are 1.5-2.5 mm long. The rice
the monsoon or premonsoon season application of granular insecticides in root aphids reproduce by viviparous
in May. Females lay eggs in June, crop furrows or hills at sowing. parthenogenesis and no male aphids
July, and August. Pupation takes have been recorded. There are four
place around February, and adults nymphal instars, each lasting an
that emerge remain under the soil average period of 2.2, 2.7, 3.8, and
until the onset of the monsoon.
Soil moisture is an important Rice root 5.7 d. The adult aphids live 15-20 d,
and each female produces
factor in egg incubation and survival
of white grubs. The most suitable soil aphids 35-45 nymphs in its lifetime. The first
nymphs are usually born shortly after
moisture is 10-15% for the hatching of the aphids have reached the adult
eggs and 15-25% for survival. The rice root aphid Tetraneura stage.
(=Tetraneurella) nigriabdominalis
(Sasaki) (Homoptera: Aphididae) is
Damage considered a major pest of upland Seasonal occurrence
rice. It occurs in Australia, Bangla- and abundance
Adults of black beetles burrow in the desh, Fiji, India, Indonesia, Japan,
soil and feed on the roots; larvae feed Republic of Korea, Malaysia, Nepal, In the Philippines, the occurrence of
only on organic matter. New Zealand, Pakistan, Philippines, both apterous and alate adults has
Chafer adults, on the other hand, Sri Lanka, Tonga, and the USA. It has been recorded. Alate forms make up
are foliage feeders. Larvae feeding on also been recorded in Africa, Central 2-3% of the population from June to
Soil-inhabiting pests 77
Selected references Litsinger J A, Barrion A T, Soekarna D Rohilla H R, Singh H V, Yadava T P
(1987) Upland rice insect pests: their (1981) White grubs- increasing menace
Akino K, Sasaki M, Okamoto D (1956) ecology, importance and control. IRRI in Haryana. Haryana Agric. Univ. J.
Studies on mole crickets attacking rice Res. Pap. Ser. 123. International Rice Res. 11:256-258.
seeds directly sown without trans- Research Institute, P.O. Box 933, Sachan J N, Gangwar S K, Katiyar J N
planting between rows of wheat and Manila, Philippines. (1980) Field crickets damaging crops in
barley. Bull. Chugoku Agric. Exp. Stn.
Martins J F D S, Ferreira E, Prabhu A S, north eastern hill region. Indian J.
3:91-110.
Zimmermann F J P (1980) Use of Entomol. 42:526-527.
Alam M S, John V T, Kaung Z (1985) pesticides for the control of the main Sithole S Z (1986) Mole cricket (Gryllotalpa
Insect pests and diseases of rice in subterranean upland rice pests. Pesqui. africana). Zimbabwe Agric. J. 83:21-22.
Africa. Pages 67-82 in Rice improve- Agropecu. Bras. 15:53-62.
ment in Eastern, Central and Southern Tan J P (1924) The rice root aphid,
Matsumura H, Ishisaki H, Oda H (1982) Dryopeia hirsuta Baker. Philipp. Agric.
Africa. Proceedings of the Interna-
Appropriate timing of insecticide 13:277-288.
tional Rice Workshop, 9-19 April 1984,
application for controlling the rice
Lusaka, Zambia, Tanaka T (1961) The rice root aphids:
plant weevil, Echinocnemus squameus
Blackman R L, Eastop V F (1984) Aphids Billberg (Coleoptera: Curculionidae). their ecology and control. (Jpn.)
on the world's crops: an identification Special Bull. Coll. Agric. Utsunomiya
Jpn. J. Appl. Entomol. Zool. 26:74-75.
and information guide. John Wiley Univ. 10:1-83.
Mrig K K, Kushwaha K S (1989) Oviposi-
and Sons. 466 p. Veeresh G K (1981) Taxonomy, bionomics
tional behaviour and larval character
Chakrabarti N K, Kulshreshtha P J, Rao of rice root weevil Hydronomodius and control of white grubs. Pages 1-79
molitor Faust. Indian J. Entomol. in Progress in white grubs research in
Y S (1971) Pests and diseases of new
51:481-483. India. G.K. Veeresh, ed. U.A.S. Dep.
varieties and remedial measures.
Entomol. Agric. Coll., Hebbal, Banga-
Indian Farming 21:53-58. Nguyen Van H, Nguyen Ngoc P, Nguyen lore. 154 p.
Ding H S (1985) A survey on the occur- Trung T (1986) Insect and vertebrate
Yano K, Miyake T (1983) The biology and
rence of root aphids and their damage pests of deep water rice in the Mekong
Delta, Vietnam. Int. Rice Res. Newsl. economic importance of rice aphids
to upland rice. Insect Knowledge
11(5):36. (Hemiptera:Aphididae): a review. Bull.
(Kunchong Zhishi) 22:255.
Entomol. Res. 73:539-566.
Eastop V F, HilleRisLambers D (1976) Patel H K, Patel R C, Patel V C (1965)
Survey of the world's aphids. Dr. D.W. Hydronomodius molitor Fst., a new pest
Junk b.v., Publishers, The Hague. of paddy in Gujarat. Indian J. Entomol.
27:496-497.
573 p.
Patel V S, Desai N D (1986) Insecticidal
Feakin S D (1970) Pest control in rice.
control of grubs of rice root weevil
PANS Manual 3. Centre for Overseas
Echinocnemus oryzae Marshall in
Pest Research, London. 270 p.
Gujarat. Pesticides 20:15-16.
Habu A (1986) Redescription of larvae of
Patil B R, Hasabe B M (1981) Studies in
Peropsophus jessoensis (Coleoptera,
the bionomics of white grub Holotrichia
Carabidae). Entomol. Rev. Jpn. 41:113-
serrata F. Pages 115-119 in Progress in
122.
white grubs research in India. G.K.
Hahn H E (1958) Investigations on the Veeresh, ed. U.A.S. Dep. Entomol.
habits and development of Gryllotalpa Agric. Coll., Hebbal, Bangalore. 154 p.
africana in Braudenberg. Beitr. Ento-
Prakasa Rao P S (1989) Bioecology and
mol. 8:334-365.
management of insect pests of rainfed
Hsieh C Y (1970) The aphids attacking upland rice ecosystem. Nutr. Ecol. Inst.
rice plants in Taiwan. II. Studies on the Environ. pp. 93-106.
biology of the red rice root aphid,
Raodeo A K, Deshpande S V (1987) White
Rophalosiphum rufiabdominalis (Sasaki)
grubs and their management. Marath-
(Aphididae, Homoptera). Plant Prot.
wada Agric. Univ. Res. Bull. 11:l-72.
Bull. (Taiwan) 12:68-78.
Reissig W H, Heinrichs E A, Litsinger J A,
Hu M (1985) An investigation on the
Moody K, Fiedler L, Mew T W, Barrion
control of Gyllotalpa africana by
A T (1986) Illustrated guide to inte-
Beauveria bassiana and Labidura sp.
grated pest management in rice in
Natural Enemies of Insects (Kunchong
tropical Asia. International Rice
Tiandi) 7:110-112
Research Institute, P.O. Box 933,
Manila, Philippines. 411 p.
The rice stem maggot Chlorops oryzae Seasonal occurrence Larvae of the first generation feed on
rice seedlings and pupate from late
Matsumura (Diptera: Chloropidae) is
an important pest of rice in several and abundance May to early June. Adults emerge
parts of Asia. The insect is also from mid- to late June, and most of
known to infest wheat, barley, rye, In Japan, the pest occurs in five them migrate to rice areas above
and about 12 species of graminaceous distinct broods: two in the north and 800 m. Larvae of the second genera-
weeds. It is not known to infest three in the south. In two-generation tion damage rice crops and pupate
nongraminaceous plants. Outbreaks areas, first-brood flies appear in June from early to mid-September. Adults
of large populations occurred in and the second-brood flies in Septem- emerge from mid-September to mid-
Japan from 1935 to 1942 and from ber. The first-brood flies oviposit on October and migrate to lower,
1945 to 1955. The fly has been consid- rice seedlings in the seedbed; those of warmer areas where oviposition
ered an important pest of rice in the second brood oviposit on various occurs. The hatching peak of third-
China since the 1970s. grasses on which the insects overwin- generation larvae is in mid-Novem-
ter as full-grown larvae. In three- ber and these larvae overwinter.
generation areas, the first-brood flies
Life history emerge at the end of May and sec-
ond-brood flies the second half of Damage
The adult insects look like small July. Both broods oviposit on the rice
houseflies. Adult life-span averages plants. The third-brood flies appear Damage is caused by the maggots
2 wk, during which the female lays in September, oviposit on grasses and boring near the growing points and
50-100 eggs. The eggs are minute, other hosts, and overwinter as full- feeding on the leaf blades. Usually,
shiny white, elongate, and are laid grown maggots. The flies from two- they feed from the margin toward the
singly on the leaf blades. The average and three-generation areas behave as midrib, causing broad, chewed
incubation period is 7 d. distinct races, but those in mixed patches along the sides of the leaf
The freshly hatched maggots are population areas freely interbreed. blades. Occasionally, they also
whitish and translucent. Each meas- The maggots of three-brood popula- puncture the leaves. Heavy infesta-
ures about 1 mm long. Maggots tions develop faster than those of the tion during the vegetative stage
migrate to the central whorl of the two-brood populations. causes stunting. Second- and third-
rice plants to feed on the leaves near In the Republic of Korea, the pest brood maggots also feed on develop-
the growing points. The larvae also also has three generations a year with ing grains within the boots, causing
infest rice panicles within the boot population peaks in the second half substantial losses. The damaged
and feed on the developing grains. of May, early July, and mid-Septem- grains do not develop and the pan-
The larval stage lasts for about 6 wk. ber. icles have blank spots.
The final instar larvae then move to In China, the pest also has three Rice varieties vary in their suscep-
the upper part of the rice stem and generations a year. Larvae overwinter tibility to the stem maggot. Fre-
pupate on the upper and inner parts in areas below 700 m, mainly in the quently, 10-20% of the panicles in
of the leaf sheath. The pupal stage grasses Alopecurus japonicus, susceptible varieties are infested
lasts for about 14 d. A. equalis, and Leersia hexandra, and in during booting. A 10% infestation
wheat. Overwintered larvae pupate results in about 4% yield loss. Early-
during mid- to late March, and adults heading varieties usually have
emerge from mid- to late April. panicles damaged at the bottom; later
varieties have panicles damaged at
the top.
79
Control methods Selected references
Heinrichs E A, Medrano F G, Rapusas
Cultural control
H R (1985) Genetic evaluation for
Effective control measures include insect resistance in rice. International
the use of irrigation against the Rice Research Institute, P.O. Box 933,
overwintering generation and de- Manila, Philippines. 356 p.
layed sowing of the crop to escape Hirao J (1963) Further observations on the
peak damage by the pest. ecotype formation in the rice stem
maggot. Jpn. J. Appl. Entomol. Zool.
Biological control 7:338-342.
No natural enemies of this pest have Hwang C Y, Lee Y B, Kim S H, Lee M H,
been recorded so far, but spiders are Choi X M (1985) Development of rice
considered probable predators in the stem maggot (Chlorops oryzae) in the
Republic of Korea. fields in Suweon. Korean J. Plant Prot.
24:61-64.
Varietal resistance Iwata T (1963) Ecological studies on the
Several rice varieties from Japan are local variation in the seasonal history
sources of resistance to the rice stem of the rice stem maggot, Chlorops oryzae
maggot. Among them are Aichi- Matsumura [in Japanese, English
asahi, Ashai, Joshu, Kanan-nansen, summary]. Bull. Hokuriku Agric. Exp.
Ou 187, Ou 188, Ou 230, Ou 231, Stn. 4:109-189.
Sakaikaneko, Sin 4, Sin 5, Tohoku 64, Kim K W (1982) Relationships between
Tugaru-asahi, Tyosen, Tyusin 203, damaged site of ears and heading time
Norin 10, and Obanazawa. and position of punctured leaves by
the rice stem maggot, Chlorops oryzae
Matsumura, in the second generation.
Chemical control
Korean J. Plant Prot. 21 :175-178.
Rice stem maggot populations can be
Kim K W, Hyun J S (1982) Studies on the
effectively controlled by insecticide
life history and damage of rice plants
treatment. by the rice stem maggot, Chlorops
oryzae Matsumura [in Korean, English
summary]. Seoul Natl. Univ. Coll.
Agric. Bull. 7(1): 43-55.
Okamoto D (1961) Damages caused by
the rice stem maggot, Chlorops oryzae
Matsumura and its control method [in
Japanese]. Agric. Hort. 36:67-70.
Yan W X, Jiang C H, Xiang D (1981)
Discussion on the occurrence and
forecasting of Chlorops oryzae
Matsumura. Insect Knowledge
(Kunchong Zhishi) 18:197-199.
Yang Z H (1984) A preliminary study on
the bionomics of Chlorops oryzae
Matsumura. Insect Knowledge
(Kunchong Zhishi) 21:l0-15.
The high-yielding, early-maturing, of infrastructure, trained personnel, effective against several others.
modern rice varieties (MVs) devel- and an active program to demon- Earlier concerns that resistance is
oped since the 1960s caused major strate IPM to farmers and extension frequently associated with poor grain
shifts in the insect pest complex. In- workers. Modification and adapta- quality and low yield have been
secticide use, which began during the tion of insect management programs overcome largely by incorporating
widespread adoption of MVs, has to a particular socioeconomic situ- resistance into high-yielding varieties
become a major part of crop manage- ation are equally important. of good quality. Depending on
ment recommendations in many The various components of IPM resistance levels, resistant varieties
developing countries. Indiscriminate are host plant resistance, cultural can be used either as the principal
use of these chemicals caused major control, biological control, and, method of control or as a supplement
outbreaks of insect pests such as finally, chemical control when pest to other methods of insect pest
brown planthopper (BPH) and green damage exceeds or threatens to management. For example, rice
leafhopper (GLH) because of the exceed the economic injury level. varieties resistant to BPH and gall
destruction of indigenous predators midge generally do not need supple-
and parasites that had kept pest mentary control measures to protect
populations in check most of the Components of them against these insects, but those
time. In addition, many MVs have integrated pest resistant to stem borer (SB) need
additional protection from infestation
resistance to insects, particularly to
leafhoppers, planthoppers, and gall management beyond the booting stage, or even
midge. earlier if the borer population is large.
During the past three decades, Host plant resistance Varietal resistance to insect vectors
considerable progress has been made Plant resistance as an approach to of plant diseases also often limits the
on various methods of pest control in pest management in rice confers spread of the viral diseases they
the tropics and subtropics. It is many advantages. Resistant rice transmit. In Colombia, fields of IR8,
important to use all of these measures varieties provide an inherent control which is resistant to the rice
in developing an integrated pest that involves no expense nor environ- delphacid Tagosodes orizicolus but
management (IPM) program that is mental pollution problems, and are susceptible to its transmitted hoja
long-lasting, inexpensive, and envi- generally compatible with other blanca virus, remains virtually virus-
ronmentally safe. In most cases, this insect control methods. The cultiva- free, while other insect-susceptible
means using nonpesticidal methods tion of resistant rice plants is not rice varieties are heavily infested.
of pest control, and resorting to subject to the vagaries of weather as Fortunately, the world’s rice
pesticides only when the pest causes are chemical and biological control germplasm is well endowed with
economic loss. IPM will require measures, and in certain circum- sources of insect and disease resis-
farmers to be trained to identify pests stances it is the only effective means tance. Identification of sources of
and their biological control agents, to of control. Resistant varieties control stable resistance is a major objective
do pest surveillance in their fields, even a low pest density whereas of rice improvement programs
and to apply various control meas- insecticide use is justifiable only throughout the world. Rice varieties
ures. when the density reaches the eco- with multiple resistance to insect
A major constraint to implement- nomic injury level. In some cases, pests and diseases are now grown on
ing an effective IPM program for rice resistance developed in plants for one more than 20 million ha in Asia and
in tropical Asia and Africa is the lack insect pest species may also be Central and South America.
81
Sources of resistance. Screening meth- Southeast Asia. Strong breeding in hybridization programs. On the
ods for more than 30 rice insect pests programs for gall midge resistance other hand, the levels of resistance to
throughout the world have been have been established in India, Sri many insect pests in cultivated rice
developed and sources of resistance Lanka, and Thailand, and several are too low for breeding purposes.
to most were identified over the past resistant rice varieties are available to For most insect pests, only a small
two decades (Table 10). Major em- farmers. Many rice varieties with portion of the world’s germplasm
phasis has been on leafhopper, moderate levels of resistance to SB collection has been screened. Screen-
planthopper, SB, and gall midge. All have been released in Asia. ing of the entire germplasm collection
the major rice-producing countries in Plant resistance for occasional and may identify varieties with higher
South and Southeast Asia have secondary pests offers advantages, levels of resistance.
breeding programs for BPH and GLH which are often overlooked. Resis- Several wild rices have high levels
resistance. BPH-resistant varieties tance sources for leaffolders, rice of resistance to insect pests and
alone occupy about 25% of the hispa, rice bugs, and whorl maggots diseases. With the development of
irrigated lowland rice area in are available, but are not widely used innovative breeding techniques, those
useful genes could be used to
broaden the primary gene pool of
Table 10. Status of screening and breeding for varietal resistance to major insect pests of cultivated rice. Genes for resistance to
rice.
grassy stunt virus have been success-
Status of resistance a fully transferred to rice from Oryza
nivara, and resistance to BPH from
Insect Resistance Resistant Resistant Genes for
Screening
methods sources breeding varieties resistance
Biotypes O. officinalis. The world collection of
encoun-
developed identified lines released identified tered more than 2,200 wild rice accessions
available at IRRI is being evaluated for resis-
+ + - - - tance to various major and minor rice
Scirpophaga incertulas +
(Walker) pests. Several accessions with high
Chilo suppressalis + + + - - - levels of resistance have been identi-
(Walker)
+ - - - - fied and may be used in wide hy-
Maliarpha separatella +
Ragonot bridization breeding programs.
Diopsis macrophthalma + + - - - -
Dalman
+ + + + +
Breeding for resistance. Breeding for
Orseolia oryzae +
(Wood-Mason) insect resistance in rice has been a
Nilaparvata lugens (Stål) + + + + + + focus of rice research programs only
Sogatella furcifera + + + + + ?
during the last two decades. Al-
(Horvath)
Laodelphax striatellus + + + + - ? though differences in varietal suscep-
(Fallen) tibility to pest infestations had been
Tagosodes orizicolus + + + + - ?
recorded for the last 50 yr or more,
(Muir)
Nephotettix virescens + + + + + + until 1962 scientists did not believe
(Distant) there was sufficient insect resistance
Nephotettix cincticeps + + + + - ?
in rice to be of practical value in
(Uhler)
Nephotettix nigropictus + + - - - - breeding programs.
(Stål) In the early screening at IRRI for
Recilia dorsalis + + + + + - genetic resistance to the striped stem
(Motschulsky)
Cofana spectra (Distant) + + - - - - borer (SSB), 10,000 rice accessions
Hydrellia philippina Ferino+ + - - - - from IRRI’s rice collection were
Lissorhoptrus oryzophilus+ + - - - -
evaluated in the field and in green-
Kuschel
Dicladispa armigera + + - - - - house experiments. Several types of
(Olivier) germplasm were identified as resis-
Spodoptera mauritia + - - - - -
tant. Most of those resistant to SSB
mauritia Boisduval
Mythimna separata + - - - - - were susceptible to other pests and
(Walker) diseases and had limited practical
Nymphula depunctalis + + - - - -
value in a breeding program. Fortu-
(Guenée)
Leptocorisa spp. + + - - - - nately TKM6, a cultivar from India,
Oebalus pugnax (Fabricius) + + - - - - was not only resistant to SSB, but was
a + = yes, - = less damaged by other common pests
no, ? = biotypes suspected.
Chemical control
Although it is well known that
chemical control is incompatible with
the ecological approach to pest
management, pesticides still remain
the first line of defense against many
rice insect pests and are used freely
when insect outbreaks occur. In
Japan, national rice production
progressively increased following the
wide adoption of parathion in 1953
and the subsequent use of other
insecticides.
Although insecticides have effec-
tively controlled many insect pest
species, their extensive use has led to
serious social and environmental
repercussions. The poisoning of
livestock, fish, wildlife, and other
beneficial organisms has been linked
with pesticide use. Likewise, there
has been a disturbing increase in
human poisonings, particularly in
developing countries. A recent
survey in a major rice-growing area
of the Philippines showed that, after
widespread adoption of insecticides
by farmers on small holdings, there
was a 27% increase in mortality from
causes other than trauma.
Pest resurgence associated with
insecticidal destruction of natural
enemies and the development of